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The PTERIDOPHYTES of

MEXICO Part I (Descriptions and Maps)

John T. Mickel and Alan R. Smith

The PTERIDOPHYTES

of

MEXICO

Memoirs of The New York Botanical Garden ADVISORY BOARD Patricia K. Holmgren, Director Emerita of the William and Lynda Steere Herbarium The New York Botanical Garden James L. Luteyn, Mary Flager Cary Curator of Botany The New York Botanical Garden Scott A. Mori, Nathaniel Lord Britton Curator of Botany The New York Botanical Garden

EDITORIAL BOARD William R. Buck, Editor The New York Botanical Garden Bronx, New York 10458-5126 USA email: [email protected] Thomas F. Daniel Department of Botany California Academy of Sciences Golden Gate Park San Francisco, California 94118 USA

The Memoirs are published at irregular intervals in volumes of various sizes and are designed to include results of original botanical research by members of The New York Botanical Garden’s staff, or by botanists who have collaborated in one or more of The New York Botanical Garden’s research programs. Ordinarily only manuscripts of 100 or more typewritten pages will be considered for publication. Manuscripts should be submitted to the Editor. For further information regarding editorial policy and instructions for the preparation of manuscripts, contact the Editor. Orders for published and forthcoming issues and volumes should be placed with: The New York Botanical Garden Press 200th St. & Kazimiroff Blvd. Bronx, New York 10458-5126 USA (718) 817-8721; fax (718) 817-8842 email: [email protected] http://www.nybg.org/bsci/spub

The PTERIDOPHYTES of

MEXICO Part I (Descriptions and Maps)

John T. Mickel and Alan R. Smith Memoirs of The New York Botanical Garden Volume 88

The New York Botanical Garden 2004

䉷 Copyright 2004 by The New York Botanical Garden Part I and Part II were originally issued as a single volume on June 30, 2004. This version is printed on demand by The New York Botanical Garden Press. All rights reserved. No portion of this book may be reproduced in any form or by any means, including electronic storage and retrieval systems, except by explicit, prior written permission of the publisher except for brief passages excerpted for review or critical purposes. Composition by Binghamton Valley Composition International Standard Serial Number 0077-8931 The paper used in this publication meets the requirements of ANSI/NISO Z39.48–1992 (R1997).

TM

Library of Congress Cataloging-in-Publication Data Mickel, John. The pteridophytes of Mexico / John T. Mickel and Alan R. Smith. p. cm. — (Memoirs of the New York Botanical Garden ; v. 88) Includes bibliographical references (p. ). ISBN 10 (part I): 0-89327-488-7 (pbk.) ISBN 13 (part I): 978-0-89327-488-7 (pbk.) ISBN 10 (original hardcover): 0-89327-458-5 1. Pteridophyta—Mexico—Classification. 2. Pteridophyta— Mexico—Identification. I. Smith, Alan Reid, 1943– II. Title. III. Series. QK1.N525 no. 88a [QK525.85] 580 s—dc22 [587/.09

The New York Botanical Garden Press 200th St. & Kazimiroff Blvd. Bronx, NY 10458-5126, USA [email protected] http://www.nybg.org/bsci/spub

When Rogers McVaugh invited the senior author to assist him on a three-month collecting expedition in western Mexico in 1957, this new graduate student was introduced to the ferns of Mexico and conceived the project of eventually treating the entire fern flora of the republic. Since this initial visit, McVaugh has been a steadfast supporter of the project, especially in connection with the treatment for Nueva Galicia (Mickel, 1992). In addition to being quite knowledgeable about ferns himself, he has been tireless in the collection and identification of Mexican pteridophytes. On that same 1957 trip, the senior author and McVaugh became acquainted with Jerzy Rzedowski, whose interest and assistance in our collecting and herbarium studies, as well as his abundant fern collections, have been invaluable in the completion of this flora. It is with great appreciation and warm affection that we dedicate this book to these two giants of Mexican botany and floristics.

CONTENTS Part I xi xiii 1 1 4 6

Contributors Acknowledgments Abstract Introduction Conspectus of the Pteridophyte Genera in Mexico Key to the Genera of Mexican Pteridophytes

17 18 19 20 21 45 47 48 60 61 63 64 71 72 132 135 139

Acrostichum Actinostachys Adenoderris Adiantopsis Adiantum Alsophila Ananthacorus Anemia Anetium Anogramma Arachniodes Argyrochosma Aspidotis Asplenium Astrolepis Athyrium Azolla

141 153 158 161

Blechnum Bolbitis Bommeria Botrychium

165 172 174 176 213 215 216 217 219 233 234 239 240

Campyloneurum Ceradenia Ceratopteris Cheilanthes Cheiloplecton Cibotium Cnemidaria Cochlidium Ctenitis Culcita Cyathea Cyclopeltis Cystopteris

242 244 249 250 251 251 268 269 270

Danaea Dennstaedtia Dicksonia Dicranopteris Didymochlaena Diplazium Diplopterygium Doryopteris Dryopteris

379 380 381 386 389 395 396 400 402

Macrothelypteris Marattia Marsilea Megalastrum Melpomene Metaxya Microgramma Micropolypodium Mildella

282 315 317 320

Elaphoglossum Enterosora Equisetum Eriosorus

403 410 410 411

Nephrolepis Neurodium Niphidium Notholaena

321

Gleichenella

322 323 324 325 327 328 329 337 348

Hecistopteris Hemidictyum Hemionanthes Hemionitis Histiopteris Holodictyum Huperzia Hymenophyllum Hypolepis

425 427 427 428 429 433

Odontosoria Oleandra Olfersia Onocleopsis Ophioglossum Osmunda

354

Isoe¨tes

357

Jamesonia

358 360 364 368 368 370 371 372 373 374 375 377

Lastreopsis Lellingeria Lindsaea Llavea Lomariopsis Lonchitis Lophosoria Loxogramme Loxoscaphe Lycopodiella Lycopodium Lygodium

435 443 453 455 460 462 463 466 467 475 477 511 525 528 529 533

Pecluma Pellaea Pentagramma Phanerophlebia Phlebodium Pilularia Pityrogramma Plagiogyria Pleopeltis Polybotrya Polypodium Polystichum Polytaenium Psilotum Pteridium Pteris

544

Radiovittaria

CONTENTS

544 546 547 548 549 550 603 603 604 607

Saccoloma Salvinia Schaffneria Schizaea Scoliosorus Selaginella Sphaeropteris Sphenomeris Sticherus Stigmatopteris

Part II 703

Plates Index to Scientific Names

609 615 621 623 638 648 663 667 668 668 669

Tectaria Terpsichore Thelypteris subg. Amauropelta subg. Cyclosorus subg. Goniopteris subg. Meniscium subg. Stegnogramma subg. Steiropteris subg. Thelypteris Trichomanes

vii

683

Vittaria

686 692

Woodsia Woodwardia

697

Literature Index to Scientific Names

CONTRIBUTORS

The following persons are attributed authorship for all or part of selected generic treatments. R. JAMES HICKEY (Isoe¨tes), Botany Department, Miami University, Oxford, OH 45056 W. CARL TAYLOR (Isoe¨tes), Milwaukee Public Museum, Milwaukee, WI 53233 IVA´N A. VALDESPINO Q. (Selaginella), Apartado 10912, Universidad de Panama´/ANAM, Panama´, Panama MICHAEL D. WINDHAM (Woodsia), Garrett Herbarium, Utah Museum of Natural History, University of Utah, Salt Lake City, UT 84112

ACKNOWLEDGMENTS

We are grateful to the curators and administrators who made their specimens available for study, both on loan and on visitation. Herbaria consulted include ANSM, ARIZ, ASU, B, BM, BR, C, CAS, CHAPA, DS, ENCB, F, FCME, GH, GUADA, IBUG, IEB, ISC, IZTA, K, LAM, LL, MA, MEXU, MICH, MO, MSC, NY, P, PH, PR, PRC, S, SD, SERBO, TEX, UAMIZ, UC, UNL, US, XAL, and YU. Assistance with manuscript preparation and specimen processing was provided by Ruth Russell and Dee Ferguson. Richard Rauh checked many details in the manuscript. Carol Mickel helped with data recording at NY and many other herbaria. We are greatly indebted to the senior author’s research assistant Lee Brown, who processed as well as critically read the full manuscript. We thank Sue Bainbridge for assistance in generating the base maps and Joan Smith for execution of the maps. Special thanks go to William Buck of the New York Botanical Garden for his meticulous reviewing and editing of the manuscript, and to Daniel Tejero-Dı´ez and George Yatskievych for their reviews and helpful comments. Brian Viger of the New York Botanical Garden Press provided valuable assistance in computerization of the manuscript.

Several botanists graciously furnished valuable taxonomic assistance: Judy Garrison Hanks (Lygodium), Richard Hauke (Equisetum), Hugo Navarette (Dennstaedtia), Benjamin Øllgaard (Lycopodium s.l.), W. H. Wagner, Jr. (Botrychium, Ophioglossum), and Michael Windham (Argyrochosma, Cystopteris, Notholaena, Woodsia). George Yatskievych brought several interesting collections to our attention. We also had valuable discussions with Rodolfo Aguirre-Clavera´n, Marı´a de la Luz Arreguı´n-Sa´nchez, Ray Cranfill, Lucrecia Garcı´a A., Francisco Lorea-Herna´ndez, Robbin Moran, Leticia Pacheco, Blanca Pe´rez-Garcı´a, Ramo´n Riba, Daniel Tejero-Dı´ez, and Ernesto Vela´zquez M. Financial support for travel and illustrations came from many members of the New York Chapter of the American Fern Society and other fern friends. Especially generous gifts were received from Irving Knobloch, Eleanor and Arden Moyer, and Gertrude and George Yourke, and bequests from F. Gordon Foster, Elinor Merrell, and Elizabeth and Harold Tovell. Without their assistance, this project could not have been completed.

Ags BCN BCS Camp Cedros Chih Chis Coah Col DF Dgo Gro Gto Guad Hgo Jal Me´x Mich Mor Nay NL Oax Pue QR Qro Rev SLP Sin Son Tab Tam Tlax Tres Marı´as Ver Yuc Zac

Aguascalientes Baja California Norte Baja California Sur Campeche Isla Cedros Chihuahua Chiapas Coahuila Colima Distrito Federal Durango Guerrero Guanajuato Isla Guadalupe Hidalgo Jalisco Me´xico Michoaca´n Morelos Nayarit Nuevo Leo´n Oaxaca Puebla Quintana Roo Quere´taro Islas Revillagigedo San Luis Potosı´ Sinaloa Sonora Tabasco Tamaulipas Tlaxcala Islas Tres Marı´as Veracruz Yucata´n Zacatecas

Mexican States and Territories

Abstract Mickel, J. T. (Institute for Systematic Botany, The New York Botanical Garden, Bronx, NY 10458-5126, USA; email: jmickel@ nybg.org) and A. R. Smith (University Herbarium, University of California, 1001 Valley Life Sciences Bldg. #2465, Berkeley, CA 94720-2465, USA; email: [email protected]). 2004. The Pteridophytes of Mexico. Memoirs of the New York Botanical Garden 88: i–xvi⫹1–1055. The pteridophytes of Mexico include 1008 species and 16 additional varieties or subspecies, a total of 1024 taxa in 124 genera. Keys, descriptions, distribution maps, and cited specimens are given for all species. Nearly all taxa are illustrated with line drawings. This treatment fills the gap between two other recent large floras, those for North America north of Mexico and for Mesoamerica, and spans the most diverse and latitudinally largest pteridophyte flora in the world. It weds the world’s richest dryland pteridoflora of northern Mexico and western United States with the rich wet forests of Mesoamerica, South America, and the West Indies. Forty new taxa are described and ten new combinations are made herein: Anemia jaliscana var. integrifolia Mickel, Asplenium arcanum A. R. Sm., A. dianae A. R. Sm., A. gentryi A. R. Sm., A. sanchezii A. R. Sm., A. sessilifolium var. occidentale A. R. Sm., A. ultimum A. R. Sm., A. venturae A. R. Sm., Botrychium tolucaense W. H. Wagner & Mickel, Cheilanthes apiacea Mickel, C. complanata var. interrupta Mickel, C. decurrens Mickel, C. longipila var. brevipila Mickel, C. maxoniana Mickel, C. spiculata Mickel, C. yatskievychiana Mickel, Ctenitis erinacea A. R. Sm., C. mexicana A. R. Sm., Dryopteris knoblochii A. R. Sm., Elaphoglossum erinaceum var. occidentale Mickel, E. leebrowniae Mickel, E. xanthopodum Mickel, E. yourkeorum Mickel, Lindsaea quadrangularis subsp. mexiae Mickel, Notholaena brachycaulis Mickel, N. brevistipes Mickel, N. meridionalis Mickel, Pecluma alfredii var. occidentalis A. R. Sm., Polypodium arcanum var. septentrionale Mickel, P. polypodioides var. knoblochianum Mickel, P. rzedowskianum Mickel, Selaginella barnebyana Valdespino, S. breedlovei Valdespino, S. hirtifolia Valdespino, S. polyptera Valdespino, S. prolifera Valdespino, Thelypteris loreae A. R. Sm., T. martinezii A. R. Sm., T. venturae A. R. Sm., and Woodsia cystopteroides Windham & Mickel. Ten new combinations are made: Asplenium semipinnatum (Hieron.) A. R. Sm., Astrolepis laevis (M. Martens & Galeotti) Mickel, Cheilanthes aurea var. palmeri (D. C. Eaton) Mickel, C. marsupianthes (Fe´e) Reeves ex Mickel, C. pringlei var. moncloviensis (Baker) Mickel, Pecluma alfredii var. cupreolepis (A. M. Evans) A. R. Sm., Pellaea ternifolia subsp. brandegeei (C. C. Hall) Mickel, Polypodium riograndense (T. Wendt) Mickel, Tectaria trichodes (C. V. Morton) A. R. Sm., and Thelypteris resinifera var. proxima (C. Chr.) A. R. Sm.

Resumen La pteridoflora de Me´xico incluye 1008 especies y 16 variedades or subespecies adicionales, para un total de 1024 taxones en 124 ge´neros. Claves, descripciones, mapas de distribucio´n y una lista de los ejemplares citados se dan para todas las especies. Casi todos los taxones esta´n ilustrados con debujos. Este tratamiento llena el espacio que existe entre las dos grandes floras recientes, la de Ame´rica del Norte norte de Me´xico ya la de Mesoame´rica, y cubre una de las pteridofloras ma´s diversas y ma´s grandes latitudinalmente del mundo. Contiene la pteridoflora ma´s diversa de las regiones secas del norte de Me´xico y de la parte occidental de los Estados Unidos con los muy diversos bosques hu´medos de Mesoame´rica, Ame´rica del Sur y las Antillas. Se describen cuarenta especies nuevas y se hacen diez combinaciones.

Introduction Mexico has one of the most exciting pteridophyte floras in the world. With 1008 species and a broad range of habitats (rain forests, cloud forests, temperate woodlands, various dry habitats), it is extremely rich and diverse. Fern floras have been produced for many parts of the world in relatively recent years. In the Old World there is coverage for Europe, Russia, Japan, China, Thailand, Malaya, Australia, New Zealand, South Africa, and Hawaii. In America there is coverage for Costa Rica-Panama (Lellinger, 1989), Surinam (Kramer, 1978), Peru (Tryon & Stolze, 1989–1994), Venezuela (Smith, 1985), Trinidad (Mickel, 1985), and parts of the West Indies (Proctor, 1977, 1985, 1989). Floras of wider coverage have been produced for North America (FNA Ed. Comm., 1993) and Mesoamerica (Davidse et al., 1995). This Mexican treatment fills the gap between these two great floras. North America north of Mexico has 441 species, Mesoamerica nearly 1400 species and varieties. If Mexico’s 186 endemic species are added, a total of ca. 2000 species, or 20% of the estimated 10,000 pteridophyte species of the world, occurs in this con-

tiguous area. Taken together, this flora includes one of the most diverse and latitudinally most extensive pteridofloras in the world, ranging from the Arctic to the heart of tropical wet forests.

The Republic of Mexico Mexico is dominated physiographically by two major uplifted mountain ranges: the Sierra Madre Oriental on the east and the Sierra Madre Occidental on the west. Where they converge in the south, the range continues as the Sierra Madre del Sur. In the east, mountains extend from Tamaulipas to Oaxaca, with peaks 2700–3400 meters high and with rain or fog nearly every day on the Atlantic slope. Consequently, the flora is extremely rich, with a wealth of tree ferns, filmy ferns, and other groups requiring constant high humidity. Wet montane forests cover the lower eastern slopes facing the Gulf of Mexico and give rise to wet pine-oak forests in a cloud zone above 2200 meters, with fir on the higher ridges at 2600–2800 meters. Inland, the descending slopes give way to drier pineoak and other dry forests before ultimately sloping down to

2

THE PTERIDOPHYTES OF MEXICO

dry interior valleys. The wetter slopes of this range are the richest part of the country for ferns, both in species diversity and abundance. The Sierra Madre Occidental runs from the border with the United States southeast to Oaxaca, with peaks mostly 1900–2500 meters in elevation. The southern part of this range (Michoaca´n to Oaxaca) is wet on the Pacific slope but without the constant rainfall and high humidity of the Atlantic slope. It has a distinctly drier winter season, and consequently there are few tree ferns, filmy ferns, and associated species. The Sierra Madre del Sur tapers down to only 80 meters elevation at the Isthmus of Tehuantepec before rising again in Chiapas as the Sierra Madre de Chiapas. Linking the two major mountain chains is the trans-Mexican volcanic belt, ranging across the country at 19–22⬚ N latitude, from Nayarit and Colima east to Veracruz. This interior mountain range includes the highest peaks of the country, including Orizaba on the east (5610 meters), Popocate´petl (5465 meters), and Iztaccihuatl (5230 meters) outside Mexico City, and the Nevado de Colima (4240 meters) on the west, in Jalisco. Between the two major chains in the north lies a great dry high plateau, mostly 1200–2000 meters elevation, with Mexico City (2238 meters) at its southern extreme. The area is characterized by dry forests and open deserts or sub-deserts with only occasional moist valleys. Other ranges include low dry mountains along the length of the peninsula of Baja California, the ridges mostly under 2000 meters (but to 3069 meters). In contrast, the Yucatan Peninsula is uniformly low, mostly 0–40 meters elevation, rarely reaching 100 meters. A more detailed description of the geography and vegetation of Mexico can be found in Rzedowski (1978).

History of Pteridological Studies in Mexico Fern collecting in Mexico began with Martı´n de Sesse´ and Jose´ Mocin˜o on the Royal Botanical Expedition to New Spain in 1787–1803 (McVaugh, 1977), followed by Christian Schiede in Ⳳ 1825–1831 (Schlechtendal & Chamisso, 1830), but the ferns gathered in these early expeditions were relatively few. The first major fern collector was the Belgian botanist/horticulturist Henri Galeotti, who collected in 1835–1840 (McVaugh, 1978), and published with Martin Martens the first book on Mexican ferns (Me´moir sur les Fouge`res du Mexique, 1842; 182 species). He was soon followed by the Danish botanist Frederik Liebmann, who collected in 1841–1843 and published his Mexicos Bregner in 1849 (247 species; Mickel, 1987). More collectors soon followed, including Friedrich Mu¨ller, Gotthold Hahn, Euge`ne Bourgeau, Alwin Aschenborn, Auguste Ghiesbreght, Jean Linden, and especially Wilhelm (Jose´ Guillermo) Schaffner (see Rzedowski, 1959), whose specimens were important in the pteridophyte treatments for Mexico by Antoine Fe´e (1857c; 523 species) and Euge`ne Fournier (1872; 542 species). Veracruz was the eastern port of entry for the country, so European collectors began there, collecting around Orizaba and eventually traveling to Puebla, Oaxaca, Me´xico, Jalisco, and San Luis Potosı´.

In the latter part of the nineteenth and early twentieth centuries a flurry of American collectors, including Edward Palmer, Charles Parry, Cyrus Pringle, and Carl Purpus, made significant collections in many parts of the country, and Townsend Brandegee collected in Baja California. The first significant collections of ferns made by a Mexican botanist were by Jose´ Rovirosa in Chiapas. Rovirosa (1909) treated 216 species, based mostly on his own collections, from Chiapas and Tabasco. Thirty years later, in 1939, an amateur botanist Cassiano Conzatti published a volume of his Flora Taxono´mica Mexicana: Pterido´fitas o Helechos (609 species). Both the Rovirosa and Conzatti works were useful at the time but lacked taxonomic rigor by modern standards. In the last 50 years there has been a wave of state and regional fern floras and annotated checklists, beginning with the Valley of Mexico (Matuda, 1956), Chihuahua (Knobloch & Correll, 1962), Sonoran Desert (Shreve & Wiggins, 1964), Baja California (Wiggins, 1980), Chiapas (Smith, 1981), Nuevo Leo´n (Aguirre-Clavera´n, 1983), Oaxaca (Mickel & Beitel, 1988), Nueva Galicia (Mickel, 1992), Veracruz (Palacios-Rios, 1992), El Bajı´o (Dı´az-Barriga & Palacios-Rios, 1992), Tabasco (Magan˜a Alejandro, 1992), Morelos (Riba et al., 1996), Guerrero (LoreaHerna´ndez & Vela´zquez M., 1998), Quere´taro (Arreguı´nSa´nchez et al., 2001), Me´xico (Tejero-Dı´ez & Arreguı´n-Sa´nchez, 2004), and the Yucatan Peninsula (Palacios-Rios, 2002a-c). Work has also begun on a Mexican fern flora by Ramo´n Riba and his associates, with the publication of fascicles dealing with a number of primarily small families of ferns and allies (Flora de Me´xico, Vol. 6, nos. 1–5, 1993–1999). This burst in floristic activity has been abetted by a wave of additional collecting, often in previously inaccessible or remote areas. A few of the principal collectors during this time have been Rzedowski, McVaugh, G. B. Hinton, and Dennis Breedlove in the southern and western parts of Mexico, Ira Wiggins, Annetta Carter, and Reid Moran in Baja California, and many botanists from the University of Arizona, Arizona State University, and the University of Texas in Coahuila, Nuevo Leo´n, San Luis Potosı´, and Tamaulipas. Other Mexican collectors who have contributed substantially to the body of collections available for study include Rodolfo Aguirre-Clavera´n, Marı´a de la Luz Arreguı´n-Sa´nchez, Francisco Lorea, Esteba´n Martı´nez S., Ramo´n Riba, Daniel Tejero, A´ngel Ventura A., Francisco Ventura A., Ernesto Vela´zquez, and Se´rgio Zamudio. This is not the final word on the Mexican fern flora, but rather another step in furthering our knowledge of the subject, just as were the works of Galeotti and others in their time. There are many species complexes that need more study, and we have pointed out problems that need resolution. The use of modern techniques of cytology, chemistry, molecular studies, and detailed comparative studies of morphology and spores can be expected to pay great dividends insofar as understanding species limits, relationships, and ranges.

Phytogeographical Connections Mexico’s pteridoflora is not just a combination of species from the western United States and Central America but is an

THE PTERIDOPHYTES OF MEXICO

exciting and taxonomically complex mixture, including several genera that appear to have Mexico as an epicenter for species radiation. In the Chihuahuan and Sonoran deserts of northern Mexico there are ties to the floras of Arizona, New Mexico, and Texas, and Baja California shares a number of species with California. In the south, the rich wet forests of Central America extend into Chiapas, Oaxaca, and Veracruz, as well as parts of Puebla and Hidalgo. Some species with Central American affinities are confined to this region, and others are more widespread and extend along the Andean Cordilleras. Still others range throughout tropical America, including the Antilles. Studies are needed to better assess many wide-ranging species, which in many cases have different names applied to them in different regions. Some of the species shared with Central America are not of the wet forests but occur in western Mexico from Sinaloa to Costa Rica (e.g., Cheilanthes skinneri). Other dryland species extend even further, for example, Pellaea ovata, P. ternifolia, Cheilanthes marginata, and C. bonariensis. A few Antillean species occur in Mexico only in the Yucatan Peninsula (e.g., Anemia cicutaria, Sphenomeris clavata, and Thelypteris augescens), whereas others are found in Oaxaca and Veracruz (e.g., Cheilanthes fimbriata) or more widely (Cheilanthes jamaicensis). Several Mexican species are highly disjunct in the Himalayas (Asplenium exiguum), or Peru (Blechnum pennamarina), or Hawaii (Pellaea ternifolia, Elaphoglossum paleaceum). There are floristic ties to eastern Asia (Huperzia serrata) and even a few pantropical species at high elevations (e.g., Dryopteris wallichiana). Relatively few species range throughout Mexico (e.g., Cheilanthes bonariensis, Adiantum capillus-veneris, and Asplenium monanthes occur in 25–26 states as well as the Distrito Federal), whereas 202 taxa are known from only a single state and 484 taxa, or nearly half the pteridophyte taxa in Mexico, are from only 1–3 states. Seventy-five taxa are known from a single collection. There are 186 species (plus 16 varieties) endemic to Mexico. The genera with the highest percentage of endemics are largely those that occur in seasonally dry habitats (Anemia 45%, Argyrochosma 50%, Cheilanthes 30%, Notholaena 42%, Selaginella 35%, and Woodsia 57%). Some genera that are predominantly of wet forests (Asplenium, Elaphoglossum, Polypodium) have species endemic to dry regions. Baja California and the Yucatan Peninsula, each with 70–75 species of pteridophytes, have only one species, Asplenium pumilum, in common. Exciting and unexpected discoveries are still being made, even in areas thought to be reasonably well collected. In this study we have found Polypodium microgrammoides, from Guanajuato, a simple-bladed species related to the more widespread P. muenchii and only the second member of this group; Polystichum schizophyllum (Guerrero) and Diplazium wendtii (Oaxaca), the most dissected members of their genera in Mexico; and Blechnum penna-marina, a widespread and well known species of Peru, Bolivia, Chile, and other austral regions but disjunct in Chihuahua and Durango, four thousand miles from the nearest known collection. Recently described is Pellaea ribae from San Luis Potosı´, with densely villous fronds in a genus of glabrous

3

or only sparsely hairy species. Much of Mexico is in need of further exploration by collectors with a keen eye for ferns. The richness of ferns in Mexico is such that in some areas it is virtually impossible to be confident that one has found everything, even in areas previously visited by botanists. For example, Galeotti, on a trip from San Pablo Nolasco to the Llano Verde in Oaxaca (a rise of about 300 meters elevation from dry pine-oak forest through subtropical deciduous forests to cloud forest of pine-yew-oak), collected 23 species of pteridophytes. One hundred thirty years later the senior author, traveling the same trail, collected 140 species. No doubt there remain uncollected species of pteridophytes along this trail.

This Treatment In writing this account, we have borrowed heavily from the recent pteridofloristic models of Smith (1981, Chiapas), Stolze (1976, 1981, 1983, Guatemala), Proctor (1985, Jamaica), and Mickel and Beitel (1988, Oaxaca, as well as multi-authored floras for North America (FNA Ed. Comm., 1993) and Mesoamerica (Davidse et al., 1995). Many of the species encountered in Mexico have been treated in some fashion in recent pteridofloras, and we have utilized them as starting points in our studies. Inasmuch as there is widespread disagreement among pteridologists regarding familial and generic phylogenetic arrangements (Alston, 1956; Ching, 1940; Christensen, 1938; Copeland, 1947; Crabbe et al., 1975; Holttum, 1947, 1949; Kubitzki, 1990; Nayar, 1970; Pichi Sermolli, 1973; Tryon & Tryon, 1982), we have chosen to ignore families, although the Mexican genera are summarized in the Conspectus (pp. 4–6). A flora is not the place to espouse a phylogenetic scheme; we believe utility should be the prevailing guide in selecting an arrangement. Consequently, we have arranged the genera and species alphabetically throughout the text. Information retrieval is much quicker in an alphabetic, rather than a phylogenetic scheme, and that to us is the most crucial element in our decision. We have discussed the relationships of genera and families under the genera, and often refer the reader to additional references that expand on classification or phylogenetic information. As a primary division of labor in writing this flora, each of the principal authors (Mickel and Smith) treated about half the genera. However, the entire text was examined by both of us. No distinction has been made between subspecies and varieties; we have followed existing treatments in selecting a rank for infraspecific taxa. New taxa are credited to the author(s) primarily responsible for the particular genus. Keys. Keys were prepared for mature plants, based primarily on Mexican specimens. Plants of some species may be precociously fertile, unusually small, and therefore not readily keyable. Synonymy. Synonymies of genera and species are not intended to be complete but do include widely used names. An attempt has been made to include all names based on Mexican types, although most certainly there remain obscure names of uncertain typification and identity, names for which we have not accounted. Few, if any, of these are likely to displace names used in this flora.

4

THE PTERIDOPHYTES OF MEXICO

Literature citation and author abbreviation. Abbreviation of references follows Lawrence et al. (1968) for journals and Stafleu and Cowan (1976, 1979, 1981, 1983, 1985, 1986, 1988) for books. Author abbreviations follow Pichi Sermolli (1996), which in turn is based primarily on Brummitt and Powell (1992). References of a general nature and works that apply to multiple genera are placed under “Literature” before the figures at the end of this treatment. Literature that applies more specifically is placed with the particular genus. Types. Information on Linnaean types was found in Jackson (1912) and Savage (1945). Whenever possible, we have attempted to see type material of Mexican species; many of the early types reside in European herbaria. Illustrations. Three hundred twenty-eight plates provide illustrations of nearly all the taxa. Most of the figures and their arrangement on the plates were done by Haruto Fukuda. Nearly all the figures in the Pteridophyte Flora of Oaxaca (Mickel & Beitel, 1988) were rearranged and supplemented and are placed here with the individual genera. Some of the figures were previously done by Bobbi Angell and are designated by BA. A few illustrations from Mickel (1992) were drawn by Karin Douthit, identified here by KD; that of Cystopteris membranifolia was prepared by Charles Crane, identified by CCC. Those for Marsilea are from David Johnson (1986), and the Equisetum stem cross sections from Richard Hauke (1963) are used with permission. Mexican material was used whenever it was deemed adequate, but extraterritorial specimens were sometimes used for illustration of uncommon or rare species. Blade details are largely of the abaxial surface but not usually so stated unless to distinguish from adaxial views on the same plate. Scale bars with broken lines give the measurement of unlabelled bars with solid lines. Descriptions. Descriptions are based on Mexican specimens unless the taxa are rare, in which case we used specimens from other parts of the range. Chromosome numbers. We report chromosome numbers, generally in 2n form, where these are known. These may be from Mexican and/or extraterritorial sources. In most cases we have not verified voucher identifications except for our own counts; indeed, vouchers do not exist for many published counts. Distribution. For all species and infraspecific taxa, we characterize habitat, followed by a statement of overall distribution. Abbreviations of Mexican states are found on the cover pages. Although the official abbreviation for Baja California Norte is BC, we here use BCN to avoid confusion with the entire peninsula of Baja California or Baja California Sur (BCS). Information on Pringle’s localities was derived from Davis (1936). Specimen citation. Only specimens actually seen are cited. We have tried to cite about ten specimens for each taxon. If fewer than ten specimens were seen, all are cited. If specimens were seen from ten or more states, generally only one specimen per state is cited. Galeotti’s numbers were applied to the species after they were processed in Brussels and have no chronological or geographical meaning (McVaugh, 1978 ). Occasionally, the same number included specimens from different localities. For determining

types of species based on Galeotti types, the locality, as well as the collection number, must be taken into account. Some Pringle numbers are also the result of repeated visits on different dates to the same or different localities. Liebmann did not use collection numbers at all. Instead, he numbered the fern specimens in two sequences, usually with a number from each series on each sheet. In his “Plantae Mexicanae” series the same number applies to an entire species or at least several sheets of a species, whereas the “Flora Mexicana” series seems to use one number per sheet. With these collections, we try to use both numbers when known, especially in citation of types. Some duplicates of Liebmann collections, especially at GH and US, do not have a number. Although we studied all Liebmann collections at C for their identity (Mickel, 1987), there was not time to record the numbers of all the specimens, and we have indicated them in the specimens cited merely as “Liebmann (C).” Unverified or mistaken reports in the literature are indicated in a separate paragraph and most are not mapped. A few collections not seen by us are very likely correctly determined, in our judgment, and are mapped. Mapping. Maps were prepared for all taxa with one dot per state, utilizing ArcView GIS 3.2. For specimens not seen but reported elsewhere, the state record was mapped if we had strong confidence in the determination. Where a species has been recorded from a given state, but we have seen no specimen, we have cited references.

Conspectus of the Pteridophyte Genera in Mexico For purposes of this flora we apply a fairly conservative and traditional circumscription of taxa at higher levels, for the most part following the informal classification of Crabbe et al. (1975), but with deviations where more recent information suggests that earlier concepts were either erroneous or inadequate. We have not adopted a specific family classification as a framework for this flora because our knowledge of family circumscriptions is in a state of flux, there is relatively little concensus among pteridologists as to the appropriate family name for certain groups, and because the morphological characters that would have to be used in any family key are sometimes difficult to assess or are obscure to all but a handful of professional pteridologists. Nevertheless, we provide the following conspectus of Mexican genera for those wishing to place a particular genus and species in a family hierarchy. Development and understanding of relationships among families and genera, and how this should be reflected in a classification, undergoes nearly continual change in systematics. With the advent and application of molecular and cladistic methods, as well as refinement and further understanding of evidence from traditional sources of morphology, anatomy, cytology, and other sources, we now have considerably more confidence in hypotheses of relationship, and these phylogenetic hypotheses are also more firmly grounded theoretically. Ideally and ultimately, the hope is to implement these phylogenetic findings in more stable and natural classifications, better reflective of evolutionary relationships.

THE PTERIDOPHYTES OF MEXICO

Within the “basal” pteridophyte families, those thought to be of older derivation (and with the oldest fossil records), relatively few changes over and beyond the classification in Crabbe et al. are necessary or advisable. The so-called basal families in our treatment include Lycopodiaceae, Selaginellaceae, and Isoe¨taceae; the eusporangiate fern families Marattiaceae, Ophioglossaceae, Equisetaceae, and Psilotaceae; and the basal leptosporangiate fern families, Osmundaceae to Azollaceae. All these constitute the first 19 families listed below. One of the few deviations from Crabbe et al. (1975) in the earlier evolved families involves the placement of Dicksonia with Culcita and Cibotium, in a family Dicksoniaceae. For the higher leptosporangiate ferns, beginning with Lindsaeaceae, ideas on circumscription have undergone significant and in some cases radical change in the last 25 years. Only the major deviations from Crabbe et al. (1975) are discussed briefly below: Morphological and molecular evidence now suggests that the lindsaeoid ferns, often included in the Dennstaedtiaceae, are a separate lineage, and so those genera can be placed in their own family. The placements of Lonchitis and Saccoloma are still in doubt, and these two genera are truly “incertae sedis,” genera of uncertain placement. Some molecular evidence suggests that Lonchitis is more closely related to the Lindsaeaceae rather than to elements here placed in the Dennstaedtiaceae. The placement of Saccoloma is even more in doubt; it is perhaps more closely related to the tree fern assemblage. The retention of Lonchitis and Saccoloma in the Dennstaedtiaceae below is more a matter of convenience rather than conviction. Our ideas on the classification of the Pteridaceae (formerly often called Adiantaceae) have changed substantially. Vittariaceae and Parkeriaceae, as well as Acrostichum, are all believed to be part of the “pteridoid” clade. Because it is large, the family Pteridaceae is often further subdivided into subfamilies (see, e.g., Tryon, Tryon, & Kramer in Kubitzki, 1990). But even within the subfamilial taxa, our concepts have been altered by recent evidence. For example, Ceratopteris and Acrostichum are now thought to be sister genera, and the cryptogrammoid ferns, with sole representative Llavea in Mexico, are now believed to be one of the basal elements in the entire family. Relationships of some

5

genera, e.g., Jamesonia and Eriosorus, are thought to be closer to the Pteridoideae, (including the taenitoid assemblage, subfam. Taenitoideae), than with the cheilanthoid genera, where they were placed by Crabbe et al. (1975). The family Polypodiaceae has also been the focus of major studies assessing relationships. There is now high confidence that the Grammitidaceae nests within the Polypodiaceae, but its sister group within the polypods, despite intensive sampling, is still uncertain. Because it is easily definable by a host of morphological synapomorphies, and is a large family, we suggest that it be maintained at family rank for the present. New understanding of relationships within the Polypodiaceae causes us to abandon previously used subfamilial ranks Polypodioideae and Pleopeltoideae as defined by Crabbe et al. (1975). Similar arguments affect genera previously regarded as comprising the dryopteroid ferns (in the broad sense). For example, the Dryopteroideae, Tectarioideae, and Lomariopsidoideae can no longer be maintained with their classical circumscriptions; they are polyphyletic and unnatural assemblages of genera. We now suggest the adoption of a family Dryopteridaceae, essentially as defined by Kramer in Kubitzki (1990), but with the addition of the Lomariopsidaceae, Oleandraceae, and Nephrolepidaceae and the segregation of the athyrioid ferns (Athyriaceae). Most genera as treated in this flora appear to be monophyletic at this stage of understanding, with the following significant exceptions: Pteris, Asplenium, Polypodium, genera of members of the Cheilanthoideae (Pteridaceae), in particular, segregates of Cheilanthes (Aspidotis, Mildella), as well as Cheilanthes itself. Further study, and no doubt recircumscription, will be required for these genera before the idealistic goal of monophyly can be attained. In the case of Asplenium, it is highly likely that the segregates Holodictyum, Schaffneria, and Loxoscaphe, as recognized in this flora, are part of an indivisible Asplenium; at least that Asplenium itself will have to be further subdivided in order for the three segregates mentioned to be accepted. Other less major examples deviating from monophyly include Cyathea (Cnemidaria is nested within Cyathea), in the Cyatheaceae, Dicksoniaceae (Cibotium may be misplaced), and Terpsichore (in the Grammitidaceae).

Classification of Pteridophyte Genera in Mexico (mostly following Crabbe et al., 1975) (Numbers following the genera are their sequence numbers in the text of this book. Parenthetical numbers are the number of species and the number of endemics, respectively, in Mexico.) Lycopodiaceae Lycopodium-70 (2:0) Lycopodiella-69 (3:0) Huperzia-55 (15:1) Selaginellaceae Selaginella-113 (80:30) Isoe¨taceae Isoe¨tes-58 (6:2) Equisetaceae Equisetum-46 (3:0)

Psilotaceae Psilotum-104 (2:0) Ophioglossaceae Botrychium-21 (6:2) Ophioglossum-89 (8:0) Marattiaceae Marattia-73 (3:0) Danaea-35 (3:0) Osmundaceae Osmunda-90 (2:0)

Hymenophyllaceae Hymenophyllum-56 (22:0) Trichomanes-121 (26:1) Gleicheniaceae Diplopterygium-41 (1:0) Sticherus-116 (4:0) Dicranopteris-38 (1:0) Gleichenella-48 (1:0) Schizaeaceae Schizaea-111 (2:0)

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Actinostachys-2 (1:0) Lygodium-71 (3:0) Anemia-8 (20:10) Plagiogyriaceae Plagiogyria-98 (1:0) Metaxyaceae Metaxya-77 (1:0) Lophosoriaceae Lophosoria-66 (1:0) Cyatheaceae Cnemidaria-28 (2:1) Cyathea-32 (8:0) Alsophila-6 (3:0) Sphaeropteris-114 (1:0) Dicksoniaceae Dicksonia-37 (1:0) Culcita-31 (1:0) Cibotium-27 (2:1) Marsileaceae Marsilea-74 (7:0) Pilularia-96 (1:0) Salviniaceae Salvinia-109 (2:0) Azollaceae Azolla-17 (2:0) Lindsaeaceae Lindsaea-62 (5:0) Sphenomeris-115 (1:0) Odontosoria-85 (2:0) Dennstaedtiaceae Dennstaedtia-36 (8:0) Hypolepis-57 (10:4) Pteridium-105 (3:0) Histiopteris-53 (1:0) Lonchitis-65 (1:0) Saccoloma-108 (2:0) Pteridaceae Adiantoideae Adiantum-5 (35:5) Vittarioideae Polytaenium-103 (4:0) Scoliosorus-112 (1:0) Anetium-9 (1:0)

THE PTERIDOPHYTES OF MEXICO

Hecistopteris-49 (1:0) Ananthacorus-7 (1:0) Vittaria-122 (4:0) Radiovittaria-107 (1:0) Cryptogrammoideae Llavea-63 (1:0) Pteridoideae (including Taenitoideae) Pteris-106 (18:1) Acrostichum-1 (2:0) Anogramma-10 (3:1) Pityrogramma-97 (4:0) Jamesonia-59 (1:0) Eriosorus-47 (2:0) Ceratopteridoideae Ceratopteris-24 (2:0) Cheilanthoideae Cheilanthes-25 (60:17) Cheiloplecton-26 (1:0) Adiantopsis-4 (2:0) Aspidotis-13 (2:1) Mildella-80 (2:0) Notholaena-84 (24:9) Pentagramma-93 (1:0) Argyrochosma-12 (12:5) Astrolepis-15 (5:0) Pellaea-92 (15:3) Doryopteris-42 (2:0) Hemionitis-52 (5:0) Hemionanthes-51 (1:1) Bommeria-20 (5:3) Aspleniaceae Asplenium-14 (86:19) Loxoscaphe-68 (1:0) Holodictyum-54 (1:1) Schaffneria-110 (1:0) Athyriaceae Onocleopsis-88 (1:0) Athyrium-16 (5:1) Diplazium-40 (25:4) Hemidictyum-50 (1:0) Cystopteris-34 (3:1) Woodsia-123 (8:3) Thelypteridaceae

Thelypteris-120 (69:8) Macrothelypteris-72 (1:0) Blechnaceae Blechnum-18 (15:0) Woodwardia-124 (4:1) Dryopteridaceae Didymochlaena-39 (1:0) Olfersia-87 (1:0) Polybotrya-100 (2:0) Lastreopsis-(2:0) Megalastrum-75 (4:0) Bolbitis-19 (8:0) Elaphoglossum-44 (58:19) Stigmatopteris-117 (2:0) Ctenitis-30 (21:5) Dryopteris-43 (13:3) Arachniodes-11 (1:0) Polystichum-102 (18:1) Adenoderris-3 (1:0) Phanerophlebia-94 (8:1) Cyclopeltis-33 (1:0) Lomariopsis-64 (3:1) Tectaria-118 (7:0) Nephrolepis-81 (8:0) Oleandra-86 (1:0) Polypodiaceae Loxogramme-67 (1:0) Pleopeltis-99 (7:1) Neurodium-82 (1:0) Microgramma-78 (6:0) Niphidium-83 (1:0) Campyloneurum-22 (9:0) Polypodium-101 (55:17) Phlebodium-95 (2:0) Pecluma-91 (10:0) Grammitidaceae Enterosora-45 (3:0) Ceradenia-23 (4:1) Lellingeria-61 (6:1) Melpomene-76 (9:1) Terpsichore-119 (9:0) Micropolypodium-79 (3:0) Cochlidium-29 (3:0)

Key to the Genera of Mexican Pteridophytes 1. Plants with no leaves (non-vascular enations present) or roots; stems dichotomously branching; sporangia trilocular. .................... 104. Psilotum. 1. Plants with leaves, rarely lacking roots; sporangia unilocular. 2. Leaves each with a single unbranched vein. 3. Leaves in whorls of 10–50, forming sheaths at nodes; stems hollow, grooved. ............................................................................ 46. Equisetum. 3. Leaves spirally arranged or rarely in whorls of 4; stems solid, not grooved. 4. Leaves mostly 3–30 cm long, grass-like; plants aquatic or in ephemeral pools. 5. Leaves circinate at young tips, borne on short- to long-creeping, filiform rhizomes; sporangia in globose sporocarps that are borne on rhizomes. .................................................................................................................................................... 96. Pilularia. 5. Leaves not circinate, clustered on short corm-like stems; sporangia embedded in leaf bases, not in separate globose sporocarps. ...................................................................................................................................................................................... 58. Isoe¨tes. 4. Leaves less than 3 cm long, not grass-like; plants terrestrial or epiphytic, not aquatic.

THE PTERIDOPHYTES OF MEXICO

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6. Strobili 4-sided; leaves oblong to ovate; plants heterosporous. ....................................................................................... 113. Selaginella. 6. Strobili round in cross section or sporangia in axils of unmodified leaves; leaves usually linear; plants homosporous. 7. Sporangia borne in axils of unmodified leaves, not in distinct cones, or if with distinct cones (H. myrsinites), then epiphytic; horizontal stems absent; upright shoots clustered; plants epiphytic or terrestrial. ...................... 55. Huperzia. 7. Sporangia borne in axils of modified leaves that form distinct cones; horizontal stems present, with upright shoots distant along rhizomes; plants terrestrial. 8. Strobili borne on leafy peduncles or nodding on lateral branches; peduncles, if present, bearing closely spaced unreduced leaves. ..................................................................................................................................... 69. Lycopodiella. 8. Strobili borne on distinct peduncles; peduncles bearing remote reduced leaves. ........................................ 70. Lycopodium. 2. Leaves complex, veins branched. 9. Plants aquatic, floating on water surface (or stranded on mud in ephemeral pools). 10. Leaves more than 5 cm long, pinnately divided; homosporous. ........................................................................................... 24. Ceratopteris. 10. Leaves less than 2 cm long, round to ovate; heterosporous. 11. Leaves 1–2 mm long, with minute papillae less than 0.1 mm long on upper surfaces. ...................................................... 17. Azolla. 11. Leaves 1–2 cm long, with hairs in groups of 4 with fused bases on upper surfaces. ...................................................... 109. Salvinia. 9. Plants terrestrial, epipetric, or epiphytic, or rooting in mud or sand in shallow water. 12. Leaves clover-like, 4 obovate leaflets at tips of stipes; plants heterosporous, with sporangia in nut-like globose or ovoid sporocarps at bases of stipes. ........................................................................................................................................................... 74. Marsilea. 12. Leaves simple or variously divided, often with numerous leaflets (pinnae) borne laterally; plants homosporous, with sporangia on abaxial blade surfaces or on specialized projections at tips of blades. 13. Sporangia borne on erect (rarely horizontal) spike-like or panicle-like clusters from near the base of the blades or stipes. 14. Erect (rarely horizontal) fertile structures (modified pinnae) 2 per blade (more than 2 in a few species); sporangia each with an apical annulus; rhizomes at ground surface; roots wiry. ........................................................ 8. Anemia. 14. Erect fertile structures 1 (rarely 6–8, then plants epiphytic); sporangia lacking an annulus; rhizomes usually subterranean (rarely epiphytic); roots fleshy. 15. Blades compound; fertile portion of each fertile blade a panicle-like structure. ........................................... 21. Botrychium. 15. Blades simple (or dichotomously forking, epiphytic); fertile portion of each fertile blade a spike-like structure (rarely 6–8 “spikes”; if so, then plants epiphytic). ........................................................................ 89. Ophioglossum. 13. Sporangia on back of blades or at least not on erect spike-like or panicle-like clusters from base of blades or stipes, sometimes borne on modified blades or parts of blades. 16. Leaves (but not rhizomes) climbing, the rachises twining on other plants; each sporangium subtended by an indusium. ........................................................................................................................................................................... 71. Lygodium. 16. Leaves not climbing (scandent or forming thickets in a few genera); each sporangium not subtended by an indusium, indusia if present covering clusters of sporangia. 17. Blades forking dichotomously, over 15 cm tall; plants terrestrial. 18. Leaves less than 0.5 m tall, lacking buds or continued growth in axils of forks; sporangia borne on flag- or finger-like appendages at blade tips; plants rare. 19. Fertile parts undivided. ......................................................................................................................... 2. Actinostachys. 19. Fertile parts pinnate. .................................................................................................................................. 111. Schizaea. 18. Leaves 0.7–4 m tall, with buds or continued growth at forks; terminal leaf branches pectinate; sporangia borne in round sori on abaxial blade surfaces; plants common. 20. Veins 1-forked between costae and margins; abaxial blade surfaces scaly, at least on major veins; rhizomes scaly; sori generally with 4–6 sporangia. 21. Frond apices always dormant with two large bipinnate or pinnate-pinnatisect pinnae; scales on dormant “buds” entire, stramineous, 6–8 ⫻ 1.5–2.5 mm; spores tetrahedral. ............ 41. Diplopterygium. 21. Frond apices with successive dormant “buds” and segments beyond the ultimate dichotomy 1-pinnate; scales on dormant “buds” ciliate, orange-brown, less than 3 ⫻ 1 mm; spores bilateral. .............................................................................................................................................. 116. Sticherus. 20. Veins 2–3-forked between costae and margins; abaxial blade surfaces glabrous, often glaucous; rhizomes hairy; sori generally with more than 6 sporangia. 22. Frond forks subtended by accessory pinnae; pinnae branching equally; axes terete; spores tetrahedral. .................................................................................................................................... 38. Dicranopteris. 22. Frond forks lacking accessory pinnae; pinnae forking unequally to produce alternate pairs of pinnules; axes other than stipes with narrow lateral ridges; spores bilateral. ........................ 48. Gleichenella. 17. Blades not forking dichotomously (or if so, then epiphytic and less than 5 cm tall); plants terrestrial, epipetric, or epiphytic. 23. Blades fleshy; fronds each with a conspicuous stipule at base; sporangia of each sorus fused to form a synangium.

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THE PTERIDOPHYTES OF MEXICO 24. Blades 1-pinnate; fronds dimorphic; synangia linear, embedded in the blades; rhizomes obviously creeping; stipules scale-like. .........................................................................................................................35. Danaea. 24. Blades 2–4-pinnate; fronds monomorphic; synangia oblong, like an open clam, on blade; rhizomes horizontal to erect, not obviously creeping; stipules large and fleshy. ............................... 73. Marattia. 23. Blades membranaceous to coriaceous; fronds lacking stipules at bases; sporangia not fused into synangia. 25. Fertile and sterile fronds dissimilar, dimorphic (holodimorphic), or blades partially dimorphic (fronds hemidimorphic, fertile and sterile pinnae different on the same blade). 26. Fertile blades simple, bilobed, or dichotomously forking (Elaphoglossum peltatum). 27. Sterile blades only 1 cell thick (filmy ferns); apices usually extended and radicant. .. 121. Trichomanes. 27. Sterile blades firm, more than 1 cell thick; apices not radicant 28. Sporangia covering abaxial surfaces of fertile blades. ............................................ 44. Elaphoglossum. 28. Sporangia in discrete sori. ......................................................................................... 78. Microgramma. 26. Fertile blades pinnatifid, 1-pinnate, or more divided. 29. Sterile blades 1-pinnate or less divided. 30. Sterile blades pectinate or pinnatifid. 31. Rhizome apices and stipe bases scaly; spores bilateral, monolete. ................... 18. Blechnum. 31. Rhizome apices and stipe bases without scales or hairs; spores globosetetrahedral, trilete. ................................................................................................. 98. Plagiogyria. 30. Sterile blades fully pinnate. 32. Veins netted. 33. Sori marginal, protected by reflexed segment margins; rhizomes poorly developed. .................................................................................................... 24. Ceratopteris. 33. Sori covering the abaxial surface of fertile blades, or protected on all sides by strongly inrolled segment margins and forming bead-like segments; rhizomes strongly developed. 34. Fertile blades 2–3-pinnate. ................................................................ 88. Onocleopsis. 34. Fertile blades 1-pinnate. 35. Plants 2–5 m tall, in swamps and marshes; spores globose, trilete. .......................................................................................... 1. Acrostichum. 35. Plants less than 2 m tall, not aquatic, terrestrial or hemiepiphytic; spores reniform, monolete. ................................ 19. Bolbitis. 32. Veins free. 36. Plants hemiepiphytic, with rhizomes climbing trees; fertile blades 1pinnate. ........................................................................................................ 64. Lomariopsis. 36. Plants terrestrial, rhizomes not climbing; fertile blades 2-pinnate. ........... 87. Olfersia. 29. Sterile blades more than 1-pinnate. 37. Rhizomes and stipe bases clothed with hairs. ............................................................... 90. Osmunda. 37. Rhizome and stipe bases clothed with scales. 38. Plants hemiepiphytic; fertile blades with no laminar tissue. ......................... 100. Polybotrya. 38. Plants terrestrial; fertile blades with some laminar tissue. 39. Sori marginal, protected by recurved margins; blades fertile only in distal 1⁄3, glabrous. .............................................................................................. 63. Llavea. 39. Sori round to elongate on abaxial blade surfaces, margins not usually recurved; blades fertile throughout, with acicular hairs. ....................... 120. Thelypteris. 25. Fronds monomorphic or nearly so. 40. Sori marginal to submarginal on the blades. 41. Sori in marginal or submarginal cups or tubes or submarginal trough facing the margins, the blade margins not recurved. 42. Blades very thin (1 cell thick); receptacle elongated, filiform, often projecting beyond the soral cup or tube; involucres (indusia) bivalvate or tubular. 43. Involucres bivalvate; receptacles usually not protruding. ........................ 56. Hymenophyllum. 43. Involucres tubular to conical; receptacles usually protruding well beyond the involucres. ........................................................................................................... 121. Trichomanes. 42. Blades generally more than 1 cell thick; receptacles not greatly elongated, not filiform; involucres (indusia) bivalvate, cup-like, or linear. 44. Stems stout, creeping, suberect, or erect and trunk-like, hairy; fronds large; indusia thick. 45. Stems forming upright trunks 1–2 m tall. ................................................. 37. Dicksonia. 45. Stems stout, creeping or suberect, but hardly emerging above the soil. 46. Stems creeping; blades Ⳳ 3-pinnate; costae and costules raised or

THE PTERIDOPHYTES OF MEXICO

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only slightly furrowed, the ribs (if any) not decurrent onto axis of next order. ............................................................................................. 27. Cibotium. 46. Stems erect, trunk-like, but very short; blades 4–5-pinnate; costae and costules deeply sulcate adaxially, the furrows flanked by prominent ribs decurrent onto axes of next order. ............................. 31. Culcita. 44. Stems slender, creeping, scaly or hairy, if erect, fronds not large; indusia thin, not coriaceous. 47. Fronds small, less than 25 cm long; plants epiphytic. ............................ 68. Loxoscaphe. 47. Fronds large, 30–200(–500) cm long; plants terrestrial. 48. Sori continuous along the margins, much longer (along margins) than wide. .............................................................................................. 62. Lindsaea. 48. Sori interrupted, short, not more than twice as long (along margins) as wide. 49. Sori slightly back from the blade margins; rhizomes bearing scales. ......................................................................................... 108. Saccoloma. 49. Sori on the blade margins; rhizomes bearing scales or hairs. 50. Fronds arching, not scrambling, blades 2–3-times divided, the segments not linear, over 5 mm wide; rhizomes hairy. ............................................................. 36. Dennstaedtia. 50. Fronds erect or scrambling, blades finely dissected, 4–5pinnate, the segments linear, 1–2 mm wide; rhizomes scaly. 51. Fronds 1.7–5 m long, scrambling; sori each served by a single vein. ................................................... 85. Odontosoria. 51. Fronds 20–60 cm long, erect; sori served by 1–4 veins. .................................................................... 115. Sphenomeris. 41. Sori not in marginal pockets facing the margins, but rather protected by recurved blade margins, or sori open without protection of either true or false indusia. 52. Rhizomes usually long-creeping, hairy; fronds large, usually 1 m or more long; stem branches sometimes arising from bases of stipes. 53. Veins netted. .......................................................................................................... 53. Histiopteris. 53. Veins free. 54. Rhizomes short-creeping, succulent; blades membranous; stipes and blades hairy. ..................................................................................................... 65. Lonchitis. 54. Rhizomes long-creeping, cord-like, firm; blades thin to coriaceous; stipes and blades hairy to glabrous. 55. Sori discrete; inner indusia absent; rhizomes at soil level; spores bilateral; axes often spiny. .................................................................... 57. Hypolepis. 55. Sori continuous; faint inner indusia present; rhizomes deep underground; spores tetrahedral; axes unarmed. ........................... 105. Pteridium. 52. Rhizomes short-creeping to erect, scaly; fronds small to large; stem branches absent from stipe bases. 56. Blades undivided to deeply lobed, but not 1-pinnate or more divided. 57. Blades deeply palmately lobed. .................................................................. 42. Doryopteris. 57. Blades unlobed. 58. Plants terrestrial; blades net-veined with included veinlets. ...................................................................................... 118. Tectaria (panamensis). 58. Plants epiphytic or epipetric; blades net-veined without included veinlets. 59. Blades linear-elliptic, 9–12 mm wide, with 2–4 rows of areoles between costae and margins. ................................... 7. Ananthacorus. 59. Blades linear, 1–4 mm wide, with a single row of areoles between costae and margins. 60. Stipes red-brown; spores bilateral. .......................... 107. Radiovittaria. 60. Stipes greenish or yellowish; spores tetrahedral, rarely bilateral. ............................................................................... 122. Vittaria. 56. Blades 1-pinnate to several times pinnately divided. 61. Sporangia borne on undersides of the indusia, not on blade tissue. .......5. Adiantum. 61. Sporangia not borne on undersides of the indusia, protected by reflexed and modified segment margins or not. 62. Sori paraphysate; awns usually present on top of axes where

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THE PTERIDOPHYTES OF MEXICO segments diverge; mostly large ferns with fronds over 80 cm long (fronds rarely lacking awns, 20–60 cm long, 1-pinnate with 20–70 pairs of linear pinnae); spores each with an equatorial flange. ........... 106. Pteris. 62. Sori lacking paraphyses; awns absent; fronds mostly less than 60 cm long; spores lacking an equatorial flange. 63. Stipes and rachises with dense, broad scales; segments beadlike; spores bilateral; stipes with several vascular bundles; plants of high elevation. .............................. 102. Polystichum (speciosissimum). 63. Stipes and rachises with narrow scales or lacking scales; segments rarely bead-like (a few species of Cheilanthes); spores tetrahedral; stipes with 1–2 vascular bundles; plants of low to high elevations. 64. Adaxial grooves of stipes and rachises densely clavatehirsute, hairs ⬍0.5 mm, unicellular, clavate, rigid; blade margins (visible as greenish teeth abaxially) projecting beyond the scarious false indusia. .................................... 80. Mildella. 64. Adaxial grooves of stipes and rachises not densely clavate-hirsute, hairs, if present, usually flexuous, septate; false margins absent. 65. Stipes stramineous to light brown, if atropurpureous, then glabrous or with flexuous, septate hairs 0.5–0.6 mm long; pinnae ternate or simply pinnate with fewer than 5 pairs of segments; axes lacking scales. .................................... 92. Pellaea. 65. Stipes atropurpureous or castaneous, or if lighter, then glandular-viscid; pinnae not ternate, rarely simply pinnate; axes often scaly in Cheilanthes. 66. Blades glabrous (except for 0.1 mm clavate hairs abaxially in Adiantopsis), thin to chartaceous; sori discrete, usually many per ultimate segment. 67. Blades digitate (pinnae radiating from stipe apices) or pinnate; segments obtuse, oblong; abaxial segment surfaces with minute (less than 0.1 mm) white clavate hairs. .......................................... 4. Adiantopsis. 67. Blades pinnately divided; segments acuminate, linear, 0.5–1.3 mm wide; abaxial segment surfaces glabrous. ........ 13. Aspidotis. 66. Blades scaly, hairy, or glabrous; sori continuous, or if discrete, blades firm to coriaceous. 68. Abaxial blade surfaces farinose; marginal indusia absent. .................................... 84. Notholaena. 68. Abaxial blade surfaces lacking farina, or if present then with conspicuous marginal indusia; marginal indusia usually present though sometimes poorly differentiated. 69. Blades 1–4-pinnate; segments beadlike to linear-lanceolate; veins usually hidden; indusia continuous to interrupted, broad to narrow, Ⳳ plane or S-shaped toward the margin. ........................................ 25. Cheilanthes. 69. Blades pinnate-pinnatifid to 2pinnate above basal pinnae; veins prominulous; indusia continuous, broad, convex, involute (inrolled so as to completely envelop sporangia. ................................. 26. Cheiloplecton.

THE PTERIDOPHYTES OF MEXICO

11

40. Sori abaxial on blades, along the veins, midveins, or spread across the entire undersurface of blades. 70. Sori (sporangia) spread across the entire abaxial surfaces of blades; indusia absent. 71. Blades undivided. ....................................................................................................... 44. Elaphoglossum. 71. Blades divided. 72. Blades with white to pale yellow farina abaxially. ....................... 97. Pityrogramma (trifoliata). 72. Blades lacking farina abaxially. 73. Fronds large (2 m or more long), coriaceous; rhizomes short-creeping, stout; plants of swamps. .............................................................................. 1. Acrostichum. 73. Fronds generally less than 1 m long; rhizomes long-creeping, slender; plants of forests. 74. Veins netted; plants terrestrial or hemiepiphytic. ................................ 19. Bolbitis. 74. Veins free; plants hemiepiphytic. .................................................... 64. Lomariopsis. 70. Sori abaxial on blades, discrete, usually running along the veins, rarely the sporangia sparsely distributed singly or in small groups between veins; indusia present or absent. 75. Sori elongate along the veins, rarely the sporangia sparsely distributed singly or in small groups, exindusiate. 76. Blades simple and unlobed, or tiny and forking at tips (Hecistopteris); rhizomes usually with clathrate scales; plants epiphytic, rarely epipetric. 77. Sporangia sparsely distributed (singly or in small groups) on the abaxial blade surfaces. ................................................................................................... 9. Anetium. 77. Sporangia densely arranged in sori along the veins. 78. Sori in grooves near and parallel to the blade margins and extending the length of the blades or at least the distal third. 79. Sori in 1 or 4 (or 6) rows. 80. Sori on each blade in one line in a groove along the costa. ............................................................................... 29. Cochlidium. 80. Sori on each blade in 4 (or 6) rows (2 or 3 rows on each side of the costa). ............................................... 103. Polytaenium. 79. Sori in 2 rows (one on each side of the costa). 81. Blades 1–4 mm wide; 1 row of areoles between costae and margins; plants generally epiphytic. 82. Stipes red-brown; spores bilateral. ................ 107. Radiovittaria. 82. Stipes greenish or yellowish; spores tetrahedral, rarely bilateral. .......................................................... 122. Vittaria. 81. Blades 9–12 mm wide; 2–4 rows of areoles between costae and margins; plants generally epipetric. 83. Sori along the length of the blades. ................. 7. Ananthacorus. 83. Sori confined to distal 1⁄3 of blades. .................. 82. Neurodium. 78. Sori not extending the full length of the blades, parallel to the costae or at an angle, or following the vein pattern. 84. Blades tiny, less than 4 cm long, forking. .......................... 49. Hecistopteris. 84. Blades larger, never forking. 85. Rhizome scales not clathrate or weakly clathrate and bicolored; blades with minute scales. ............................. 99. Pleopeltis. 85. Rhizome scales clathrate, lumina clear; blades lacking scales. 86. Costae reaching blade apices. .......................... 67. Loxogramme. 86. Costae not reaching blade apices. 87. Blades thin to coriaceous, but not fleshy; sori surficial or in grooves; paraphyses absent; spores tetrahedral; rhizome scales 2–4 mm long, less than 1 mm wide; blades 1.5–2 mm wide at bases. ............................................. 103. Polytaenium. 87. Blades fleshy; sori short and parallel to one another, embedded in the laminae, rarely surficial; paraphyses abundant (rarely scattered), short (0.3–0.5 mm long), reddish; spores bilateral; rhizome scales 10–12 ⫻ 1.5–2 mm; blades (2–)6–7 mm wide at bases. ... 112. Scoliosorus.

12

THE PTERIDOPHYTES OF MEXICO 76.

Blades deeply lobed to 3-pinnate; rhizomes usually lacking clathrate scales; plants terrestrial or epipetric. 88. Rhizomes with hairs. 89. Blades broadly elliptic to deltate, 3–4-pinnate, not permanently circinate at apices. ..... 47. Eriosorus. 89. Blades linear, 1-pinnate, Ⳳ permanently circinate at apices. ............................................ 59. Jamesonia. 88. Rhizomes with scales. 90. Blades simple, palmately lobed. 91. Veins netted; sporangia following veins more than the distal 2⁄3 of their length. ............................................................................................................................. 52. Hemionitis. 91. Veins free or netted; sporangia limited to distal 1⁄3–2⁄3 of the veins. ..................... 20. Bommeria. 90. Blades 1-pinnate or more divided. 92. Blades linear, pinnate to pinnate-pinnatifid, not pedate, the pinnae densely scaly abaxially, scales totally obscuring the blade surfaces. .................................................. 15. Astrolepis. 92. Blades linear, lanceolate, ovate or pedate, 1–3-pinnate, the pinnae not densely clothed with scales, or if scaly, blades more than 2-pinnate. 93. Blades with white to pale yellow farina abaxially. 94. Blades pedate, pinnate-pinnatifid above the proximal pair; Baja California. .............................................................................................. 93. Pentagramma. 94. Blades not pedate, bipinnate to bipinnatifid above the proximal pinna pair; only rarely in Baja California. 95. Fronds 10–30 cm tall; ultimate segments petiolulate. .............. 12. Argyrochosma. 95. Fronds 30–200 cm tall; ultimate segments adnate. ................ 97. Pityrogramma. 93. Blades lacking farina abaxially. 96. Ultimate segments petiolulate, discrete; stipes and rachises castaneous to atropurpureous or black. ................................................................. 12. Argyrochosma. 96. Ultimate segments not petiolulate, not discrete; stipes and rachises stramineous to tan. 97. Fronds 2–25 cm long; blades 2-pinnate to 4-pinnatepinnatifid. ......................................................................................... 10. Anogramma. 97. Fronds (12–)20–200⫹ cm long; blades pinnate to pinnatepinnatifid, rarely 2-pinnate. 98. Blades pinnate-pinnatifid, bearing acicular hairs. ............. 120. Thelypteris. 98. Blades pinnate or pinnate-pinnatifid to 2-pinnate; hairs, if present, jointed, not acicular. 99. Blades 1-pinnate; sori extending nearly the entire length of the veins. ......................................................... 52. Hemionitis. 99. Blades pinnate-pinnatifid to 2-pinnate at bases; sori limited to the distal 1⁄3 to 1⁄2 of the veins. ............. 51. Hemionanthes. 75. Sori round to reniform or oblong-elliptic, if elongate along the veins, then generally indusiate. 100. Sori costal, elongate. 101. Sori continuous along costae; blades 1-pinnate, or less divided. ................................................ 18. Blechnum. 101. Sori interrupted along costae, chain-like; blades pinnate-pinnatifid. ...................................124. Woodwardia. 100. Sori not costal, borne between midveins and margins. 102. Sori elongate. 103. Sori hooked over vein at end, horseshoe-shaped, or occasionally back-to-back on both sides of veins. 104. Veins obscure, free; sori continuous, each one from one side of the vein to the other around the end of the vein, horseshoe-shaped. ....................................... 39. Didymochlaena. 104. Veins visible, free or netted; sori sometimes double, but then not continuous around ends of veins. 105. Fronds very large (2–3 m long), 1-pinnate; veins free near the midveins and netted near the margins. ............................................................................... 50. Hemidictyum. 105. Fronds less than 1 m long, 1–3-pinnate; veins free or netted, but not both on the same pinna. 106. Sori one per vein, borne on one side of vein or hooked over veins .... 16. Athyrium. 106. Sori, at least some, double, the indusia back-to-back on the same vein. ............................................................................................................ 40. Diplazium. 103. Sori simple, or if double, then face to face rather than back to back. 107. Blades simple to 3-pinnate; sori single (one per vein). 108. Blades pinnate-pinnatifid; blades and rhizome scales usually with acicular hairs. .................................................................................................................... 120. Thelypteris.

THE PTERIDOPHYTES OF MEXICO

13

108. Blades simple to 3-pinnate; blades and rhizome scales mostly glabrous, lacking acicular hairs (except Asplenium pumilum). 109. Veins free; blades simple to 3-pinnate. ................................................... 14. Asplenium. 109. Veins netted; blades simple. 110. Indusia present; blades without scales. ....................................... 54. Holodictyum. 110. Indusia absent; blades with minute scales abaxially. ...................... 99. Pleopeltis. 107. Blades simple; most sori double, face-to-face on adjacent veins. 111. Blades linear-lanceolate, 7–24 cm long, 4–8 times as long as wide; stipes green. ..................................................................................................................... 14. Asplenium. 111. Blades flabellate, 2–6 cm long, about as wide as long; stipes atropurpureous. .................................................................................................................... 110. Schaffneria. 102. Sori round (to slightly oblong). 112. Tree ferns with trunks 2–20 m tall, usually with large fronds; indusia globose, cup-like or scale-like, attached at base of sori, or indusia absent. 113. Stipe scales linear, whitish, with short dark marginal teeth; cells of scale body uniform in size, shape, and orientation; indusia globose. .................................. 114. Sphaeropteris. 113. Stipe scales broadly ovate to linear, whitish to atropurpureous, without dark marginal teeth (except Cyathea bicrenata and C. myosuroides); cells of scale body differentiated in size, shape, and orientation (C. myosuroides only slightly so); indusia globose, cup-shaped, scale-like, or absent. 114. Scales of the stipes, rachises, and costae without black acicular setae; stipe bases unarmed, or if spinose, the spines not black. ........................................... 32. Cyathea. 114. Scales of the stipes, rachises, and costae with black apical and usually lateral acicular setae; stipe bases generally spinose, the spines black and stout. ........ 6. Alsophila. 112. Plants not trees; indusia various or absent. 115. Fronds large, 2–4 m long; rhizomes with scales or hairs. 116. Sori with saucer-like indusia; rhizomes with scales. ..................................... 28. Cnemidaria. 116. Sori exindusiate; rhizomes with dense, long hairs. 117. Fronds 3–4-pinnate, glaucous abaxially. ................................................ 66. Lophosoria. 117. Fronds 1-pinnate, not glaucous abaxially. ................................................. 77. Metaxya. 115. Fronds generally less than 2 m long; rhizomes with scales. 118. Blades simple; stipes distinctly articulate on phyllopodia 5–10(–30) mm long; veins free, Ⳳ parallel; sori indusiate. ................................................................ 86. Oleandra. 118. Blades simple to several times divided; stipes not articulate, or if articulate, then cutting off cleanly from the rhizomes, lacking phyllopodia or phyllopodia less than 5 mm long; veins free or anastomosing, not parallel; sori indusiate or exindusiate. 119. Blades simple to 1-pinnate, exindusiate. 120. Fronds with long hairs, or glabrous; spores tetrahedral, green. 121. Hydathodes absent or inconspicuous adaxially. 122. Blades simple, differentiated into sterile and fertile parts, the sterile leaves and at least the sterile portion of fertile leaves serrate-dentate. ................ 29. Cochlidium (serrulatum). 122. Blades simple to pinnate, not subdimorphic, nor differentiated into sterile and fertile portions. 123. Sori of young leaves with conspicuous globose paraphyses, these more or less whitish, at times yellowish. ................................................................. 23. Ceradenia. 123. Sori lacking conspicuous globose paraphyses. ... 45. Enterosora. 121. Hydathodes present at the vein endings adaxially. 124. Blades simple, repand or at most serrate-dentate in the sterile leaves or in the sterile portion of fertile leaves; castaneous setae absent. ............................................... 29. Cochlidium. 124. Blades deeply lobed to pinnate in the fertile portion of the leaves; hairs often branched; castaneous setae present or absent. 125. Rhizome scales strongly clathrate, sometimes iridescent, entire, without marginal setae or cilia, but sometimes glandular at the apices; rhizomes long- or short-creeping, dorsiventral; stipes and/or rachises generally with castaneous setae. .......... 76. Melpomene.

14

THE PTERIDOPHYTES OF MEXICO 125. Rhizome scales non-clathrate or clathrate, if clathrate then generally with marginal hairs; rhizomes compact and radially symmetrical or sometimes short-creeping; stipes and/or rachises with castaneous setae or hyaline hairs. 126. Veins of the segments (pinnae) simple or with a single acroscopic branch; blades less than 1 cm wide, linear. 127. Rhizome scales clathrate, blackish brown; veins of segments simple; castaneous setae absent on axes and blades. .................................................. 61. Lellingeria. 127. Rhizome scales concolorous, golden to castaneous, not clathrate; veins of the segments simple or each with a single acroscopic branch; castaneous setae generally present on the blade axes and often on blades. ....................... 79. Micropolypodium. 126. Veins of the segments (pinnae) branched more than once; blades more than 1 cm wide, linear to narrowly ovate. 128. Rhizome scales clathrate, setose; sori generally slightly to deeply sunken; stipes and rachises with hairs and/or small hyaline setae 0.1–0.3 mm long, without castaneous setae; rhizomes radially symmetrical. .......................... 61. Lellingeria. 128. Rhizome scales not clathrate; sori surficial (not sunken); stipes and rachises with castaneous setae generally more than 0.5 mm long; rhizomes short-creeping to ascending, usually dorsiventral. .......................................119. Terpsichore. 120. Fronds with scales, at least at the base, or glabrous; spores bilateral, golden or whitish. 129. Blades simple. 130. Sori in one row on each side of costae. 131. Sori round, with filamentous, sometimes branched paraphyses. ...................................... 78. Microgramma. 131. Sori oval to elongate, with peltate, scale-like paraphyses (often shed prior to sorus maturity). .................................................................. 99. Pleopeltis. 130. Sori in 2 or more rows on each side of costae. 132. Sori in 2 rows between main lateral veins; sori lacking paraphyses; sporangia glabrous. .... 22. Campyloneurum. 132. Sori in 1 row between main lateral veins; sori with paraphyses (aborted sporangia) present; sporangia with multicellular hairs. ...................................... 83. Niphidium. 129. Blades pinnatifid, pinnate, or more divided. 133. Veins free. 134. Rhizome scales peltate, clathrate or not; stipes grooved, stramineous, rarely dark; blades pinnatifid, pectinate, or fully pinnate toward blade base. ........................................................... 101. Polypodium. 134. Rhizome scales basally attached, never clathrate; stipes terete, black to reddish brown; blades pectinate. ................................................................... 91. Pecluma. 133. Veins netted. 135. Sori with peltate, scale-like paraphyses (often shed prior to sorus maturity). ......................................... 99. Pleopeltis.

THE PTERIDOPHYTES OF MEXICO

15

135. Sori with branched or filamentous paraphyses, or paraphyses absent. 136. Sori at junction of double included veins; costal areoles lacking included veinlets, some additional areoles lacking included veins, some with single included veinlets, and others with paired included veinlets; sori non-paraphysate; sporangia glabrous. ...................................................... 95. Phlebodium. 136. Sori terminal on free included veins; areoles each with a single included veinlet; sori paraphysate or non-paraphysate; sporangia glabrous or setose. .................................... 101. Polypodium. 119. Blades 1–4-times pinnate; sori mostly indusiate (if exindusiate, blades more than 1-pinnate). 137. Indusia forming a hood or basal cup. 138. Indusia hood-shaped. .............................................................. 34. Cystopteris. 138. Indusia cup-shaped or in 4 flat lobes beneath the sori, the lobes sometimes fimbriate. ....................................................... 123. Woodsia. 137. Indusia reniform, peltate, or absent. 139. Veins netted. 140. Stipe vascular strands 2; blades with acicular hairs. ............................................................................... 120. Thelypteris. 140. Stipe vascular strands more than 2; blades lacking acicular hairs. 141. Veins anastomosing only toward pinna margins; blades coriaceous. .......................................... 94. Phanerophlebia. 141. Veins abundantly and finely anastomosing throughout pinna width; blades herbaceous to chartaceous. ............................................................... 118. Tectaria. 139. Veins free. 142. Indusia peltate (sometimes shriveling or shed early). 143. Fronds 7–10 cm long; blades pinnatepinnatifid. ............................................................... 3. Adenoderris. 143. Fronds (15–)30–200 cm long; blades 1–3-pinnate. 144. Blades 2–3-pinnate, if 1-pinnate, the pinnae more than 20 pairs per blade (Polystichum acrostichoides, P. munitum) or less than 2.5 cm long (P. muenchii) and not reduced proximally. ................................................ 102. Polystichum. 144. Blades 1-pinnate, fewer than 20 pairs per blade, or if more than 20 pairs then 2–5 pairs of proximal pinnae reduced. 145. Proximal pinnae not shortened or deflexed; pinnae⬍20 pairs per blade. ............................................ 94. Phanerophlebia. 145. Proximal 2–5 pinna pairs somewhat shortened and deflexed; pinnae 30⫹ pairs per blade. .................................... 33. Cyclopeltis. 142. Indusia reniform or absent. 146. Stipe bases with 2 vascular strands; blades with acicular hairs. 147. Blades 2-pinnate-pinnatifid. .............. 72. Macrothelypteris. 147. Blades simple or pinnate to pinnatepinnatifid. ................................................... 120. Thelypteris. 146. Stipe bases with more than 2 vascular strands; blades lacking acicular hairs. 148. Blades 1-pinnate. 149. Pinnae thin, oblong to narrowly deltate, often abscising, often with

16

THE PTERIDOPHYTES OF MEXICO lime dots adaxially; sori indusiate; plants with wiry stolons. .................. 81. Nephrolepis. 149. Pinnae coriaceous, ovate to linearlanceolate, not abscising, lacking lime dots adaxially; sori exindusiate; plants lacking stolons. ........................... 94. Phanerophlebia. 148. Blades more than 1-pinnate. 150. Blades catadromous, at least in proximal portion. 151. Hairs on adaxial axes more than 0.5 mm long, whitish, acicular, round, at least at their bases; basal basiscopic veins arising from costae rather than costules; vein endings clavate adaxially; indusia absent. ..................................... 75. Megalastrum. 151. Hairs on adaxial axes less than 0.5 mm long, reddish, with blunt apices, flattened, adjacent cells twisted; basal basiscopic veins arising from costules; vein apices not clavate adaxially; indusia present or absent. 152. Rachis buds present near blade tips. .......................... 60. Lastreopsis. 152. Rachis buds absent. 153. Rachises and costae adaxially with short, reddish, pluricellular hairs. ............................. 30. Ctenitis. 153. Rachises and costae adaxially without short, reddish, pluricellular hairs. .......................... 43. Dryopteris. 150. Blades anadromous. 154. Indusia absent. 155. Blades pinnate to pinnatepinnatifid; segments rounded or acute at tips. ............................... 117. Stigmatopteris. 155. Blades 2–4-pinnate. 156. Segments rounded at tips. ....................... 75. Megalastrum. 156. Segments spinulose or mucronate at tips. ....................... 102. Polystichum. 154. Indusia present. 157. Blades 3–4-pinnate; segments mucronate, dark green. ................................ 11. Arachniodes. 157. Blades 2–3-pinnate; segments green to yellowish green, with tips acute to acuminate, but not mucronate. ......................... 43. Dryopteris.

ACROSTICHUM

1 . A C R O S T I C H UM Acrostichum L., Sp. Pl. 2: 1067. 1753. Lectotype (chosen by J. Sm., Hist. Fil. 146. 1875): Acrostichum aureum L. Chrysodium Fe´e, Me´m. Foug. 2: 22, 97. 1845, nom. superfl. for Acrostichum L.

Terrestrial in swamps and marshes; rhizomes massive, shortcreeping to erect; roots stout, fleshy; rhizome scales thick, blackish brown, to 22 ⫻ 2 mm; fronds large, 1-pinnate, each with a conform terminal pinna; pinnae elliptic, entire, coriaceous, glabrous or with minute hairs; veins finely netted, without included veinlets; sori acrostichoid on all or only the distal pinnae, with many capitate paraphyses; indusia absent; spores tetrahedral, whitish tan; x⫽30. Acrostichum inhabits marshes, swamps, and stream margins,

17

forming extensive stands 2–5 m tall in fresh or brackish water. It is a genus of perhaps as few as three tropical species: one American, one Australasian, and one pantropical. Although the genus has been generally allied with Pteris, its affinity is not close. It is especially distinct in its acrostichoid sori, many stipe bundles, large fleshy roots, large plant size, marshy habitat, and coriaceous, 1-pinnate fronds. The species are often confused due to their subtle differences. Our descriptions are based on Adams and Tomlinson (1979). References Adams, D. C. & P. B. Tomlinson. 1979. Acrostichum in Florida. Amer. Fern J. 69: 42–46; Garcı´a de Lo´pez, I. 1978. Revisio´n del ge´nero Acrostichum en la Repu´blica Dominicana. Moscosoa 1(3): 64–70; Lloyd, R. M. & T. L. Gregg. 1975. Reproductive biology and gametophyte morphology of Acrostichum danaeifolium from Mexico. Amer. Fern J. 65: 105–120.

Key to the Mexican Species of Acrostichum 1. Fronds fertile only in the distal 1⁄4–1⁄3 (5 distal pinna pairs and terminal pinna); pinna pairs usually not more than 30, proximal pinnae distant, long-stalked to 3 cm; areoles next to costae narrow, 3 times longer than wide; rachises decidedly grooved adaxially, short spines (aborted pinna midribs) frequent on distal portion of stipes; paraphyses with isodiametric, multilobed tips; pinna areoles divergent from costae at 45–60⬚; pinnae glabrous abaxially. ............................................................................................................ 1. A. aureum. 1. Fronds fertile for entire length, rarely only in distal 1⁄4; pinna pairs usually 40–60, closely set and overlapping, proximal pinnae short-stalked less than 2 cm; areoles next to costae broad, never 3 times longer than wide; rachises scarcely grooved, lacking short spines on distal portion of stipes; paraphyses with long, nearly horizontal, few-lobed or entire tips; pinna areoles divergent from pinna midveins at 60–85⬚; pinnae usually with small, stiff hairs on the abaxial surfaces. ............................................................... 2. A. danaeifolium.

1. ACROSTICHUM AUREUM Acrostichum aureum L., Sp. Pl. 2: 1069. 1753. Lectotype (chosen by Lellinger, 1985: Proc. Biol. Soc. Wash. 98: 366). Plumier, Descr. Pl. Ame´r., pl. 7, 1693, based on a specimen collected by Plumier at F. Royal, Martinique; LINN 1245.5 was not present in LINN prior to 1756 (Jackson, 1912). Figs. 1D–F. Rhizomes short-creeping, branched, massive; rhizome scales linear to ovate, dark brown, to 1.5 cm long; fronds 1.5–3 m long; stipes ca. 1⁄3 of the frond length, terete to decidedly grooved distally, short spines (aborted pinna midribs) frequent on distal stipe; pinnae 10–14(–30) pairs, obtuse to acute at apices, coriaceous, overlapping, margins recurved; proximal pinnae with stalks to 3 cm long; costal areoles narrow, 3 times longer than wide, extracostal areoles divergent from costae at 45–60⬚; fertile pinnae 1–5 pairs in distal portion of blades, similar in size and shape to the sterile pinnae; paraphyses each with isodiametric, multilobed tip; 2n⫽60 (Jam).

Distribution. In mangrove swamps and brackish marshes; 0– 30 m. USA (Fla); Mexico; Guat, Bel, Hond, Nic, CR, Pan; Bah, Gr & L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Braz, Parag; Old World tropics. Specimens Examined. Camp (Flores G. 318, ENCB, MEXU; Wendt & Chiang 292, XAL). Chis (Matuda 2672, K, MEXU, NY, UC, US). Gro (Lorea 4621, IEB, FCME). Nay (Ferris 5435, DS, US). Oax (Elorsa C. 1160, NY). Tab (Cowan 3193, CAS, ENCB, MEXU, NY). QR (Torres 13, UAMIZ). Ver (Rzedowski 17171, ENCB, NY). Yuc (Rico-Gray 74, UC, XAL).

Acrostichum aureum is restricted to coastal saline mangrove communities. It is distinguished from A. danaeifolium by the former being fertile only in the distal 1⁄4–1⁄3 of the blades (rather than nearly the entire length).

18

ACTINOSTACHYS

2. ACROSTICHUM DANAEIFOLIUM

guished by this character. Acrostichum danaeifolium is much more common than A. aureum.

Acrostichum danaeifolium Langsd. & Fisch., Pl. Voy. Russes Monde 1: 5, t. 1. 1810. Type. Brazil. Santa Catarina Island, Krusenstern s.n. (LE). Figs. 1A–C.

2 . A C T I N O ST A C H Y S

Chrysodium lomarioides Jenman, Timehri 4: 314. 1885. Acrostichum lomarioides (Jenman) Jenman, Bull. Bot. Dept., ser. 2, 5: 154. 1898, hom. illeg., non Bory, 1833. Acrostichum excelsum Maxon, Proc. Biol. Soc. Wash. 18: 224. 1905. Type. British Guiana [Guyana]: Jenman s.n. (NY!). Chrysodium lomarioides Jenman var. hastatum Christ, Bull. Herb. Boissier, se´r.2, 5: 725. 1905. Type. Mexico. Chiapas: Hot sulphur spring near El Carmen (200 miles from the coast), Mu¨nch s.n. (P, not found; isotypes DS!, US!).

Rhizomes massive, erect, infrequently branched, shortcreeping; rhizome scales linear, to 2 cm long, dark brown, bases black, narrowed, margins pale; fronds 3.5 m long, clumped; stipes stout, slightly grooved, dark brown; pinnae usually 40–60 pairs, linear-oblong, chartaceous, margins slightly recurved and cartilaginous; proximal pinnae short-stalked, stalks less than 2 cm; costal areoles broad, never more than 3 times longer than wide, extracostal areoles divergent from costae at 60–85⬚; fronds either fertile or sterile (otherwise sterile fronds rarely fertile only in distal 1⁄4), the fertile pinnae resembling the sterile in size and shape, fertile fronds somewhat taller than sterile fronds; paraphyses with long, nearly horizontal tips, entire or few-lobed; 2n⫽60 (Jam).

Actinostachys Wall., Numer. List. 1. 1829. Type: Actinostachys digitata (L.) Wall. [⬅ Acrostichum digitatum L.]. Terrestrial; rhizomes erect, bearing hairs; fronds upright, unbranched, nearly triangular in cross section, the sterile ones pointed, the fertile ones tipped with 2 to many erect to spreading terminal rays that bear sporangia in 2–4 rows; sporangia in two rows on each fertile segment, sporangia erect, oblong with terminal annuli; spores bilateral, verruculate or striate; gametophytes subterranean, not green, tuber-like, brown-hairy; x⫽134, 140, 168. Actinostachys is a genus of about 20 species (one in Mexico), nearly worldwide in tropical regions. It is related to Anemia and Lygodium, but most closely to Schizaea, with which it is sometimes combined (e.g., Tryon & Tryon, 1982). Reference Wagner, W. H., Jr. & V. Quevedo. 1985. [Abstract] Polymorphism in Actinostachys pennula (Sw.) Hook. and the taxonomic status of A. germanii (Fe´e) Prantl. Amer. J. Bot. 72: 927–928.

1. ACTINOSTACHYS PENNULA Actinostachys pennula (Sw.) Hook., Gen. Fil., t. 111A. 1842. Figs. 2A, B. Schizaea pennula Sw., Syn. Fil. 150, 379. 1806. Type. “America meridionalis,” “Habitat in America calidore,” s. coll. s.n. (S). Actinostachys germanii Fe´e, Me´m. Foug. 11: 123, t. 29, f. 3. 1866. Schizaea germanii (Fe´e) Prantl, Schiz. 132. 1881. Type. Guadeloupe. Sainte-Rose, PetitBourg, Deshaies, M. Germain s.n. (P!).

Distribution. Brackish swamps, fresh-water swamps or by streams in tropical rain forest; 0–300 m. USA (Fla); Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Berm, Bah, Gr & L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, s Braz, Bol, Parag. Selected Specimens Examined. Camp (Chan 4258, MEXU). Chis (Breedlove 33175, DS, MEXU, NY). Col (Lott 2755, ENCB, MEXU, NY, UC). Gro (Vela´zquez 739, ENCB, FCME). Jal (McVaugh 21025, MEXU, MICH, NY). Mich (Dı´az B. & Pe´rez de la R. 7502, IEB). Nay (Gonza´lez & Vela´zquez 866, MEXU). Oax (Conzatti 4418, US). QR (Ucan & Flora 984, XAL). SLP (Carranza & Dı´az B. 4731, IEB, MEXU) Tab (Cowan 1946, CAS, MEXU). Tam (Rzedowski 24586a, ENCB, NY). Ver (Novelo 424, MEXU, MO, XAL). Yuc (Darwin & White 2198, MEXU, MO).

Most collections of A. danaeifolium bear numerous, minute, erect hairs on the abaxial blade surfaces and can often be distin-

Rhizomes erect, ca. 1 mm diam. with stiff, bristle-like hairs; fronds dimorphic, the sterile ones 5–50 cm ⫻ 0.5–1 mm, glabrous; stipes nearly the entire length of the frond, only the fertile portions extending beyond; fertile fronds similar to the sterile except bearing at apex 2–6 linear rays 3–6 ⫻ 0.7–0.9 mm, bearing yellowish hairs and abaxially two rows of sporangia; 2n⫽168 (Trin).

ADENODERRIS

19

alliance with the athyrioid ferns (Tryon & Tryon, 1982), and it was treated as a member of the Woodsiaceae (containing Athyrium, Diplazium, et al.) by Moran (in Davidse et al., 1995: 227). That alliance was refuted by Kato (1984), who suggested that a relationship with the dryopteroids was more likely. Unpublished molecular work by Cranfill (pers. comm.) suggests that Adenoderris is indeed a member of the dryopteroid clade, possibly closely related to Polystichum. The base chromosome number for the genus is unknown, but would be expected to be x⫽41. References Kato, M. 1984. Notes on the two fern genera Adenoderris and Cheilanthopsis. J. Jap. Bot. 59: 13–19; Maxon, W. R. 1905. Adenoderris, a valid genus of ferns. Bot. Gaz. 39: 366–369.

Distribution. Subperennials or perennials, in swamps and moist hummocks, on rotten stumps and decomposing litter in damp forests and open wet areas; 180–250 m. USA (Fla); Mexico; Guat, Bel, CR; Cuba, PR, L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol, Uru. Specimens Examined. Chis (Martı´nez S. 10069, MEXU, 14891, NY, XAL, 15662, 16281, ENCB, MEXU, 19306, IEB). QR (Castillo R. & Tun G. 235, UADY, XAL).

Plants treated as A. germanii are apparently dwarf or juvenile specimens of A. pennula.

3 . A DE N O DE R R I S Adenoderris J. Sm., Hist. Fil. 222. 1875. Type: Adenoderris glandulosa (C. Presl) J. Sm. [⬅ Polystichum glandulosum C. Presl, nom. nov. for Aspidium glandulosum Hook. & Grev., 1929, non Blume, 1828]. Terrestrial or epipetric (ours); rhizomes erect, small, bearing glandular scales; fronds fascicled, monomorphic, commonly less than 20 cm long, not articulate; stipes with two vascular bundles at the base, scaly primarily at the base, stipitate-glandular throughout; blades herbaceous, deeply pinnatisect to 1-pinnatepinnatifid, much longer than the stipes, anadromous; pinnae inequilateral at base, slightly more developed along acroscopic side; rachises slightly grooved adaxially, narrowly winged, costae not grooved; veins free, commonly 1–3-forked, reaching the segment margin or terminating short of the margin; indument of viscid-glandular hairs (short-stalked glands), scales and acicular hairs absent; sori round, borne along the veinlets or at the tips of veinlets; indusia peltate or subcordate and with a narrow sinus (ours); paraphyses absent; spores dark brown, ellipsoid, bilateral, with a rugose or winged perispore; x⫽? Adenoderris comprises two species, one in Cuba and Jamaica, the other in Chiapas and Guatemala; however, the relationship of the two species to each other has been questioned (Tryon & Tryon, 1982), primarily because of their rather different perispore surfaces. Affinities of Adenoderris have been suggested with Polystichum (because of the perceived peltate indusia) and Dryopteris. The vasculature of the stipe, with two bundles, suggests an

1. ADENODERRIS SORORIA Adenoderris sororia Maxon, Bot. Gaz. 39: 368. 1905. Type. Guatemala. Alta Verapaz: Between San Pedro Carcha and Senahu´, Sesisp [Sesibche´], Tu¨rckheim s.n. [Donn. Sm. 868] (US; isotypes NY!–2 sheets, UC!, YU!). Figs. 3A–C. Rhizomes short, erect; rhizome scales orangish brown or reddish brown, lanceolate, appearing minutely ciliate, with minute glands along their margins; fronds clustered, to 8 ⫻ 2.5 cm; stipes stramineous, 0.5–1.5 cm long, subterete, often very narrowly winged for part of their length, bearing a few small scales as on the rhizome and abundant, minute, gland-tipped hairs (shortstalked glands); blades herbaceous, lanceolate to oblanceolate, pinnate-pinnatifid, slightly reduced at base, narrowing to a pinnatifid apex, minutely glandular throughout; pinnae 5–12 pairs, sessile or slightly adnate and decurrent, lobed more than halfway to a flexuous costa, the basal acroscopic lobe much the largest; veins simple or 1-forked (or 2–3-forked in largest basal lobes), not reaching the margin; sori commonly 1 per lobe, terminal on the veinlets, each commonly comprising 6–12 sporangia; indusia roundish, attached at narrow sinus, minutely glandular; spore perispore winged.

Distribution. Epipetric in montane rain forests; 1200 m. Mexico; Guat. Specimens Examined. Chis (Breedlove 56215, CAS, NY).

20

ADIANTOPSIS

Only two collections of this very rare and curious species are known other than the type, the Breedlove collection and the other from Guatemala made over 100 years ago (see Stolze, 1981). Pacheco and Moran (in Davidse et al., 1995) and others have referred to the indusia as peltate, but in an isotype seen, indusia are clearly subcordate and attached at a narrow sinus.

inner edge; indusia pale, thin, 0.5–0.8 orange-tan; 2n⫽60 (Trin.)

⫻

0.3–0.5 mm; spores

4 . A D I A N T O PS I S Adiantopsis Fe´e, Me´m. Foug. 5: 145. 1852. Lectotype (chosen by C. Chr., Index Filic. XLI. 1906): Adiantopsis radiata (L.) Fe´e [⬅ Adiantum radiatum L.]. Terrestrial; rhizomes erect to short-creeping, compact, scaly; fronds small to medium-sized; stipes atropurpureous, sulcate or adaxially flattened, with narrow, membranous, brown wings; blades 1–4-pinnate to pedate or digitate, generally 2–3-pinnate in Mexican species, essentially glabrous; veins free; sori marginal, short, several per segment; spores tetrahedral; x⫽30. There are about seven species known from tropical America, but perhaps other species within Cheilanthes belong in Adiantopsis. Two species occur in Mexico. The group is only marginally distinct from Cheilanthes, being close to the C. microphylla group in its dark, sulcate axes, discrete sori, and echinate spores. Reference Maxon, W. R. 1908. The Cuban species of Adiantopsis. Contr. U.S. Natl. Herb. 10: 485, 486.

Key to the Mexican Species of Adiantopsis 1. Fronds bipinnate; 5–9 pinnae palmately arranged at distal stipe tips; veins inconspicuous. .............................. 1. A. radiata. 1. Fronds bipinnate-pinnatifid to tripinnate; pinnae pinnately arranged; veins evident. ................................ 2. A. seemannii.

1. ADIANTOPSIS RADIATA Adiantopsis radiata (L.) Fe´e, Me´m. Foug. 5: 145. 1852. Figs. 4C, D. Adiantum radiatum L., Sp. Pl. 2: 1094. 1753. Cheilanthes radiata (L.) J. Sm., J. Bot. (Hooker) 4: 159. 1841. Hypolepis radiata (L.) Hook., Sp. Fil. 2: 72. 1852. Lectotype (chosen by Lellinger, Mem. New York Bot. Gard. 23: 3. 1972). Jamaica(?). Sloane “Adiant. 3 radiatum” (BM). LINN 1252.1 was not in LINN until after 1755 (Jackson, 1912).

Rhizomes compact, ascending; rhizome scales 2–3 ⫻ 0.3 mm, bicolorous, each with a narrow, thickened black central streak and light brown margins; fronds to 45 cm tall, clumped; stipes 1⁄2–3⁄4 of the frond length, atropurpureous to black, lustrous, glabrous, terete (to shallowly grooved distally); blades bipinnate, the 5–9 pinnae radiating from the tips of stipes; costae grooved with narrow brown wings, segments dimidiate, auriculate; adaxial surfaces glabrous; abaxial surfaces with sparse, clavate, white hairs less than 0.1 mm long; veins obscure; sori marginal, short, 6–10 on both long edges of segments and even 1–3 on narrow

Distribution. Moist, wooded rocky slopes; 100–900 m. Mexico; Guat, Bel, Hond, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol, Parag, Arg. Selected Specimens Examined. Chis (Mell 2076, NY). Col (Miranda 8752, MEXU). Gro (Hinton 14703, NY). Oax (Mickel 4007, NY). Pue (Sa´nchez 759, MEXU). Rev (Patin˜o 80, MEXU). SLP (Rzedowski 9843, ENCB). Tab (Matuda 3330, MEXU). Tam (Palmer 302, NY). Ver (Purpus 2931, NY).

The distinctive frond architecture sets A. radiata apart from the other Adiantopsis species, which more closely resemble species of Cheilanthes.

2. ADIANTOPSIS SEEMANNII Adiantopsis seemannii (Hook.) Maxon, Proc. Biol. Soc. Wash. 52: 113. 1939. Figs. 4A, B. Cheilanthes seemannii Hook., Sp. Fil. 2: 85, t. 97A. 1852. Type. Northwest Mexico. Seemann 1931 (mixed with Cheilanthes microphylla) (K!, frag. NY!).

Rhizomes short, stout, ascending; rhizome scales 5 ⫻ 0.5–0.8 mm, each bicolorous with black center and brown margins, varying to dark brown, hair-like scales 1 mm long; fronds to 65 cm tall, clumped; stipes 1⁄3–1⁄2 of the frond length, atropurpureous, lustrous, glabrous, terete; blades bipinnate-pinnatifid to tripinnate, pinnately arranged, deltate, rachises terete to shallowly grooved, glabrous, the costae grooved adaxially, with brown wings; adaxial surfaces glabrous, papillate with dark vein ends; abaxial surfaces with sparse, clavate, white hairs less than 0.1 mm long; veins visible; sori ca. 0.5 mm long; indusia pale with white cells on outer surfaces, thin, broken into flaps, 0.5– 0.8 ⫻ 0.3–0.5 mm; spores tan.

ADIANTUM

21

oblong to linear, lunate, or arcuate, one to usually several per ultimate division; paraphyses absent; spores tetrahedral, usually tan to yellowish; x⫽29, 30.

Distribution. Lowland wet forests; 100–1400 m. Mexico; Guat, Bel, Hond, Nic, CR, Pan. Selected Specimens Examined. Gro (Lorea 2574, FCME, 4012, FCME, IEB). Jal (Alava 1676, UC; McVaugh 19978, IEB, MEXU, NY). Nay (Castillo C. 6075, XAL). Oax (Mickel 5214, NY; Rivera H. 1732, NY). Sin (Breedlove 43808, CAS; Gonza´lez Ortega 6026, MEXU; Vega 3815, UC).

These specimens were previously treated as A. chlorophylla (Sw.) Fe´e (Mickel & Beitel, 1988; Mickel 1992), but apparently that species is limited to southern Brazil, Paraguay, and Bolivia. Typically, A. chlorophylla has castaneous to black-centered rhizome scales, castaneous frond axes, and sori 2–3 mm long along the segment margins, as opposed to A. seemannii which has rhizome scales entirely black (or nearly so), frond axes black, and sori only 1 mm long. Andean material identified as A. chlorophylla may be distinct from both A. seemannii and A. chlorophylla.

5 . A DI A N T UM Adiantum L., Sp. Pl. 2: 1094. 1753. Lectotype (chosen by J. Smith, Hist. Fil. 274. 1875): Adiantum capillus-veneris L. Hewardia J. Sm., J. Bot. (Hooker) 3: 432, t. 16, 17. 1841. Type: Hewardia adiantoides J. Sm. [⬅ Adiantum adiantoides (J. Sm.) C. Chr.].

Terrestrial or epipetric; rhizomes short- to long-creeping or suberect, scaly; fronds small to large, usually monomorphic; stipes castaneous to atropurpureous, lustrous, glabrous to densely covered with narrow scales or hairs, brittle; blades simple to 5 times pinnate (or pedate); ultimate divisions (pinnae, pinnules, or pinnulets, depending on blade dissection) often rhomboidal, trapezoidal, or flabellate, sessile to short-petiolulate, never adnate, deciduous (articulate) in some species; laminae generally glabrous, sometimes glaucous, some species with stiff hairs or hairlike scales on the blades; veins free, forking, rarely anastomosing, sometimes with linear epidermal idioblasts (false veins) between the true veins; sporangia borne on the veins of recurved, membranous margins, indusia (and thus also sori) reniform to short-

Worldwide, there are about 200 species of Adiantum, with more than half of them in tropical America; a few reach into temperate regions. About 35 species occur in Mexico, of which 13 are widespread into South America and the West Indies. Two species (A. aleuticum, A. jordanii) are found primarily in temperate North America and reach Mexico only in the extreme northwestern states. Five species are known so far only from Mexico (A. amblyopteridium, A. galeottianum, A. mcvaughii, A. oaxacanum, A. shepherdii), and two others are relatively rare outside of Mexico (A. feei, A. tricholepis). Adiantum species grow mostly at low to middle elevations in wet forests; one (A. capillusveneris) is pantropical and subtropical in moist microhabitats in otherwise dry regions. Adiantum andicola and A. poiretii generally occur at middle and high elevations, above the elevation of most other adiantums. Characters important in grouping and keying the species include the branching pattern of the blades (pinnate or imparipinnate; with or without a conform terminal pinna), degree of division of blades (pinnate, 2-pinnate, or more divided), presence or absence of venuloid idioblasts (epidermal false “veins”), shape of the ultimate divisions (whether dimidiate or flabellate), indument of blades and axes (whether of hairs, scales, or neither), scale margins (entire, denticulate, or ciliate), venation (free or anastomosing), vein ends (whether in teeth or sinuses), and presence or absence of articulation zones at the bases of the ultimate divisions. This last feature is presumably an adaptation to periodically drier habitats. The affinities of Adiantum are with genera of Pteridaceae, but the genus is clearly somewhat isolated and is often treated in a family of its own, Adiantaceae, or in its own subfamily, Adiantoideae, within Pteridaceae (e.g., by R. Tryon et al. in Kubitzki, 1990). In most published trees, Adiantum is sister to the vittarioid ferns, which are usually treated as a separate family Vittariaceae (e.g., Hasebe et al., 1995; Cranfill, unpubl. data). Sori are borne on the veins (and, reportedly, between the veins) of the strongly recurved and highly modified margins (false indusia), so that the sporangial capsules oppose the abaxial blade surfaces. This character is an autapomorphy for the genus. The indusia may be either round-reniform, lunate, oblong, or linear. References Fernald, M. L. 1950. Adiantum capillus-veneris in the United States. Rhodora 52: 201–208; Morton, C. V. 1955. Notes on Adiantum in Mexico, I. Amer. Fern J. 45: 113–117; Palacios-Rios, M. & R. Riba. 1983. Helechos de Veracruz: Adiantum (Pteridaceae). Bol. Soc. Bot. Me´xico 44: 43–62; Paris, C. A. & M. D. Windham. 1988. A biosystematic investigation of the Adiantum pedatum complex in eastern North America. Syst. Bot. 13: 240–255; Rojas, A. F. 2001. Informes en la familia Pteridaceae (Filicales) para el Neotro´pico. Revista Biol. Trop. 49: 429–434; Scamman, E. 1960. The maidenhair ferns (Adiantum) of Costa Rica. Contr. Gray Herb. 187: 3–22.

22

ADIANTUM

Key to the Mexican Species of Adiantum 1. Blades 1-pinnate. 2. Proximal pinnae on long stalks about 20–30 mm; stalk color ending at pinna bases; veins ending in sinuses; stipes, rachises, and blade surfaces glabrous. 3. Blades often rooting at tips; pinnae more than 5 lateral pairs, lunate or semicircular, ca. 1–2(–4) cm long; blades dull green abaxially. ................................................................................................................................................................................ 15. A. lunulatum. 3. Blades not rooting at tips; pinnae 1–4 lateral pairs, ovate, 5–10 cm long; blades glaucous abaxially. ................................. 25. A. seemannii. 2. Proximal pinnae on short stalks to 6 mm, or subsessile; stalk color usually continuing into pinna bases; veins ending in teeth; stipes, rachises, and blade surfaces glabrous or with hairs or narrow hair-like scales. 4. Largest pinnae 12–20 mm long, with rounded or flabellate tips; idioblasts absent from lamina. 5. Veins ending in teeth; pinnae stalked 3–8 mm, flabellate, cuneate at bases; rhizome scales bicolorous with dark brown center. .......................................................................................................................................................................................... 10. A. deflectens. 5. Veins ending in thickened, generally untoothed margins; pinnae sessile or short-stalked less than 1.5 mm, orbicular to oblong or rhomboid, rounded or obtusely angled at bases; rhizome scales concolorous, golden brown. 6. Pinnae not overlapping, 3–10 pairs, not dimidiate, suborbicular, distal ones about the same size as the middle and proximal ones; stipes 1⁄3–1⁄2 the frond length. .................................................................................................. 12. A. galeottianum. 6. Pinnae strongly overlapping, 34–56 pairs, middle ones strongly dimidiate and Ⳳ twice as long as broad, distal ones very gradually reduced and much smaller than the middle and proximal pinnae; stipes 1⁄10–1⁄5 the frond length. ................................................................................................................................................................................ 26. A. shepherdii. 4. Largest pinnae 35–120 mm long, with acute to acuminate tips; idioblasts present on both blade surfaces. 7. Sori 35–120 mm long, one on each pinna; stipes and rachises glabrous to sparsely filiform-scaly; largest pinnae 5–10 cm long. 8. Proximal pinnae sessile, truncate at their bases; stipes and rachises glabrous; veins free. ............................... 16. A. macrophyllum. 8. Proximal pinnae stalked 2–6 mm, rounded to subcordate at their bases; stipes and rachises sparsely filiform-scaly to hairy, glabrescent; veins anastomosing toward pinna margins. ................................................................................ 35. A. wilsonii. 7. Sori 1–6 mm long, many on each pinna; stipes and rachises with sparse to dense scales; largest pinnae 3.5–6.5 cm long. 9. Sterile pinnae coarsely serrate or biserrate; blades dull green abaxially; rachis scales moderate to dense and persistent with age. ........................................................................................................................................................... 19. A. obliquum. 9. Sterile pinnae minutely and evenly serrulate; blades glaucous abaxially; rachis scales sparse to nearly absent with age. ..................................................................................................................................................................................... 21. A. petiolatum. 1. Blades 2-pinnate or more divided, or appearing pedate (actually imparipedate). 10. Ultimate divisions with dark color stopping abruptly at their bases; veins ending in teeth. 11. Largest ultimate divisions 35–50 mm long; blades ending in a conform terminal pinna. ................................................ 30. A. trapeziforme. 11. Largest ultimate divisions 7–20 mm long; blades gradually tapering at apices. 12. Stipes and rachises with dense, stiff, reddish hairs 0.1–0.3 mm long; pinnulets abaxially with pale hairs 0.2–0.5 mm long. ..................................................................................................................................................................................................... 11. A. feei. 12. Stipes and rachises glabrous; pinnulets abaxially glabrous or with minute papillae. 13. Blade axes with pale patch at base of each division; pinnulets non-articulate; rhizome scales entire, dark brownblack. ....................................................................................................................................................................................... 5. A. braunii. 13. Blade axes uniformly dark, no pale patch at base of each division; pinnulets articulate (except in A. andicola); rhizome scales denticulate or ciliate-fimbriate, dark brown-black to golden brown. 14. Rhizome scales concolorous, dark brown to black, 1–1.5 ⫻ 0.1–0.3 mm; abaxial leaf surfaces glabrous to papillate; pinnulets mostly flabellate; rhizomes long-creeping; sori lacking yellow farina. ............................... 4. A. andicola. 14. Rhizome scales concolorous, golden brown, or bicolorous with light brown margins, more than 2 ⫻ 0.8–1 mm; abaxial leaf surfaces glabrous; pinnulets flabellate, rhombic, or trapeziform; rhizomes short-creeping, compact; sori sometimes with yellow farina among sporangia. 15. Rhizome scales 6–10 mm long, concolorous, golden brown, denticulate; segments rhombic to trapeziform; mostly Pacific lowlands. ................................................................................................................ 3. A. amplum. 15. Rhizome scales 2–3 mm long, bicolorous with light brown margins, ciliate; segments mostly cuneateflabellate; Atlantic lowlands. ............................................................................................................................. 27. A. tenerum. 10. Ultimate divisions (often pinnulets) having dark color passing into their bases; veins ending in teeth or at sinuses, or pinnulet margins without teeth, Ⳳ entire or cleft. 16. Blades gradually tapered to apices. 17. Axes with pale patch at bases of each division. ........................................................................................................................ 5. A. braunii. 17. Axes uniformly dark, no light patch at bases of each division. 18. Stipes often with minute glandular hairs; blades with only the proximal pinnae pinnate; ultimate divisions (pinnae or pinnules) suborbicular; spores brown. .................................................................................................. 12. A. galeottianum. 18. Stipes glabrous; blades with more than proximal pinnae pinnate; ultimate divisions (pinnulets) suborbicular to flabellate to trapeziform; spores tan.

ADIANTUM

23

19. Adaxial and abaxial blade surfaces and indusia with short white hairs 0.3 mm long; rhizome scales 5–6 mm long, denticulate. ....................................................................................................................................................... 32. A. tricholepis. 19. Adaxial and abaxial blade surfaces and indusia glabrous; rhizome scales less than 4 mm long, entire. 20. Sterile pinnulets serrate; veins ending in teeth; rhizome scales 2 ⫻ 0.1–0.3 mm. 21. Pinnulets distally incised or lobed often ⬎1⁄4 of the way to bases, margins proximally converging at 45–90⬚; sori (and indusia) (2–)3–11 per pinnulet, generally ⬍5 mm long; widespread in Mexico. ................................................................................................................................................ 6. A. capillus-veneris. 21. Pinnulets distally incised or lobed ⬍1⁄4 of the way to bases, or not incised at all, margins proximally converging at 90–180(–240)⬚; sori (and indusia) 1–3(–5) per pinnulet, generally ⬎5 mm long; BCN, BCS. ......................................................................................................................................... 13. A. jordanii. 20. Sterile pinnulets not serrate, or if serrate, veins ending in sinuses; rhizome scales 2.5–4 ⫻ 0.5–1 mm. 22. Pinnae subsessile, stalks 1–2 mm; pinnulets of basal pinnae overlapping rachises. ........................ 8. A. concinnum. 22. Pinnae long-stalked, stalks 2–5 mm; pinnulets of basal pinnae not overlapping rachises. 23. Pinnulets mostly rhombic or flabellate, 5–8 mm wide; sori with yellow farina; rhizome scales golden brown; sterile margins with few or no teeth; soral sinuses broad and shallow, almost absent. ..................................................................................................................................................... 22. A. poiretii. 23. Pinnulets mostly trapeziform, 8–12 mm wide; sori lacking yellow farina; rhizome scales brown; sterile margins serrate; soral sinuses very narrow, deep. ........................................................... 24. A. raddianum. 16. Blades each with a conform terminal pinna. 24. Blades appearing pedate or digitate-pedate, actually imparipedate, the two proximal pinnae 2–4 times basiscopically branched; rachises glabrous or with minute whitish hairs; idioblasts absent. 25. Rachises glabrous; blades glabrous abaxially; sterile pinnulets (or portions) with veins ending in teeth; sori oblong, each lacking a distinct sinus at the point of attachment; Chih. ................................................................... 1. A. aleuticum. 25. Rachises with minute whitish, often branched hairs 0.05–0.2 mm long; blades often with scattered hairs 0.5–1 mm long abaxially; sterile pinnulets (or portions) with veins meeting somewhat cartilaginous, entire margins; sori roundish, each with a distinct sinus at the point of attachment; widespread in w. and s. Mexico. ......................... 20. A. patens. 24. Blades pinnate; rachises with stiff, brown hairs 0.1 mm, on abaxial surfaces only, or dense, narrow, pectinate scales 2– 3 mm long; idioblasts present (absent in A. wilesianum). 26. Rachises with stiff, brown hairs 0.1 mm long on abaxial surfaces only; blade surfaces glabrous; idioblasts absent. .......................................................................................................................................................................................... 34. A. wilesianum.

26. Rachises with dense, narrow, pectinate scales 2–3 mm long; blade surfaces glabrous or with scattered hair-like pectinate scales; idioblasts present on blades adaxially. 27. Pinnules equilateral, 2.5–4.5(–8) ⫻ 1.5–2(–3) cm; terminal pinna on each blade larger than the few (1–3) lateral ones; lateral pinnae with to 6 pairs of pinnules; Chis. .......................................................................... 7. A. caryotideum. 27. Pinnules inequilateral, dimidiate, to ca. 3(–5) ⫻ 1(–3) cm; terminal pinna on each blade about the same size as the lateral ones; lateral pinnae with usually 10 or more pairs of pinnules; s. and w. Mexico. 28. Pinnules abaxially bearing golden hairs (unbranched and without flared bases) on veins; distal pinnules on each pinna greatly reduced; blades thin-textured. .............................................................................. 28. A. terminatum. 28. Pinnules abaxially glabrous or with narrow brownish scales, these branched (pectinate) at their bases; distal pinnules on each pinna reduced or not; blades thin to often thick-textured. 29. Rhizomes long-creeping, usually ca. 2 mm diam.; blades glabrous abaxially; largest pinnules 25–55 mm long. ................................................................................................................................................... 14. A. latifolium. 29. Rhizomes short-creeping, often ⬎4 mm diam.; blades glabrous abaxially, or with scattered to dense, entire to toothed or branched scales; largest pinnules usually ⬎25 mm long. 30. Stipes and rachises with dense, stiff, spreading, entire, subulate scales. ................................... 9. A. decoratum. 30. Stipes and rachises with sparse to moderately dense, twisted, subappressed or slightly spreading, entire to denticulate scales. 31. Largest pinnules of each pinna 5–20 mm long; midveins indistinct, near basiscopic pinnule margins; basal pinnules cuneate with equal sides; abaxial blade surfaces with scattered, hair-like scales; terminal pinnules pinnatifid, elongate, non-conform. 32. Sori usually one per pinnule, 5–10 mm long; pinnule apices acute, directed toward pinna apices; indusia glabrous. ................................................................................. 23. A. pulverulentum. 32. Sori two to many per pinnule, 1–4 mm long; indusia glabrous or with hairs. 33. Indusia glabrous; pinnules rounded at apices. .............................................. 2. A. amblyopteridium. 33. Indusia with hairs 0.5–0.8 mm long; pinnules acute to rounded at apices, sterile ones directed toward pinna apices. .................................................................. 31. A. trichochlaenum. 31. Largest pinnules of each pinna 20–40 mm long; midveins distinct to indistinct, running diagonally near middle of pinnules; basal pinnules rectangular, longer than wide; abaxial blade surfaces glabrous or with scattered, hair-like scales; terminal pinnules conform or non-conform.

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ADIANTUM 34. Terminal pinnules pinnatifid, non-conform; lateral pinnules often strongly falcate at tips, strongly serrate or biserrate. ............................................................................... 29. A. tetraphyllum. 34. Terminal pinnules conform; lateral pinnules usually not falcate at tips, serrulate (except sometimes A. villosum). 35. Pinnules glabrous or with scattered golden hairs abaxially, rounded at tips. ... 17. A. mcvaughii. 35. Pinnules with scattered scales or hair-like scales abaxially, truncate or rounded at tips. 36. Sides of pinnules forming acute angle at pinnule bases; rhizome scales golden brown and with occasional teeth; pinnules rounded at apices, reduced at pinna bases. ....................................................................................................... 18. A. oaxacanum. 36. Sides of pinnules forming right angle at pinnule bases; rhizome scales light brown with thickened, dark brown cell walls, with many recurved marginal teeth; pinnules acute at apices, not much reduced at pinna bases. .............. 33. A. villosum.

1. ADIANTUM ALEUTICUM

Specimens Examined. Chih (Correll & Gentry 23215, LL, MO, UC, US). Sin? (see comment below).

Adiantum aleuticum (Rupr.) C. A. Paris, Rhodora 93: 112. 1991. Figs. 10G, H, J.

This species is subtly different from A. pedatum L., from eastern United States and southeastern Canada. Adiantum aleuticum differs by ultimate divisions (pinnulets) with length usually more than 3.2 times the width (vs. less than 3.2 times the width in A. pedatum), apices with sharply denticulate, angular lobes (vs. apices with rounded, crenulate lobes), and lobes separated by deep sinuses 0.6–4 mm (vs. 0.1–2 mm). Traditionally, the western North America A. aleuticum has been treated as an infraspecific variant of A. pedatum, but recent biosystematic and electrophoretic work has pointed out the differences among this and several closely related north temperate species (Paris & Windham, 1988). See A. patens, the most similar Mexican species, for differences between that and A. aleuticum. Vega A. et al. (1989) reported A. pedatum for Sinaloa; we have not seen the specimen on which this is based, and none was cited, but it is possible that this refers to A. aleuticum, which would be a range extension for Mexico.

Adiantum pedatum L. var. aleuticum Rupr., Distr. Crypt. Vasc. Ross. 49. 1845. Adiantum pedatum L. subsp. aleuticum (Rupr.) Calder & Roy Taylor, Canad. J. Bot. 43: 1388. 1965. Type. U.S.A. Alaska: Aleutian Islands, Unalaska, Chamisso s.n. (HAL?, LE?).

Rhizomes short-creeping or suberect, to 5 mm diam.; rhizome scales castaneous, concolorous, 3–5 ⫻ 0.5–1 mm, margins entire; fronds densely clustered, mostly 30–60 cm long, laxly arching or pendent; stipes castaneous, lustrous, to 30 ⫻ 1–2 mm, ca. 1⁄2 the frond length, glabrous; blades fan-shaped or appearing palmate, actually imparipedate, mostly ca. 15–25 ⫻ 15–25 cm; rachises glabrous, sometimes glaucous; pinnae 2 lateral pairs plus a 1-pinnate terminal pinna, lateral pinnae repeatedly and unequally branching basiscopically, forked 2–4 times, to ca. 20 cm long; pinnulets oblong, tapered at tips, occasionally flabellate, basiscopic margins straight to oblique, generally ca. 2.5–4 times as long as broad, stalks to 3 mm, dark color passing into pinnulet bases or not, non-articulate; veins free, forking, ending in minute teeth in sterile pinnulets; indument absent on both surfaces; idioblasts absent; sori 6–10 per pinnulet, confined to acroscopic margins; indusia 2–3(–6) mm long, oblong to lunate, glabrous; 2n⫽58 (Canada, USA).

Distribution. In spray in recessed area beneath waterfall; 2100 m. Canada, USA; Mexico.

2. ADIANTUM AMBLYOPTERIDIUM Adiantum amblyopteridium Mickel & Beitel, Mem. New York Bot. Gard. 46: 20. 1988. Type. Mexico. Oaxaca: Distrito Pochutla, 15 km N of Pochutla, Mickel 5140 (NY!; isotypes MEXU!, UC!). Figs. 8K–M. Rhizomes short-creeping, to 5 mm diam.; rhizome scales somewhat bicolorous, light brown with cells toward middle having thickened, dark brown cell walls, 2–3 ⫻ 0.3–0.5 mm, margins entire; fronds clumped, 60–100 cm long, arching; stipes atropurpureous, 30–60 cm ⫻ 2–3 mm, 1⁄3–2⁄3 the frond length, with pectinate scales but these abraded and stipes then subglabrous; blades deltate, 2-pinnate, each blade with a conform terminal pinna, 30–50 cm wide; rachises with persistent, tan, lanceolate scales 1–1.5 mm long, these ciliate or denticulate at bases; pinnae 6–9 pairs, 20–35 ⫻ 3–4 cm, terminal pinnules non-conform, pinnatifid, elongate, basal pinnules triangular or flabellate; pinnules (largest) 12–20 ⫻ 8–10 mm, rounded or obtuse at apices, non-articulate; veins free, forking, ending in minute teeth in sterile pinnules or portions of pinnules; indument adaxially ab-

ADIANTUM ANDICOLA

sent, abaxially the laminar surfaces with scattered, tan, pectinate scales 0.8 mm long; idioblasts present and conspicuous on both surfaces; sori ca. 6–10 per pinnule, on acroscopic and distal margins; indusia 2–3 mm long, oblong to lunate, glabrous.

Distribution. Wooded stream banks; 50–600 m. Mexico. Specimens Examined. Oax (Conzatti et al. s.n., MEXU, 3147, US; Mickel 5163, NY–3 sheets). QR (Palacios-Rios 3661, XAL).

Adiantum amblyopteridium is distinguishable from the other bipinnate, idioblast-bearing species with small pinnules (A. pulverulentum, A. trichochlaenum) by the combination of several sori per pinnule, glabrous indusia, and broadly rounded pinnule tips. Adiantum amblyopteridium may be one parent of A. oaxacanum (which see). These two taxa and the other presumed parent, A. obliquum, have been collected near each other in Dto. Pochutla, Oaxaca.

3. ADIANTUM AMPLUM Adiantum amplum C. Presl, Reliq. Haenk. 1(1): 63. 1825. Type. Mexico [“Mexico et in Guayaquil”]. Haenke s.n. (NY! frag. ex PR). Figs. 7A–C. Adiantum princeps T. Moore, Gard. Chron. II, 4: 197, f. 43–44. 1875. Type. Described from cultivated material originally received from Colombia, Hort Veitch, 1875, no. 16, Thomas Moore’s Fern Herbarium (K!; isotype K!).

Rhizomes short-creeping, stout, 3–5 mm diam.; rhizome scales concolorous, bright golden brown, 6–10 ⫻ 0.8–1 mm, margins ciliate-denticulate; fronds clumped, 45–80(–130) cm long, arching; stipes atropurpureous, lustrous, 20–70 ⫻ 3–5 mm, 1⁄3–1⁄2 the frond length, glabrous except at bases, or with minute tubercles or raised areas; blades broadly ovate, 3–4-pinnate proximally; rachises atropurpureous, glabrous; pinnae 4–12 compound pairs, 15–35 ⫻ 10–20 cm, alternate, long-stalked; pinnulets mostly rhombic or obovate-flabellate, 7–15(–20) ⫻ 1–2.5 mm, bases broadly cuneate, sterile margins incised-lobate and denticulate, stalked 2–6 mm, color of stalks stopping abruptly at bases of pinnulets, articulate; veins free, forking, ending in minute teeth; indument absent on both sides of blades; idioblasts absent; sori to ca. 10 per pinnulet, confined to acroscopic and distal margins,

25

with a little yellowish farina among sporangia; indusia 1–3 mm long, oblong to lunate, glabrous.

Distribution. Terrestrial in moist forests, often along streams, roadbanks and woods, tropical deciduous forests, mostly in Pacific lowlands; 50–1500 m. Mexico; Guat, Hond, Salv, Nic, CR, Pan; Col, Ven. Selected Specimens Examined. Chis (Breedlove 23439, DS, NY). Col (Gonza´lez 16919, GUADA, UC). Gro (Palmer 518, NY, UC). Jal (Lott et al. 1012, MEXU, NY, UC). Mich (Hinton 15956, DS, ENCB, NY, P, UC, US). Nay (McVaugh 19220, IEB, MEXU). Oax (Mickel 5981, NY, UC). Sin (Breedlove 35623, CAS). Tab (Cowan 3163, NY). Ver (Tryon & Tryon 357, NY).

The rhomboid pinnulets and long, concolorous rhizome scales distinguish this taxon from the somewhat similar articulate species, A. tenerum. Adiantum amplum and A. tenerum seldom, if ever, grow together, the former occurring primarily on Pacific slopes, the latter on Atlantic slopes.

4. ADIANTUM ANDICOLA Adiantum andicola Liebm., Mexic. Bregn. 266 (reprint 114). 1849. Type. Mexico. Oaxaca: Cerro de Sempoaltepec, Liebmann s.n. [Pl. Mex. 2203, Fl. Mex. 434] (C!; isotypes NY! frag. ex US, US!). Figs. 7G, H, J, O–W. Adiantum cuneatum Langsd. & Fisch. var. angustifolium M. Martens & Galeotti, Me´m. Foug. Mexique 70. 1842. Lectotype (chosen by Morton, Contr. U.S. Natl. Herb. 38: 216. 1973). Mexico. Veracruz: “Pic d’Orizaba,” Galeotti 6266 (BR! photo US; isotype K!, photo US). Adiantum tenerum Sw. var. dissectum M. Martens & Galeotti, Me´m. Foug. Mexique 71. 1842. Type. Mexico. Oaxaca: Galeotti 6361 (BR!, P! photo ex BR, US! photo ex BR). Adiantum amabile Liebm., Mexic. Bregn. 265 (reprint 113). 1849, non A. amabile T. Moore. Lectotype (chosen by Smith, 1981: 19). Mexico. Oaxaca: “inter Tonaguia et Roayaga,” Liebmann s.n. [Pl. Mex. 2206, Fl. Mex. 826] (C!; isolectotype C!, Fl. Mex. 430). Adiantum glaucophyllum Hook., Sp. Fil. 2: 40. 1851. Lectotype (chosen by Lellinger, Proc. Biol. Soc. Wash. 89: 704. 1977). Panama. Veraguas, Seemann s.n. (K!). For additional synonymy, see Lellinger (1989: 145).

Rhizomes compact to long-creeping, 2–5 mm diam.; rhizome scales usually black, or castaneous only at rhizome tips, appressed, 1–1.5 ⫻ 0.1–0.3 mm, margins entire; fronds clumped to often

26

ADIANTUM

well spaced, (23–)35–70 cm long, arching; stipes atropurpureous, lustrous, 10–40 cm ⫻ 1–3 mm, 1⁄2–2⁄3 the frond length, glabrous except at bases; blades broadly ovate to deltate, 3–5-pinnate proximally, 15–50 ⫻ 15–60 cm, tapering and less divided towards apices; rachises atropurpureous, glabrous, sometimes glaucous; pinnae 4–8 compound pairs, alternate, lowermost stalked 2–7 cm; pinnulets obovate to rhombic or flabellate, narrowly (to broadly) cuneate at bases, tiny (4 ⫻ 4 mm) to large (22 ⫻ 25 mm), distal margins incised-lobate, often articulate; veins free, forking, ending in teeth (when teeth are present); indument absent on both sides of blades, abaxially often glaucous and with or without a dense covering of minute papillae; idioblasts absent; sori 2–8 per pinnulet, confined to distal margins; indusia 1–2 mm long, reniform, round-reniform, or lunate, glabrous.

(not found in other species with articulate pinnulets). Maxon and Morton’s unpublished A. hintonii (represented by King & Soderstrom 5043, NY; Rzedowski 45333, IEB; both from Michoaca´n), with larger, broader pinnulets, is one of the more distinct variants; it is most common in western and central Mexico but extends to Panama. Other variants have more deeply incised pinnulets. Breedlove 42784 (DS), from Chiapas, also has more flabellate pinnulets and wider indusial sinuses, and Balls B4703 (UC), from Veracruz, has more lunate sori that lack pronounced sinuses, reminiscent of sori in A. braunii. Another extreme, with numerous, small, narrow pinnulets, and large laminar papillae (longer than wide), appears to be most common in Oaxaca, Veracruz, Chiapas, and Guerrero, but other small-pinnuleted specimens from these states and elsewhere in Mexico lack papillae. Specimens from Hidalgo and Tamaulipas are atypical in having more rounded, less incised pinnulets; identifications of several of these (e.g., Herna´ndez Magan˜a 6092, MEXU; Hinton 24893, TEX) are problematic.

5. ADIANTUM BRAUNII Adiantum braunii Mett. ex Kuhn, Linnaea 36: 75. 1869. Type. Mexico. In 1830, Karwinski s.n. (B!; isotypes BR!, P! photo ex BR, US! photo ex BR). Figs. 5A–C.

Distribution. Terrestrial in moist, oak or pine-oak forests, alder forests, at middle to high elevations, in dry woods, montane rain forests, evergreen cloud forests; 1000–3150 m. Mexico; Guat, Hond, Salv, Nic, CR, Pan; Col. Selected Specimens Examined. Chih (Jenkins et al. s.n., 27 July 1988, MO). Chis (Breedlove 32455, DS, NY). DF (Purpus 1584, NY, UC, US). Dgo (Palmer 176, K, NY, US). Gro (Cowan 4948, CAS, NY, TEX, UC). Gto (Dı´az Barriga 3054, ENCB, IEB). Hgo (Gimate 753, ENCB, NY). Jal (McVaugh 26137, NY). Me´x (Pringle 15638, CAS, UC, US). Mich (Hitchcock & Stanford 7142, DS, NY). Mor (Copeland herb. 113, MEXU, NY, UC). Nay (Boutin & Brandt 2646, CAS). NL (Meyer & Rogers 3057, MO). Oax (Mickel 6150, ENCB, NY). Pue (Orcutt 4025, DS, US). Qro (Rubio 2117, MEXU). SLP (Rzedowski 27710, IEB). Sin (Vega et al. 3014, ENCB, FCME). Tam (Hinton 24893, TEX). Tlax (Chimal et al. 28, IEB). Ver (Garcı´a & Delgado 950, CHAPA, MO, NY, XAL). Unverified, Doubtful, or Mistaken Reports. Ags (De la Cerda & Garcı´a 1350, HUAA, cited by Siqueiros-Delgado & Gonza´lez-Adame, 2004, but not verified). Tab (reported by Magan˜a, 1992, probably based on a misidentification).

Adiantum andicola is most likely a complex comprising several species and is in need of monographic study. The papillate nature of the surface of pinnulets in some variants has been overlooked, and is most evident on specimens from Oaxaca and Guerrero. There is considerable variation in this species in pinnulet size, shape, blade dissection, glaucousness, and rhizome form (compact to long-creeping), but some correlation exists between pinnulet size and papilla size. The variants lumped together here have in common dark, usually black, rhizome scales

Adiantum braunii Mett. ex Kuhn var. cuneatum Mett. ex Kuhn, Linnaea 36: 75. 1869 [“cuneata”]. Type. Mexico. Berlandier 1151 (B, not seen). Adiantum convolutum E. Fourn., Mexic. Pl. 1: 127, t. 6. 1872. Syntypes. Mexico. Veracruz: Orizaba, Mpio. Orizaba, environs d’Orizaba, Botteri & Sumichrast 18 (P!–2 sheets; isosyntype BR!); Ghiesbreght 406 (P!–2 sheets; isosyntype US!).

Rhizomes short-creeping, 4–6 mm diam.; rhizome scales dark castaneous at rhizome apices to mostly black back from the tips, lustrous, 1.5–3 ⫻ 0.5 mm, margins entire; fronds 25–80 cm long, clumped, arching; stipes atropurpureous to blackish, lustrous, 20–40 cm ⫻ 1–2.5 mm, 1⁄2–2⁄3 the frond length, glabrous except for few broad scales at bases; blades broadly ovate, 3–4-pinnate proximally; rachises atropurpureous, glabrous; pinnae 5–8 compound pairs, alternate, stalked to 4 cm, axes with a pale spot at each branching point along rachises; pinnulets rhombicflabellate, orbicular, or obovate, mostly broadly cuneate (120– 180⬚, rarely less than 90⬚) at bases, 8–12 mm long, margins incised-lobate, stalked 1–10 mm, stalk color passing into pinnulet bases, non-articulate; veins free, forking, ending at the entire, cartilaginous margins or ending in very small teeth; indument absent on both sides of blades, blades not glaucous; idioblasts absent; sori 1–4 per pinnulet, confined to distal margins; indusia 1–8 mm long, linear-oblong to lunate, glabrous; 2n⫽60 (Jal).

ADIANTUM CAPILLUS-VENERIS

27

ca. 1⁄2 the frond length, glabrous except at bases; blades ovate to ovate-lanceolate, 2–3 times pinnate, ca. 5–22 cm wide, tapering to apices; rachises castaneous, glabrous; pinnae 3–7 compound pairs, alternate, stalked; pinnulets obovate, flabellate, or rhombic, 5–30 mm long, cuneate at bases, margins often deeply lobed or incised distally, 2 or more clefts per pinnulet, sterile margins strongly denticulate, color passing into base of pinnulets, nonarticulate; veins free, forking, ending in teeth; indument absent on both sides of blades, laminae not glaucous; idioblasts absent; sori 2–6 per pinnulet, confined to distal margins; indusia 1–5 mm long; oblong to arcuate, glabrous; 2n⫽60 (Old World), 120 (USA). Distribution. Terrestrial in oak and pine woods, montane forests, deciduous forests, subtropical forests, tropical deciduous forests, and on lava flows; (150–)350–2200 m. Mexico; Guat, Salv, Nic, CR; Col, Ven, Ec. Selected Specimens Examined. Chis (Davidse et al. 29737, MO, UC). Col (Zu´n˜iga 18302, UC). Dgo (Ortega 4299, US). Gro (Hinton 9766, MO, NY, US). Hgo (Riba 4, NY). Jal (Pringle 1868, B, BR, CAS, DS, K, MO, NY, P, UC, US). Me´x (Mickel 683, NY). Mich (Pringle 3385, UC). Mor (Lyonnet 2564, CAS, MO, UC, US). Nay (Mexia 536, CAS, UC, US). Oax (Mickel 6230, ENCB, NY, UC). Qro (Ferna´ndez & Acosta 2149, ENCB, IEB). Sin (Mexia 427, B, CAS, MO, UC, US). Son (Gentry et al. 19325, NY, US). Ver (Purpus 6185, MO, NY, UC). Unverified, Doubtful, or Mistaken Reports. Ags (Garcı´a s.n., HUAA, cited by Siqueiros-Delgado & Gonza´lez-Adame, 2004, but not verified).

This is a very common species on the western slopes of the Sierra Madre Occidental, at moderately low to middle elevations, but uncommon or absent elsewhere in Mexico. Among the 2–5-pinnate species with tapering blade apices, this species is distinguished by having non-articulate stalks with the color passing onto the pinnulet bases, and raised whitish patches at the bases of the costae. The specimen cited from Hidalgo is atypical in the small fronds and pinnulets and the lack of whitish patches at the bases of the pinnae.

6. ADIANTUM CAPILLUS-VENERIS Adiantum capillus-veneris L., Sp. Pl. 2: 1096. 1753. Lectotype (chosen by Pichi Sermolli, Webbia 12: 678, 1957). France. Magnol s.n. (LINN 1252.9, microfiche UC). Figs. 5M–O. Adiantum schaffneri E. Fourn., Bull. Soc. Bot. France 27: 328. 1880. Type. Mexico. San Luis Potosı´: “in montibus circa San Luis de Potosı´,” Schaffner 64 (P!, frag. NY!, photos BM!, BR, US!). Adiantum capillus-veneris L. var. protrusum Fernald, Rhodora 52: 203. 1950. Type. U.S.A. Georgia: Clay Co., Samochechobee Creek, Harper 1791 (GH). Adiantum modestum Underw., Bull. Torrey Bot. Club 28: 46. 1901. Adiantum capillus-veneris L. var. modestum (Underw.) Fernald, Rhodora 52: 206. 1901. Type. U.S.A. New Mexico: North Spring River, Roswell, Earle 261 (GH; isotype GH).

Rhizomes short-creeping, 2–4 mm diam.; rhizome scales castaneous, concolorous, ca. 2–2.5 ⫻ 0.1–0.3 mm, margins entire; fronds clumped, 15–45(–70) cm long, arching or pendent; stipes castaneous to atropurpureous, lustrous, 7–35 cm ⫻ 0.5–1.5 mm,

Distribution. Terrestrial on wooded stream banks or epipetric on limestone in seepage areas; ravines with Taxodium and Ficus, tropical deciduous forests, matorral, lower montane rain forests, evergreen rain forests; 150–2450 m. USA; Mexico; Guat, Bel, Hond; Berm, Bah, Gr & L Ant; Ven, Trin, Peru; Eur, Africa, Asia, Pac Is. Selected Specimens Examined. Ags (Siqueiros D. 4295, IEB). BCN (Wiggins 10030, DS, UC). BCS (Carter & Sharsmith 4240, NY, UC, US. Cedros (Anthony 114, DS, MO, NY, UC, US). Chih (Johnston et al. 11375, CAS). Chis (Matuda 2423, MEXU, NY, US). Coah (Palmer 71, MO, UC, US). DF (Schaffner 120, in 1875, NY). Dgo (Palmer 77, MO, UC). Gro (Lorea 1141, ENCB, FCME). Gto (McVaugh 14834, NY). Hgo (Ortega 86, IEB). Jal (Jones 497b, US). Me´x (Ventura 422, ENCB, NY). Mich (Hinton 15895, NY, UC). Mor (Hinton 17079, ENCB, NY). Nay (Te´llez 9164, IEB p.p.). NL (Hinton 17508, ENCB, MEXU). Oax (Mickel 6301, NY, UC). Pue (Orcutt 2098, DS). Qro (Ferna´ndez N. 3773, ENCB, IEB). SLP (Copeland herb. 112, UC, US). Son (Hartman 292, UC, US). Tam (Palmer 189, MO, NY, UC, US). Tlax (Langman 2584, US). Ver (Va´zquez et al. 379, NY). Zac (Lloyd 13, UC, US). Unverified, Doubtful, or Mistaken Reports. Sin (reported by Vega A. et al., 1989, but not verified).

Of the Adiantum species with blades gradually tapering distally, only a few have the veins ending in teeth. Adiantum capillus-veneris is the most conspicuous in this respect, with its strongly denticulate margins. Adiantum andicola is sparingly and shortly toothed and differs in its blackish rhizome scales. There is significant variation within A. capillus-veneris, even within Mexico, in such characters as depth of lobing or incision of the pinnulets (both fertile and sterile), size of segments, length-to-width ratio of pinnulets, size and degree of dissection of blades, and extent of creeping of the rhizomes. Fernald (1950)

28

ADIANTUM

recognized two intergrading varieties of this widespread species in the United States and Mexico, while restricting the type variety to Europe. Variety modestum was said by him to differ chiefly in its firmer texture, small, barely notched fertile pinnulets and Cordilleran distribution in the United States. We are unable to apply the characters discussed by Fernald to distinguish varieties in the New World, or even to distinguish the European plants from those of the United States and Mexico. In general, sterile and/or juvenile plants or fronds, in many parts of the species range, often have more deeply cleft pinnulets than fertile plants or fronds.

7. ADIANTUM CARYOTIDEUM Adiantum caryotideum Christ, Bull. Soc. Bot. Gene`ve 1: 230. 1909. Type. Costa Rica, 1903, Werckle´ s.n. (P!). Figs. 11D–G. Rhizomes short-creeping, 5–10 mm diam.; rhizome scales dark brown to blackish, clathrate, scales ca. 2 ⫻ 0.3–0.5 mm, margins denticulate; fronds clumped, 50–75 cm long, arching; stipes atropurpureous to blackish, lustrous, ca. 35 cm ⫻ 3 mm, 1⁄2 the frond length, with appressed scattered scales, glabrescent; blades deltate, 2-pinnate, ca. 20–25 ⫻ 7–15 cm, broadest at bases, terminal pinna of each blade larger than lateral pinnae and with about 9 pairs of pinnules; rachises atropurpureous, with scattered, spreading, linear scales 1–5 mm long, scales often caducous; pinnae 2–3 lateral pairs, to ca. 15 ⫻ 7 cm, with ca. 6–7 pairs of pinnules; pinnules ovate to ovate-lanceolate, 2.5–4.5(–8) ⫻ 1.5–2(–3) cm, sterile margins serrate to biserrate, distal pinnules of each pinna not greatly reduced, terminal pinnules hastate, to 6 ⫻ 2 cm, stalked 2–5 mm, stalk color stopping rather abruptly at bases of pinnulets, non-articulate; veins free, forking, ending in marginal serrations; indument absent on both sides of blades, not glaucous; idioblasts present and conspicuous on both blade surfaces; sori 5–10 per pinnule, along acroscopic and distal part of basiscopic margins; indusia mostly 3–5 mm long, linear-oblong, glabrous.

Distribution. Terrestrial in lowland rain forests; 200–300 m. Mexico; Guat, CR, Pan.

Specimens Examined. Chis (Breedlove 34205, DS, MO, NY; Gonza´lez-Espinosa et al. 466, XAL; Sohns 1615a, US).

This is a very distinct but rare species in Mexico and appears to be related to A. seemannii. It is amply distinct from all other Mexican species by the bipinnate fronds with relatively few pinnae, small number of pinnules per pinna, large terminal pinna that exceeds the lateral ones in length and width, and the large pinnules, exceeded only by those of A. seemannii in Mexico.

8. ADIANTUM CONCINNUM Adiantum concinnum Humb. & Bonpl. ex Willd., Sp. Pl., ed. 4, 5(1): 451. 1810. Lectotype (chosen by R. Tryon, Contr. Gray Herb. 194: 168. 1964). Venezuela. Caracas [“Cumana”], Humboldt & Bonpland s.n. (B-Willd. 20099-2!, microfiche UC!; frag. NY! ex B-Willd.). Figs. 5K, L. Adiantum lutescens Mougeot ex Fe´e, Me´m. Foug. 5: 119. 1852. Type. Mexico. Oaxaca: s. coll. s.n. (Herb. Mougeot, BR?, MPU?).

Rhizomes short-creeping to suberect, compact, ca. 5 mm diam.; rhizome scales castaneous, concolorous, 3–4 ⫻ 0.8–1 mm, margins entire; fronds clumped, 20–75 (–95) cm long, arching to pendent; stipes castaneous to atropurpureous, lustrous, to 20 cm ⫻ 2(–3) mm, 1⁄4–1⁄3 the frond length, glabrous except at the bases; blades lanceolate to ovate-lanceolate, 2–3-pinnate proximally, 6–30 cm wide, long-attenuate at apices; rachises castaneous to atropurpureous, glabrous; pinnae 10–15(–18) compound pairs, gradually reduced to apices, alternate, basal acroscopic pinnule of each pinna often overlapping the rachises; pinnulets obovate to flabellate, cuneate at bases, sterile pinnulets entire to obscurely denticulate, 7–10 mm long, stalked 1–5 mm, lobed or incised distally, color passing into the pinnulet bases, non-articulate; veins free, forking, ending at entire thickened pinnulet margins or shallow sinuses; indument absent on both sides of blades; idioblasts absent; sori 4–8 per pinnulet, confined to distal margins; indusia 1–1.5 mm wide, reniform to round-reniform, glabrous; 2n⫽114 (Chis), 116 (Nay, Jam), 120 (PR).

Distribution. Terrestrial on moist banks in lightly shaded woods; 50–2300 m. Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Ec, Peru, Braz.

ADIANTUM DEFLECTENS

29

Selected Specimens Examined. BCS (Brandegee 664, NY, UC, US). Chis (Breedlove 21551, DS, MO, NY). Col (Palmer 1127, MO, NY, UC, US). DF (Rzedowski 2055, ENCB). Dgo (Orcutt 3455, DS). Gro (Mexia 8724, NY, UC). Jal (Pringle 1866, DS, MO, UC, US). Me´x (Matuda 27188, UC, US). Mich (Rzedowski 40741, ENCB, IEB). Mor (Lyonnet 2874, CAS, FCME, IEB, MO, UC). Nay (Mexia 534, MO, UC). Oax (Smith 2089, MO, UC, US). Pue (Turra 1979, ENCB). Qro (Rzedowski 42931, ENCB, IEB). SLP (Pringle 3394, UC). Sin (Breedlove 18049, CAS, NY). Son (Gentry 764, DS). Tam (Kemp s.n., Jan 1902, NY). Ver (Purpus 1964, UC). Unverified, Doubtful, or Mistaken Reports. Ags (reported by de la Cerda Lemus & Siqueiros D., 1985, but not verified).

This species is widespread and common nearly throughout Mexico, with over 500 collections seen. Among the 2–3-pinnate taxa that lack a conform terminal pinna, the short-stalked pinnules with stalk color passing into the pinnulets, veins ending at entire margins or shallow sinuses, and basal acroscopic pinnules overlapping the rachises distinguish this species.

9. ADIANTUM DECORATUM Adiantum decoratum Maxon & Weath., Amer. J. Bot. 19: 165. 1932. Type. Panama. Panama´: Rı´o Tapia, Standley 26216 (US!). Figs. 12H, J, K. Rhizomes short-creeping, nodose, 4–7 mm diam.; rhizome scales concolorous, dark brown, 3–4 ⫻ 0.3–0.5 mm, margins denticulate; fronds clumped, 50–90 cm long, arching; stipes atropurpureous, 25–50 cm ⫻ 3–5 mm diam., with dense filiformsubulate spreading scales, these persistent, giving the stipes a shaggy appearance; blades 2-pinnate (rarely 3-pinnate proximally), each with a conform terminal pinna; rachises atropurpureous, with dense spreading dark brown scales similar to those of stipes, scales entire or denticulate, especially toward their bases; pinnae 5–7 pairs, each pinna ending in a small lanceolate pinnule, distal pinnules on each pinna reduced to less than half the size of largest pinnules; pinnules oblong, truncate or rounded at tips, 10–20 ⫻ 4–7 mm, the sterile serrulate along distal and acroscopic margins, oblique to spreading, not or only slightly falcate, short-stalked, non-articulate; veins free, forking, ending in marginal serrations; indument absent on both sides of blades, often glaucous abaxially; idioblasts present and conspicuous on both blade surfaces; sori 4–9 per pinnule, along acroscopic and distal margins of pinnules; indusia 1–3 mm wide, oblong to linear, glabrous.

Distribution. Terrestrial in lowland rain forests and lower montane forests; 100–350 m. Mexico; Guat, Bel, Hond, CR, Pan; nw Col. Selected Specimens Examined. Chis (Breedlove 33179, DS, F, LL, MEXU, 34171, DS; Castillo C. et al. 3718, 9503, XAL; Martı´nez S. 7786, UAMIZ, 7999, 8405, MEXU, NY, 8448 MEXU, NY, UAMIZ; Sohns 1701, US). QR (Uca´n & Flores 935, XAL, UADY, UC). Tab (Matuda 3328, F, LL, US). Unverified, Doubtful, or Mistaken Reports. Ver (Ventura s.n., ENCB, cited by Palacios-Rios, 1992, but not found and likely misidentified).

This differs from other bipinnate species with idioblasts and conform terminal pinnae by the dense and spreading hair-like scales on the stipes and rachises. Several of the collections from Chiapas have the proximal pinnae (to three pairs) fully bipinnate at the bases, with three pairs of pinnate pinnules on both acroscopic and basiscopic sides. Of Mexican species, A. decoratum appears most closely related to A. tetraphyllum.

10. ADIANTUM DEFLECTENS Adiantum deflectens Mart., Icon. Pl. Crypt. 94. 1834. Type. Brazil. Para´: Santare´m, Martius s.n. (M). Figs. 5D, E. For additional synonymy, see Lellinger (1989: 147).

Rhizomes short-creeping to erect, 2–4 mm diam.; rhizome scales bicolorous, center dark brown, margins light brown, 2 ⫻ 0.1–0.3 mm, margins entire; fronds clumped, (8–)12–30 cm long, arching; stipes castaneous to atropurpureous, dull to sublustrous, to 15 cm ⫻ 0.5–0.8 mm, 1⁄4–1⁄2 the frond length, glabrous; blades lanceolate, 1-pinnate, tapering to apices, 2–5 cm wide; rachises castaneous, glabrous, outside Mexico occasionally flagelliform and rooting at tips; pinnae 3–10 pairs, alternate, stalked 3–8 mm, stalk color stopping Ⳳ abruptly at pinna bases, articulate; ultimate divisions (pinnae) cuneate-flabellate, asymmetrical to symmetrical, 12–14 mm long, blades usually with a Ⳳ conform terminal pinna, pinnae cleft or lobed to 3 mm deep, 2–7 lobes per pinna, sterile margins sharply serrulate; veins free, forking; ending in sharp and conspicuous teeth on sterile pinnae; indument absent on both sides of blades; idioblasts absent; sori 3–7 per pinna, confined to distal margins; indusia 1–5 mm long, oblong to slightly arcuate, glabrous.

30

ADIANTUM

ially on costae, costules, and pinnulet stalks of dense, stiff, reddish hairs 0.1–0.3 mm long, abaxially the pinnulets glandularhirsute with stiff to flexuous pale hairs 0.2–0.5 mm long on the veins; idioblasts absent; sori 2–6 per pinnulet, confined to distal margins; indusia 1–3 mm long, oblong to often lunate or reniform, glabrous.

Distribution. Terrestrial in lowland moist woods near streams, tropical deciduous forests; 150–780 m. Mexico; Guat, Hond, Salv, Nic, CR, Pan; Col, Ven, Fr Gui, Ec, Peru, Braz, Bol, Parag. Specimens Examined. Chis (Breedlove 27288, DS). Gro (Lorea 4773, FCME). Oax (Gallardo H. 1900, MEXU; Mickel 5974, NY, UC).

This is often confused with A. lunulatum. Adiantum deflectens is distinct in its articulate pinnae, short pinna stalks, veins ending in teeth, and tan spores. The rather few Mexican specimens have pinnae that are not as sharply serrate as in South American material. Also, the Mexican specimens seen show no evidence of the rachises being prolonged and rooting at the tips, as is common elsewhere in the range of the species. The closest relatives of A. deflectens are a small group of South American species that include A. delicatulum Mart. in Brazil and Paraguay, A. calcareum Gardn. from the same two countries, and A. mendoncae Alston from Mozambique and Zimbabwe!

11. ADIANTUM FEEI Adiantum feei T. Moore ex Fe´e, Me´m. Foug. 7: 29, t. 24, f. 1. 1857. Type. Mexico. Veracruz: “prope` Orizabam,” Schaffner 446 (isotype K!). Figs. 6A, B. Adiantum flexuosum Hook., Sec. Cent. Ferns, t. 61. 1861. Type. Guatemala. Baja Verapaz: Santa Rosa, Salvin s.n. (K!–2 sheets; isotype Salvin 464 US!).

Rhizomes short-creeping, ca. 3–6 mm diam.; rhizome scales dark brown to atropurpureous, blackish with age, 2–4 ⫻ 0.3–0.5 mm, margins entire; fronds clumped, (20–)35–100⫹ cm long, arching or scandent; stipes atropurpureous, lustrous, ca. 1⁄2 the frond length, with stiff, erect, uniform reddish hairs ca. 0.1–0.2 mm long; blades ovate to ovate-lanceolate, 3–4-pinnate proximally, 15–30 cm wide, gradually tapering towards apices; rachises atropurpureous, often flexuous or zig-zag, with dense, stiff, reddish hairs 0.1–0.2 mm long; pinnae 6–8 compound pairs per blade, alternate, stalked 1–2 mm long; pinnulets flabellate to rhombic or orbicular, 7–12 mm long and broad, bases cuneate to rounded, margins incised-lobate, stalk color stopping Ⳳ abruptly at pinnulet bases, articulate; veins free, forking, ending at the thickened cartilaginous pinnulet margins; indument adax-

Distribution. Terrestrial among rocks on shrubby trailsides, pine-oak forests, pine-oak-Liquidambar forests, montane rain forests; 1300–1900 m. Mexico; Guat, Bel, CR, Nic. Selected Specimens Examined. Chis (Breedlove 21160, DS, MO, 23179, DS, NY, 53164, 56188, CAS, ENCB). Oax (Mickel 1608, ISC, UC, US, 4839, ENCB, NY, UC, US). Qro (Carranza 4608, IEB, MEXU, UAMIZ; Gonza´lez P. 690, IEB, MEXU). Tam (Sharp et al. 50103, US). Ver (Palacios-Rios 3507, XAL; Schaffner 446, K).

This species is by far most commonly collected in Chiapas, with nearly 30 collections seen; fewer than half this number of collections are known from elsewhere in Mexico, where the species appears to be rare or very local. The flexuous, densely hairy rachises, costae, costules, and pinnulet stalks distinguish this species from all others in Mexico. It has the peculiar habit, for a maidenhair, of scrambling over low brush.

12. ADIANTUM GALEOTTIANUM Adiantum galeottianum Hook., Sp. Fil. 2: 10, t. 80B. 1851. Type. Mexico. Oaxaca: [Juquila del Sur], Galeotti 6561 (K!; isotypes BR!–4 sheets, P!–3 sheets). Figs. 6G, H. Rhizomes short-creeping, compact, ca. 2–4 mm diam.; rhizome scales golden brown, 2–3 ⫻ 0.1–0.3 mm, margins ciliolate; fronds clumped, 17–35 cm long, arching; stipes castaneous (to atropurpureous), sublustrous, ca. 10 cm ⫻ 1 mm, 1⁄3–1⁄2 the frond length, glabrous or with minute stubble toward the bases; blades usually linear, 1-pinnate, sometimes 2-pinnate (and hence more broadly deltate) with a single proximal pair of divided pinnae; rachises castaneous to atropurpureous, glabrous, sometimes glaucous; pinnae 3–10 pairs, opposite to subopposite, sessile or shortstalked ca. 1–1.5 mm; ultimate divisions (pinnae) suborbicular, subcordate to rounded at bases, apices rounded, sterile margins irregularly crenate or subentire, short-stalked 1–2 mm, stalk color stopping near the pinna bases, non-articulate; veins free, forking,

ADIANTUM LATIFOLIUM

31

ending at the whitish cartilaginous pinna margins; indument absent on both sides of blades; idioblasts absent; sori to ca. 15 per pinna, on all but the proximal margins; indusia ca. 2 mm long, reniform or round-reniform with often deep sinuses, glabrous.

Distribution. Terrestrial on moist ground in shade, on rocky banks, north slopes in chaparral, in damp shady gullies; 50– 500 m. USA (Calif, Ore); Mexico. Distribution. Terrestrial on wet roadside banks in pine woods, pine-oak woods, moist banks of streams; 600–900 m. Mexico. Selected Specimens Examined. Gro (Clark 7201, MO; Gray & Rowell 3004, US; Hinton et al. 11375, ENCB, K, MEXU, MO, NY, US; Te´llez 6168, MEXU; Vela´zquez 661, FCME). Oax (Cedillo T. & Torres 1471, CAS, MEXU; Conzatti 4374, MEXU, US; Hallberg 1765, NY; Mickel 3992, 7004, NY, UC).

Adiantum galeottianum is one of the most distinctive species in the genus with its nearly orbicular pinnae and thickened pinna margins. It is most closely related to A. shepherdii, with which it agrees in the thickened pinna margins; Adiantum shepherdii differs by the much more numerous, dimidiate ultimate divisions (pinnae). Another relative is A. patens, which differs in having imparipedate blade form.

13. ADIANTUM JORDANII Adiantum jordanii Mu¨ll. Hal., Bot. Zeitung (Berlin) 1864: 26. 1864. Type. U.S.A. California: Napa Valley, Jordan s.n. (KR?, STU?). Figs. 10K–M. Rhizomes short-creeping, 2–3 mm diam.; rhizome scales castaneous, 2–3.5 ⫻ 0.4–0.6 mm, margins entire; fronds clumped, 18–35(–45) cm, arching or pendent; stipes castaneous, lustrous, 9–20 ⫻ 1–1.5 mm, glabrous except at bases, not glaucous; blades ovate-lanceolate, 2–3-pinnate proximally, gradually tapered distally, 10–24 ⫻ 8–10 cm; rachises castaneous, glabrous; pinnae 4–6 compound pairs, alternate, proximal pinnae 1–2-pinnate; pinnulets flabellate to orbicular, usually broader than long, bases truncate or broadly cuneate, distal margins rounded, slightly incised, sterile margins serrulate, stalks 1–4 mm, with dark color ending Ⳳ abruptly at pinnulet bases, non-articulate; veins free, forking, ending in marginal teeth; indument absent on both sides of blades; idioblasts absent; sori 2–5 per pinnulet, confined to distal margins; indusia 3–8(–10) mm long, oblong to linear, glabrous; 2n⫽60 (USA).

Selected Specimens Examined. BCN (Bell 1378, CAS, DS, MEXU; Fisher et al. 2200, ARIZ, ENCB; Mason et al. 3451, ARIZ, ENCB; Moran 26389, MEXU, MO; Shreve 6815, MO, US; Wiggins 4225, CAS, DS, US, 5150, DS, US, 7541, DS, NY, UC, US). BCS (Epling & Robison s.n., 27 Mar 1940, DS; Epling & Stewart s.n., 11 Apr 1936, DS).

This species is characterized by flabellate pinnulets with veins ending in teeth, 2–3(–5) sori per pinnulet, and color stopping more or less abruptly at the bases of pinnulets. It appears to be related to A. capillus-veneris and differs from that especially by the last character mentioned. It is a seasonal species, in California appearing with the winter (Nov–Apr) rains, and then drying and seemingly disappearing during the dry, nearly rainless summers. The name Adiantum emarginatum Hook. (Sp. Fil. 2: 39, t. 75A. 1851, non Bory ex Willd., 1810, which has as its type a plant from Mauritius) has often been used for this species, but Hooker did not intend to describe a new taxon. Subsequent authors (e.g., D. C. Eaton, Ferns N. Amer. 1: 285. 1879) adopted A. emarginatum in a new (and illegitimate) sense, applying it to plants from California. Hooker’s original figure probably does represent A. jordanii and was drawn from a collection that had been mislabeled, according to Eaton.

14. ADIANTUM LATIFOLIUM Adiantum latifolium Lam., Encycl. 1: 43. 1783. Neotype (designated by Proctor, 1977: 114). Guadeloupe. Basse Terre, Proctor 20110 (A; isoneotype US!). Fig. 8D. Adiantum lucidum Sw. var. bipinnatum Mett. ex E. Fourn., Mexic. Pl. 1: 129. 1872. Lectotype (chosen by Lellinger, Mem. New York Bot. Gard. 23: 13. 1972). Mexico. Tabasco: Teapa, Linden s.n. (P, photo US!). For additional synonymy, see Lellinger (1989: 148).

Rhizomes long-creeping, ca. 2–3 mm diam.; rhizome scales concolorous, light brown with thick, dark brown cell walls, 2–3 ⫻ 0.5–0.8 mm, margins entire to sparsely denticulate; fronds distant, 30–75 cm long, arching; stipes atropurpureous, 30–50 cm ⫻ 2–3 mm, 1⁄2–2⁄3 the frond length, with pectinate scales to 3 mm, glabrescent; blades deltate, 2-pinnate, 20–32 cm wide, each

32

ADIANTUM

blade with a conform terminal pinna; rachises atropurpureous, with tan pectinate scales mostly 1–2 mm long; pinnae 1–4 pairs, alternate, proximal pinnules reduced, stalk color stopping Ⳳ abruptly at pinnule bases; pinnules denticulate, stalked to 1 mm, largest 20–55 ⫻ 6–12 mm, distal pinnules on each pinna ca. 1⁄2 the length of the longest pinnules, tips acute to rounded, nonarticulate; veins free, forking; ending in small teeth; indument absent on both sides of blades, often glaucous abaxially; idioblasts present on both blade surfaces but often rather faint abaxially; sori to 18 per pinnule, on acroscopic and basiscopic margins; indusia 1–5 mm long, oblong to linear, glabrous; 2n⫽120 (Jam).

Distribution. Terrestrial in lowland and lower montane rain forests, deciduous tropical forests; 50–600 m. Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol, Parag, Arg. Selected Specimens Examined. Chis (Breedlove 34142, DS; Matuda 3678, MEXU, NY, US). Oax (Hallberg 1483, MEXU, NY; Mexia 9182, GH, MICH, NY, UC; Mickel 5774, 6849, NY, UC). Tab (Jime´nez Gonza´lez 17139, GUADA, UC; Ventura A. 20383, IEB). Ver (Copeland herb. 121, MEXU, MICH, NY, UC; Rzedowski 18890, MEXU). Unverified, Doubtful, or Mistaken Reports. Jal (reported by Mickel & Beitel, 1988, but the specimens are all A. mcvaughii). Nay (reported by Mickel & Beitel, 1988, but the specimens are all A. mcvaughii). QR (Torres 715, CIQRO, cited by Palacios-Rios, 2002c, but not verified).

Among the Mexican species with idioblasts on the adaxial epidermis, this bipinnate species is distinguishable by its longcreeping rhizomes and large, acute pinnules. The closest relative in Mexico is probably A. mcvaughii.

15. ADIANTUM LUNULATUM Adiantum lunulatum Burm. f., Fl. Indica 235. 1768. Type. India. Malabar Coast, s. coll. s.n. (G). Fig. 5J. Rhizomes short-creeping, 1–2 mm diam.; rhizome scales brown, often with dark brown to black central streak, lustrous, 3–4 ⫻ 0.3–0.5 mm, margins entire; fronds clumped, 15–45(–60) cm long, arching to pendent; stipes castaneous, lustrous, 8–15 cm ⫻ 0.7–1.5 mm, 1⁄5–2⁄5 the frond length, glabrous except at bases; blades linear, 1-pinnate, often ending in flagelliform tips and

eventually rooting; rachises castaneous to atropurpureous, glabrous; ultimate divisions (pinnae) 10–16 pairs, with subconform terminal pinna, alternate, largest pinnae at blade bases, 2–4 ⫻ 1–2 cm, stalked to 2(–4) cm, gradually reduced at blade tips, basiscopic margins straight to concave, outer margins subentire to incised-lobate, sometimes deeply so, lobes rounded, distal segments broadly cuneate proximally, terminal segments (if any) flabellate, color ending at pinna bases, not usually articulate; veins free, forking, ending at the Ⳳ entire margins; indument absent on both sides of blades; idioblasts absent; sori 4–10 per pinna, confined to acroscopic and distal pinna margins; indusia 2–4(–8) mm long, oblong to linear, glabrous; 2n⫽60, 90, 120 (India), apogamous forms known.

Distribution. Terrestrial, on banks near streams, low elevations on Pacific slopes; pine-oak forests, deciduous and semideciduous tropical rain forests, tropical subdeciduous forests with Brosimum, seasonal evergreen forests; 50–1000 m. Mexico; Guat, Hond, Salv, Nic, CR, Pan; Cuba; Col, Ven; Africa, Madagascar, India, se Asia, Malesia, Polynesia. Selected Specimens Examined. Chis (Purpus 6744, MO, UC, US). Col (Orcutt 4670, DS, MEXU, MO, US). Gro (Mexia 8721, B, MO, UC, US). Jal (Feddema 2580, ENCB, MO). Me´x (Tejero-Dı´ez 2651, IZTA). Mich (Ventura 2477, ENCB, NY). Nay (McVaugh 19197, CAS, IEB, MEXU, MO, NY). Oax (Mickel 5139, 5980, NY, UC). Sin (Mexia 371, CAS, MO, UC).

This species often goes under the name A. philippense L. (Sp. Pl. 2: 1094. 1753), based on a Petiver illustration (Gazophylacii Naturae et Artis decas prima pol. 4, f. 4. 1702), but Verma (Nova Hedwigia 3: 463–468. 1961) argued that this plate is so poor that this name cannot be typified. For a different opinion, see Pichi Sermolli (Webbia 12: 665–669. 1957). Radicant blade apices are also seen in extraterritorial specimens of A. deflectens, which differs in its sharply denticulate pinna margins and shorter pinna stalks.

16. ADIANTUM MACROPHYLLUM Adiantum macrophyllum Sw., Prod. 135. 1788. Type. Jamaica. Swartz s.n. (S). Fig. 8J.

ADIANTUM MCVAUGHII

Rhizomes short-creeping, 4–7 mm diam.; rhizome scales concolorous, linear-lanceolate, brown with dark brown, thickened cell walls, 2–3 ⫻ 0.8–1 mm, margins ciliate-denticulate; fronds clumped, mostly 30–65 cm long, arching; stipes atropurpureous to black, lustrous, mostly 20–45 cm ⫻ 2–4 mm, ca. 1⁄2–2⁄3 the frond length, glabrous; blades oblong; 1-pinnate, 15–25 ⫻ 10–16 cm; rachises atropurpureous to black, glabrous except at bases; pinnae 3–8 pairs, each blade with a conform terminal pinna, fertile pinnae entire, sterile pinnae incised or serrulate, with a costa running down middle of each pinna, ovate-deltate, broadly cuneate proximally, acuminate distally, largest pinnae at blade bases, 5–9 ⫻ 2.5–5 cm, subsessile or with stalks 1–2 mm long, the dark color of axis entering the pinna bases, non-articulate; veins free, forking, ending in teeth; indument absent on both surfaces, blades often glaucous abaxially; idioblasts present on both blade surfaces; sori continuous, one on each side of pinnae, interrupted only by the sterile pinna tips; indusia 35–60 mm long on fully fertile pinnae, glabrous; 2n⫽60 (Jam).

Distribution. Terrestrial in wet lowland forests; pine-oakLiquidambar forests, seasonal evergreen forests, especially near streams; 100–1000(–1400) m. Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol. Selected Specimens Examined. Chis (Breedlove 22133, 31510, DS, NY, 32987, DS, ENCB). Gro (Hinton et al. 10811, NY, US; Lorea 2605, FCME). Oax (Hallberg 1470, NY, UC, US, 1650, NY, UC). Pue (Vera Santos 2573, US). Ver (Copeland herb. 122, B, MICH, NY, UC, US; Ventura A. 8415 ENCB, IEB).

Among the 1-pinnate, idioblastic taxa, A. macrophyllum is easily distinguished from A. obliquum and A. petiolatum by the broader, opposite pinnae, a single sorus on each side of the pinnae, the glabrous stipes, rachises, and pinna surfaces, and the roughly equilateral pinnae. The young fronds are often very reddish or pinkish, a characteristic of many species in the genus, but most pronounced in A. macrophyllum.

17. ADIANTUM MCVAUGHII Adiantum mcvaughii Mickel & Beitel in McVaugh, Fl. NovoGaliciana 17: 147. 1992. Type. Mexico. Nayarit [“Jalisco”:] Tuxpan, Mexia 1030 (MICH!; isotypes MO, NY!, UC!). Figs. 9A–D.

33

Rhizomes short-creeping, 4–7 mm diam.; rhizome scales dark brown, lustrous, 1.5–2 ⫻ 0.1–0.3 mm, margins sparsely denticulate to entire; fronds spaced ca. 5 mm apart, 35–65 cm long, arching; stipes atropurpureous, lustrous, 15–30 cm ⫻ 1–3 mm, ca. 1 ⁄2 the length of fronds, with narrow brown scales to 2 mm long; blades broadly ovate, 2-pinnate, 16–35 ⫻ 19–30 cm; rachises castaneous to atropurpureous, with a mixture of hairs and hair-like scales; pinnae 2–5 pairs plus a conform terminal one, alternate; pinnules not much reduced at the bases or tips of pinnae, the largest 20–34 ⫻ 10–15 mm, tips rounded, the sides forming nearly right angles at pinnule bases, non-articulate; veins free, forking, ending in small teeth; indument adaxially absent, abaxially the costae with a mixture of hairs and hair-like scales, also with scattered golden hairs 0.5–0.8 mm along the veins and tissue between veins, or blades glabrous; idioblasts present on both blade surfaces; sori mostly 3–10 per pinnule, along both the acroscopic and distal margins of pinnules, also sometimes along basiscopic side toward pinnule tips; indusia 2–4 mm long, glabrous.

Distribution. Growing in damp soil, semi-deciduous tropical forests; 20–1000(–2900) m. Mexico. Specimens Examined. Chis (Breedlove 48568, CAS). Nay (Alava & Cook 1521, UC, 1603, MEXU, UC; Gonza´lez Ortega 6657, CAS; McVaugh 19018, MEXU, 19196, CAS, IEB, MICH; Mexia 1030, MICH, MO, NY, UC; Miller & Te´llez 3198, MO, UC; Te´llez & Miller 10494, MEXU, MO). Unverified, Doubtful, or Mistaken Reports. Jal (Maciel s.n., GUADA; Villalobos 164, ZEA, both cited by Mickel & Beitel, 1988, but not verified). Mich (Rzedowski 46175, IEB; Santos Martı´nez 1999, IEB, both cited by Mickel, 1992, but not verified).

It is surprising that there are two collections of this species from very high elevations in Michoaca´n, at 2750 and 2900 m; ordinarily this group of Adiantum occurs below 1500 m. It is most closely related to A. latifolium, differing by the blunt, larger pinnules, more greenish color of the dried blades, and shortercreeping rhizomes. Most specimens, including the type, bear a few, scattered, golden hairs on the veins abaxially. The sole specimen seen from Chiapas, from the Central Depression, is noticeably disjunct from the other collections, but appears not to differ significantly from specimens further north.

34

ADIANTUM

18. ADIANTUM OAXACANUM Adiantum oaxacanum Mickel & Beitel, Mem. New York Bot. Gard. 46: 25. 1988. Type. Mexico. Oaxaca: Distrito Tuxtepec, 2 km N of Chiltepec, Mickel 5804 (NY!; isotype UC!). Figs. 11H, J, K. Rhizomes short-creeping, 2–4 mm diam.; rhizome scales golden brown, 1–3 ⫻ 0.1–0.3 mm, margins sparsely denticulate; fronds spaced 3–8 mm apart, 40–100 cm long, arching; stipes atropurpureous, 25–50 cm ⫻ 2–4 mm, 1⁄2–2⁄3 the frond length, with scattered pectinate appressed scales; blades deltate to oblong, 2-pinnate, 20–35 cm wide; rachises atropurpureous, with tan, appressed, lanceolate scales mostly ca. 1–1.5 mm long, these denticulate or ciliate at margins; pinnae (1–)2–4 pairs plus a conform terminal one, alternate, 12–20 ⫻ 4–7 cm, terminal pinnules triangular and acute at tips, basal pinnules reduced, asymmetrical; pinnules 20–35 ⫻ 10–15 mm, pinnule bases ca. 90⬚ or slightly less, apices rounded, margins finely serrulate, nonarticulate; veins free, forking, ending in small teeth; indument adaxially absent, abaxially the surfaces with scattered, tan, pectinate, hair-like scales to 0.5–1 mm long; idioblasts present on both blade surfaces, inconspicuous; sori mostly 10–14 per pinnule, along acroscopic and distal half of basiscopic margins, as well as on pinnule tips; indusia 2–4 mm long, glabrous.

Distribution. Terrestrial along stream banks and roadsides in wet lowland forests, lower montane forests; 0–800 m. Mexico. Selected Specimens Examined. Oax (Herna´ndez G. 1610, 1611, MEXU, NY; Herna´ndez G. & Gonza´lez 1738, CAS, CHAPA, MEXU; Martı´nez C. 104, NY; Mickel 5132a, NY, UC, 5172, 5804, 6914, NY; Torres C. 627, MEXU). Ver (Copeland herb. s.n., 31 Jan 1938, UC; Delgadillo 12274, UC; Martı´nez C. 1035, XAL; Reeves R5908, ASU). Unverified, Doubtful, or Mistaken Reports. Tab (reported by Magan˜a, 1992, not verified, probably based on a misidentification).

Among the bipinnate, idioblastic species with large pinnules, this species is distinguished by its glabrous indusia, rounded pinnule apices, and pinnule bases ca. 90⬚ or slightly less. It appears most closely related to A. mcvaughii, of western Mexico, from which it differs primarily by the persistent narrow scales and hairs on the blades abaxially. Mickel (1992) stated that the basal pinnules of the latter were less strongly reduced than in A. oa-

xacanum, but this difference seems rather slight, if significant at all. Both species are closely related to A. latifolium and differ from that in the broader, round-tipped pinnules. Mickel and Beitel (1988) thought Adiantum oaxacanum to be intermediate in morphology between A. obliquum and A. amblyopteridium and possibly the fertile hybrid between these two species. Part of the evidence for hybridization was based on the fact that both putative parents and A. oaxacanum were from the same general locality, near Pochutla. This hypothesis needs testing, and it seems just as likely that A. oaxacanum has been derived from the widespread A. latifolium. Herna´ndez & Gonza´lez 1738 (CAS) has prominently ciliate indusial margins, and thus deviates from other specimens seen. Martı´nez C. 1035 (CAS), from Veracruz, has strongly malformed spores and is possibly a hybrid involving A. latifolium and A. oaxacanum.

19. ADIANTUM OBLIQUUM Adiantum obliquum Willd., Sp. Pl., ed. 4, 5(1): 429. 1810. Lectotype (chosen by Lellinger, Mem. New York Bot. Gard. 23: 13. 1972). Venezuela. Caracas, Bredemeyer s.n. (B-Willd. 20067, sheet 1!). Figs. 10A–C. Rhizomes short-creeping, 3–4 mm diam.; rhizome scales lustrous, castaneous, entire to denticulate, 2.5–3 ⫻ 0.5 mm, margins with short teeth; fronds clumped, 25–70 cm long, arching; stipes black, lustrous, to ca. 35 cm ⫻ 1–2.5 mm, ca. 1⁄2–2⁄3 the frond length, with numerous hairs and hair-like scales to 2 mm long; blades oblong to oblong-deltate, 1-pinnate (ours), occasionally the blades with a pair of 1-pinnate proximal pinnae smaller than the terminal part of the blades; rachises atropurpureous, with dense, tan, filiform, contorted scales and hairs; pinnae 6–12 pairs, ovate-lanceolate, proximal pinnae often largest, short-stalked to 5 mm, inequilateral, largest pinnae 3.5–7 ⫻ 1.5–2.5 cm, bases oblique, excised basiscopically, rounded acroscopically, apices acuminate, margins finely denticulate to unevenly biserrate, midveins close to middle of pinnae, nonarticulate; veins free, forking, ending in teeth; indument adaxially absent, abaxially of pectinate, hair-like scales to ca. 1 mm, not glaucous; idioblasts present on both blade surfaces; sori to ca. 20 along both acroscopic and distal portions of basiscopic margins; indusia 2–6 mm long, glabrous or often with numerous short hairs to ca. 0.3 mm long.

ADIANTUM PETIOLATUM

35

to truncate at tips, shallowly lobed to crenate on acroscopic and distal margins, entire on basiscopic margins, subsessile with stalks less than 1 mm long, stalk color passing into pinnulet bases, non-articulate; veins free, forking, ending at the cartilaginous margins; indument on both surfaces of scattered glandular, septate hairs 0.2–1 mm long on veins; idioblasts absent; sori to 10 per pinnulet along acroscopic and distal margins, absent on basiscopic margins; indusia 1–1.5 mm long, lunate to reniform or round-reniform, glabrous.

Distribution. Terrestrial in wet, tropical rain forests, lowland forests, calcareous soils; 100–600 m. Mexico; Guat, Bel, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Tob, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol. Selected Specimens Examined. Chis (Breedlove 33136, DS, NY, 34143, DS). Oax (Herna´ndez G. 249, CAS, CHAPA, ENCB, MEXU, NY). Tab (Cowan 3067, CAS; Gilly & Herna´ndez X. 258, US; Matuda 3325, US, 3336, US; Rovirosa 49, US). Ver (Calzada 7336, XAL; Wendt et al. 2680, CHAPA, NY; Zambrano 1019, CHAPA, MEXU, MO).

The 1-pinnate species (A. obliquum-petiolatum complex) are in need of careful revision over their entire range. Material identified as this species from Oaxaca by Mickel and Beitel (1988) is mostly A. petiolatum. Adiantum obliquum and A. amblyopteridium may have formed the fertile hybrid A. oaxacanum (which see), which is intermediate in morphology. The rounder apices, reduced basal pinnules, and tan spores (easily seen against a white background) distinguish A. oaxacanum from A. obliquum. Occasional specimens of A. obliquum are bipinnate with a single pair of 1-pinnate lateral pinnae: Herna´ndez G. 1751, 1828 (NY), from Oaxaca. In such cases, the terminal pinna is larger (longer and wider) than the lateral pinnae.

20. ADIANTUM PATENS Adiantum patens Willd., Sp. Pl., ed. 4, 5(1): 439. 1810. Type. Venezuela. Caracas, Bredemeyer s.n. (B-Willd. 20078!, photo GH!, frag. NY!; isotype W). Figs. 6E, F. Adiantum lobatum C. Presl, Reliq. Haenk. 1(1): 62, t. 10, fig 5. 1825. Type. Mexico. s. coll. [presumably Ne´e] s.n., (PR).

Rhizomes short-creeping, 3–5 mm diam.; rhizome scales light brown, ciliate-fimbriate, 2–3 ⫻ 0.8–1 mm; fronds clumped, 20– 45(–70) cm long, laxly arching; stipes castaneous to atropurpureous, 12–40 cm ⫻ 1–2(–3) mm, 1⁄2–2⁄3 the frond length, glabrous except at bases; blades imparipedate, almost round in outline, to 30 ⫻ 30 cm, the two proximal pinnae alternate, each appearing 2–3-forked by virtue of unequal basiscopic branching; rachises castaneous, with dense, branched whitish hairs 0.1 mm long; pinnae a single lateral pair plus a terminal one, the lateral ones to 20 cm long, unequally basiscopically branched 2–3 times, the terminal one 1-pinnate, to 25 cm long; pinnulets oblong, rounded

Distribution. Terrestrial on road banks and open woods, pineoak forests; 200–1900(–2100) m. Mexico; Guat, Hond, Salv, CR; Col, Ven, Ec, Peru, Bol; Africa (subsp. oatesii) Selected Specimens Examined. Chis (Purpus 7113, UC). Col (Jones 500, UC, US). Dgo (Sa´nchez 747, MEXU, US). Gro (Mexia 8802a, B, UC). Jal (Pringle 1869, B, BR, DS, K, NY, UC, US). Me´x (Hinton 4751, NY). Mich (Dı´az Luna 7491, GUADA, UC). Mor (Lyonnet 1374, CAS, ENCB, MEXU, US). Nay (Te´llez 12821, IEB, MEXU). Oax (Mickel 6105, NY, UC). Sin (Gonza´lez Ortega 586, ENCB, MEXU). Son (Boutin & Kimnach 3673, NY).

This species is extremely common on the Pacific slopes in Mexico (e.g., over a hundred collections have been seen from Nayarit and Jalisco alone) but absent elsewhere in the country. The seemingly pedate branching of the fronds and the minute, branching rachis hairs distinguish this species from all others. Although superficially similar to A. aleuticum, which has similar blade architecture, the closest relatives appear to be A. galeottianum and A. shepherdii, which see. A closely related subspecies occurs in Africa, subsp. oatesii (Baker) Schelpe.

21. ADIANTUM PETIOLATUM Adiantum petiolatum Desv., Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck. Gesammten Naturk. 5: 326. 1811. Type. Brazil or French Guiana, not found at P by R. Tryon or others. Figs. 8H, 10D. Adiantum lucidum (Cav.) Sw. var. pinnatum E. Fourn., Mexic. Pl. 1: 129. 1872. Lectotype (chosen by Lellinger, Proc. Biol. Soc. Wash. 89: 704. 1977). Mexico. Tabasco: Linden s.n. (P, Morton photo 2605, US!).

Rhizomes short-creeping, 2–4 mm diam.; rhizome scales castaneous, entire to faintly denticulate, linear-lanceolate, 2.5–3 ⫻

36

ADIANTUM

0.5 mm; fronds distant, mostly 2–5 mm apart, 22–45 cm long, arching; stipes black, lustrous, 10–25 cm ⫻ 1–1.5 mm, ca. 1⁄2–2⁄3 the frond length, glabrous or with sparse pectinate scales to 2 mm long; blades oblong-deltate, 1-pinnate; rachises atropurpureous, with scattered scales and hairs, glabrescent, the scales linear, branched or pectinate at the bases, otherwise entire or nearly so; pinnae 6–9 pairs, the proximal pinnae often largest, 3.5–6.5 cm long, distal pinnae 1⁄2–3⁄4 as long, lateral pinnae inequilateral, stalked 1–5 mm, stalk color passing into pinna bases, terminal pinna subconform, equilateral, pinna bases oblique, margins finely and evenly serrulate, apices acute or acuminate, costae near mid-pinna or slightly closer to basiscopic margins, nonarticulate; veins free, forking, ending in marginal teeth; indument absent on both surfaces; idioblasts present on both blade surfaces, conspicuous adaxially, sometimes faint abaxially; sori 6–14 along acroscopic margins, also to 9 along distal part of basiscopic margins; indusia 1–6 mm long, oblong, linear, or slightly arcuate, glabrous; 2n⫽60 (Braz).

Distribution. Terrestrial along shaded wet streams banks, seasonal evergreen forests, lowland and lower montane rain forests; 100–600 m. Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Tob, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol.

closely related, differing primarily in being bipinnate, rather than simply pinnate.

22. ADIANTUM POIRETII Adiantum poiretii Wikstr., Kongl. Vetensk. Akad. Handl. 1825: 443. 1826. Based on A. crenatum Poir. in Lamarck, Encycl. Suppl. l: 137. Sep 1810, non Willd., Mar 1810. Type. Tristan da Cunha, Petit-Thouars s.n. (P-JU 1427!). Figs. 10E, F. Adiantum thalictroides Willd. ex Schltdl., Adumbr. Pl. 5: 53. 1832. Type. Mauritius, Petit-Thouars s.n. (B-Willd. 20101!, BR! photo ex P). Adiantum pellucidum M. Martens & Galeotti, Me´m. Foug. Mexique 72. 1842. Type. Mexico. Oaxaca: Tanetze & Talea [Llano Verde; isotype], Galeotti 6461 (BR!, photo P! ex BR, photo US! ex BR; isotypes BR!, C!, K!, LE!, P!). One of the specimens from BR is A. concinnum. Adiantum gratum Fe´e, Me´m. Foug. 5: 119. 1852. Type. Mexico. Oaxaca: near Nolasco, Galeotti 6542 (P? not found) [perhaps 6562 (BR!, K!)].

Rhizomes short- to usually long-creeping, 1.5–3(–4) mm diam.; rhizome scales castaneous, lanceolate, 3–4 ⫻ 0.5–0.8 mm, entire to short-ciliate; fronds distant, (10–)20–60(–75) cm long, arching; stipes ca. 1⁄4–1⁄2 the frond length, 7–35 cm ⫻ 0.8–2 mm, castaneous to atropurpureous, glabrous except at bases; blades ovate-deltate, (2–)3–4-pinnate proximally, (5–)15–45 ⫻ 4–35 cm, gradually tapering to apices; rachises castaneous to atropurpureous, glabrous; pinnae 3–9 pairs, alternate, tapering at apices; pinnulets orbicular to flabellate, 5–8 ⫻ 5–10(–15) mm, mostly symmetrical, not articulate, cuneate or truncate at bases, entire or with a few teeth distally, stalk color passing into pinnulet bases and often into veins, non-articulate; veins free, forking, ending in sinuses of sterile pinnulets; indument absent on both surfaces, or glaucous abaxially; idioblasts absent; sori mostly 3–8 per pinnulet, confined to distal margins, yellow farina mixed with sporangia; indusia 1–3 mm long, lunate to reniform or roundreniform, glabrous; 2n⫽114 (Africa).

Selected Specimens Examined. Chis (Breedlove 49150, 56781, CAS; Ventura & Lo´pez 2110, ENCB, NY). Oax (Mickel 4184, MEXU, NY, UC, 5175, 5832, 6834, NY, UC). Tab (Matuda 3336, NY). Ver (Martı´nez C. 1035, ENCB, MEXU, XAL; Orcutt 3201, DS, MEXU, NY, US).

All specimens here cited from Oaxaca were identified as A. obliquum by Mickel and Beitel (1988). However, we now believe that they are more properly assigned to A. petiolatum because of their simply and finely serrulate pinna margins, glaucous abaxial blade surfaces, and relatively sparsely scaly rachises. Specimens agreeing with the Mexican ones are relatively common in Mesoamerica. Adiantum obliquum differs in having doubly and more coarsely serrate pinna margins, green blade surfaces, scalier blade surfaces abaxially, and denser rachis scales. The character of the idioblasts used by Moran et al. (in Davidse et al., 1995), conspicuous on both sides in A. obliquum, but conspicuous on the adaxial side only in A. petiolatum, while perhaps a tendency, does not seem to be definitive. Adiantum latifolium is also very

Distribution. Terrestrial, higher elevations, pine-oak woods, oak woodlands with tropical deciduous forest elements, near streams; 800–3150 m. Mexico; Guat, Hond, Salv, Nic, CR, Pan; Cuba?, Hisp; Col, Ven, Ec, Peru, Bol, Chile, Arg; Old World.

ADIANTUM PULVERULENTUM Selected Specimens Examined. Ags (Garcı´a R. 2417, HUAA, IEB). BCS (Jones 24558, NY, UC, US). Chih (Correll & Gentry 23045, UC). Chis (Breedlove 22694, DS, NY). Col (Alava & Cook 1705, UC). DF (Mexia 2666, UC). Dgo (Tenorio L. 2146, ENCB, MEXU). Gro (Mexia 9009, B, UC). Gto (Flores 72, ENCB, IEB). Hgo (Knobloch 678, ENCB, US). Jal (Pringle 1867, B, UC, US). Me´x (Lyonnet 250, NY). Mich (Bartholomew 2835, CAS, NY). Mor (Lyonnet 1846, MEXU, NY). NL (Hinton 22687, TEX). Oax (Mickel 1170, ENCB, NY, UC). Pue (Marcks & Marcks 747, ENCB). Qro (Herrera 16, ENCB, IEB). SLP (Rzedowski 10060, ENCB). Sin (Gentry 7278, UC, US). Son (Sanders et al. 13481, UC). Tlax (Vela´zquez s.n., ENCB). Ver (Nee 33077, NY, UC).

37

coarsely serrate, terminal pinnule of each pinna non-conform, elongate, proximal pinnules reduced and deltate, non-articulate; veins free, forking, ending in teeth in sterile pinnules; indument adaxially absent, abaxially usually of narrow tan scales 0.8 mm long, blades not glaucous; idioblasts present on both blade surfaces; sori generally one per pinnule (infrequently with 1 or 2 smaller sori distal to the linear main one), confined to acroscopic margins; indusia 5–10 mm long, linear, glabrous; 2n⫽60, 90 (Jam).

Unverified, Doubtful, or Mistaken Reports. Nay (reported by Mickel & Beitel, 1988, and Mickel, 1992, but not verified; Calzada 18618, MEXU, US, determined as this in various herbaria, is A. tricholepis). Tam (reported by Mickel & Beitel, 1988, and Mickel, 1992, but not verified; also reported by Sharp, 1954, from Rancho del Cielo, but not verified). Tres Marı´as (Ferris 5652, DS, cited by Lenz, 1995, but not verified).

This species and A. andicola are by far the commonest species of Adiantum in Mexico at middle to high elevations, and A. poiretii is especially common in and around the Valley of Mexico. Material from Tristan da Cunha is sometimes considered distinct, in which case all other material is called A. thalictroides Schltdl. Adiantum poiretii is distinguished from A. raddianum by the golden brown scales, usually longer-creeping rhizomes, smaller pinnulets, and yellow farina in the sori. Specimens from northwestern Mexico and Baja California Sur often show a pronounced dimorphism, with the sterile fronds having much larger pinnulets, nearly twice as large as the fertile ones. Given only a sterile frond, it is easy to mistake this species for A. braunii, which may occur in the same general area.

23. ADIANTUM PULVERULENTUM Adiantum pulverulentum L., Sp. Pl. 2: 1096. 1753. Lectotype (chosen by Proctor, 1977: 185). Plumier, Descr. Pl. Ame´r., pl. 47, 1693 (⫽ Traite´ Foug. Ame´r., t. 55, 1705), based on a plant from Martinique or Hispaniola. Fig. 8G. ?Adiantum monotis Nees, Linnaea 19: 684. 1847. Type. Mexico. Aschenborn 348 (B?). This was placed as a synonym of A. villosum by Lellinger (1989: 159), but based on the description, it seems more likely to be A. pulverulentum; no one recently has examined the type. For additional synonymy, see Lellinger (1989: 153).

Rhizomes short-creeping, 4–8 mm diam.; rhizome scales dark brown, 1.5–2.5 ⫻ 0.3–0.8 mm, linear-lanceolate, with marginal hairs 0.1–0.3 mm long; fronds clumped, 40–105 cm long, arching; stipes atropurpureous, lustrous, 20–55 cm ⫻ 2–5 mm, ca. 1⁄2– 2 ⁄3 the frond length, with small (to 3 mm long), brown, pectinate scales; blades deltate, 2-pinnate, (20–)25–50 cm wide, each blade with a conform terminal pinna; rachises atropurpureous, with rather dense tan lanceolate scales and hair-like scales; pinnae 3– 8(–10) pairs, alternate, gradually tapering at the tips, stalked 3– 8 mm; pinnules dimidiate, oblong, largest 10–20 ⫻ 4–6 mm, fertile pinnules obtuse or blunt, straight, or the apices (beyond sori) curved toward pinna tips, sterile pinnules acute, falcate,

Distribution. Terrestrial in wet forests at low elevations, lower montane rain forests, pine-oak-Liquidambar forests; 50–1000 (–1400) m. Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Guy, Sur, Fr, Gui, Ec, Peru, Braz, Bol. Selected Specimens Examined. Chis (Breedlove 22457, DS, NY; Purpus 6854, UC). Gro (Gonza´lez 341, FCME). Oax (Mexia 9196, CAS, GH, MICH, NY, UC, US; Mickel 7223, NY, UC). QR (Uca´n & Flores 935, CICY, this sheet containing also A. decoratum; in other herbaria this number is A. decoratum). SLP (Copeland herb. s.n., UC). Tab (Cowan 3433, CSAT, UC). Ver (Palacios-Rios 43, ENCB, IEB, UAMIZ; Purpus 5942, B, MO, NY, US).

This species is very common in lowland areas of Veracruz, Oaxaca, and Chiapas, but rare elsewhere in Mexico. Adiantum pulverulentum is distinctive among the short-pinnuled, bipinnate, idioblastic species in usually having one long sorus per pinnule, along the acroscopic margin (and occasionally at the distal end). Occasional specimens, especially those that are only partially fertile, have the sori somewhat lobed: Ver (Smith 2084, UC), Oax (Hallberg 1264, NY). Abortive-spored hybrids between A. pulverulentum and A. villosum (Fig. 8F) have large, acute pinnae, one sorus per pinnule, and glabrous indusia: Ver (Mickel 7226b, NY, UC; Purpus 7765, UC, US), Oax (Conzatti et al. 31475, US; Mickel 5777, NY, UC, Fig. 41F), Chis (Breedlove 38671, DS, ENCB; Fisher 35400, NY). Another probable hybrid involving A. pulverulentum is Breedlove 34993 (DS), from Chiapas. This hybrid may involve either A. tetraphyllum or A. decoratum as the second parent. This specimen has strongly malformed spores and variably continuous or discrete sori along the acroscopic pinnule margins.

38

ADIANTUM

24. ADIANTUM RADDIANUM Adiantum raddianum C. Presl, Suppl. Tent. Pterid. 158. 1836. Type. Based on Raddi, Pl. Bras. Nov. Gen., t. 78, f. 2. 1825. Figs. 5F–H, P, Q. Adiantum cuneatum Langsd. & Fisch., Pl. Voy. Russes Monde 1: 23. 1810 (t. 26, 1818), hom. illeg., non G. Forst., 1786. Type. Brazil. Santa Catharina Island, Langsdorff s.n. (LE! photo GH; isotypes B-Willd. 20096!, microfiche UC!, BM). Adiantum mexicanum C. Presl, Suppl. Tent. Pterid. 158. 1836. Presl cited only Adiantum cuneatum Schltdl., which is an indirect reference to a specimen (Schiede & Deppe “797”) cited by Schlechtendal in Linnaea 5: 615 (1930), without further description. The Schiede & Deppe collection (frag. NY!) is A. andicola. But because A. mexicanum has no description of its own, it must be regarded as an illegitimate renaming of A. cuneatum, which is here and by others treated as a synonym of A. raddianum. For additional synonymy, see Lellinger (1989: 153–154).

Rhizomes short-creeping, 3–5 mm diam.; rhizome scales dark to light brown, linear-lanceolate, entire, 2.5–3 ⫻ 0.5 mm; fronds clumped, ca. 20–45 cm long, laxly arching; stipes dark reddish brown, 10–20 cm ⫻ 1–2 mm, ca. 1⁄2 the frond length, glabrous except at bases; blades ovate-deltate to ovate-lanceolate, 2–3pinnate proximally, 1–2-pinnate distally, tapering to apices; rachises red-brown, glabrous; pinnae 6–9 pairs, pinnae and pinnules with stalks 1–5 mm long; pinnulets obovate to rhomboid, 8–12 mm long, sterile margins dentate, the stalk color passing into pinnulet bases, non-articulate; veins free, forking, ending in sinuses; indument absent on both surfaces; idioblasts absent; sori 4–6 per pinnulet, confined to distal margins; indusia 1–1.5 mm long, reniform to round-reniform, glabrous.

Distribution. Terrestrial on open or lightly shaded banks, oak forests, seasonal evergreen forests; 800–2200 m. Mexico; Guat, Nic, CR; Gr & L Ant; Col, Ven, Trin, Ec, Peru, Braz, Bol, Parag, Arg, Uru; widely naturalized in OW tropics. Selected Specimens Examined. Chis (1971, Breedlove 21589, DS; 1972, 27375, DS). Mich (1985, Leopo 17084, GUADA, UC). Mor (1938, Lyonnet 2565, CAS, MO, UC, US; 1966, Mitastein 168, ENCB, NY; 1994, Pacheco 4669, UAMIZ; 1992, Posadas & Sandoval s.n., UAMIZ; 1993, Riba 1909, UAMIZ; 1991, Sandoval 6, UAMIZ). Oax (1969, Mickel 4020a, NY, UC, 1971, 5999, 6026, NY, UC). Pue (1971, Weber 834, 852, ENCB). Sin (1969, Breedlove 17107, NY). Tlax (1970, Weber 734, ENCB, NY). Ver (Fisher s.n., DS, questionably det.).

Because of the spottiness and recency of collections of this species in Mexico, we suspect that A. raddianum may not be native, but rather recently introduced; dates of collection, where known, are given in the specimen citations, and all but one of the Mexican collections were made since 1966. Proctor (1985) discussed the possible naturalization of A. raddianum in Jamaica. It is certainly widely cultivated throughout the world, including Mexico, and it would be surprising if it had not escaped. However, nowhere, except possibly in the state of Morelos, does it seem to be common. Adiantum raddianum is closely related to A. poiretii, which see for distinguishing characters. The Fisher collection cited from Veracruz is immature and lacks rhizome, but it appears to belong to this species (although not matching other Mexican specimens very well); we wonder if it is a horticultural variant of A. raddianum.

25. ADIANTUM SEEMANNII Adiantum seemannii Hook., Sp. Fil. 2: 5, t. 81A. 1851. Type. Panama. Veraguas: San Lorenzo, Seemann 1124 (K! photo UC!). Figs. 12E–G. Rhizomes short-creeping, rather compact, 5–10 mm diam.; rhizome scales castaneous, concolorous, entire to minutely denticulate; fronds clumped, mostly 30–70 cm long, arching; stipes atropurpureous to black, to 45 cm ⫻ 3 mm, glabrous except at bases; blades ovate to deltate, 1-pinnate or some specimens 2pinnate proximally, each blade with a Ⳳ conform terminal pinna; rachises atropurpureous to black, glabrous; pinnae 1–4 pairs, broadly ovate, mostly 5–10 ⫻ 2.5–6 cm, rounded at bases, acuminate at apices, serrate (sterile) to entire (fertile), with stalks 1–3 cm long, stalk color stopping abruptly at pinna bases, pinnae not articulate; pinnules absent on pinnae, or the proximal pinna pair 1-pinnate; veins free, forking, ending in sharp teeth on sterile pinnae/pinnules; indument absent on both surfaces, blades often glaucous abaxially; idioblasts present on both blade surfaces, more conspicuous adaxially; sori linear, usually interrupted (but contiguous) on both acroscopic and basiscopic margins of pinnae or pinnules; indusia mostly 5–20 mm long, especially near bases of pinnae/pinnules, glabrous.

ADIANTUM TENERUM

Distribution. Terrestrial in lowland and lower montane forests on steep rocky slopes; 200–450 m. Mexico; Guat, Hond, Nic, CR, Pan; w Col. Specimens Examined. Chis (Breedlove 31586, DS, NY, 34206, DS; Croat 43786, CAS, ENCB, MO; Sohns 1615, MEXU, US).

Adiantum seemannii is readily distinguished from congeners in Mexico by the 1-pinnate blades, relatively few lateral pinnae (1–4 pairs), and large, ovate pinnae 5–10 cm long. It is related to species in South America, e.g., A. platyphyllum Sw., from Ecuador to Bolivia and southern Brazil.

26. ADIANTUM SHEPHERDII

39

Distribution. Terrestrial on rocky banks, oak woods, pine-oak forests; 400–1850(–2200) m. Mexico. Selected Specimens Examined. Gro (Hinton 11347, DS, MEXU; Langlasse´ 322, MEXU, US; Mexia 8804, B, CAS, NY, UC, US). Me´x (Hinton 1602, MEXU, 4758, K, US; Matuda 31053, MEXU, UC, US). Mich (Hinton 13485, ENCB, NY, US; King & Soderstrom 4806, MEXU, UC; Rzedowski 51698, IEB, MEXU; Ventura 2484, ENCB, NY).

This species is characterized by the linear fronds with many pairs of strongly overlapping and very gradually reduced distal pinnae at the almost indeterminate blade apices. The closest relatives are A. galeottianum and A. patens, which occur in the same general area. Adiantum shepherdii is a relatively rare endemic at middle elevations in western Mexico.

Adiantum shepherdii Hook., Sp. Fil. 2: 9, t. 73B. 1851. Type. Mexico, 1834, Bates s.n. (K!). Figs. 11A–C.

27. ADIANTUM TENERUM

Rhizomes short-creeping, compact, 3–6 mm diam.; rhizome scales golden brown, linear-lanceolate, 2–3 ⫻ 0.3–0.5 mm, sparsely ciliate-denticulate; fronds clumped, 20–46 cm long, arching; stipes castaneous to atropurpureous, 2–6(–15) cm ⫻ 1–2 mm, 1⁄10–1⁄5 the frond length, glabrous except at bases; blades linear, 1-pinnate, 2–3.5 cm wide, infrequently 2-pinnate at the bases; rachises castaneous or red-brown, glabrous; pinnae 34–56 pairs, to 1.7 ⫻ 1 cm, alternate, very gradually reduced toward the blade apices, sessile, obliquely flabellate or dimidiate, bases often overlapping the rachises, sterile margins entire to broadly crenate, distinctly thickened; pinnules absent on pinnae, except infrequently on the proximal pair, those sometimes 1-pinnate, to 15 ⫻ 2 cm with up to 15 pinnules, non-articulate; veins free, forking, ending at the cartilaginous margins; indument absent on both blade surfaces; idioblasts absent; sori 3–6 per pinna, confined to acroscopic and distal margins; indusia 1–2 mm long and wide, round-reniform and with deep sinuses, glabrous.

Adiantum tenerum Sw., Prodr. 135. 1788. Type. Jamaica. Swartz s.n. (S). Figs. 7D–F. Adiantum trapezoides Fe´e, Me´m. Foug. 5: 117. 1852. Type. Mexico. Veracruz: Puente Nacional, Galeotti 6317 (P!; isotype RB). ?Adiantum extensum Fe´e, Me´m. Foug. 8: 72. 1857. Syntypes. Mexico. Veracruz: near Orizaba and Huatusco, Schaffner 40 (P?), 41 (P?, RB).

Rhizomes short-creeping, 5–10 mm diam.; rhizome scales ferrugineous, becoming dark brown in center portion (and so somewhat bicolorous), lustrous, 2–3 ⫻ 0.8–1 mm, margins ciliate-fimbriate; fronds clumped, 25–110 cm long, laxly arching; stipes atropurpureous to blackish, lustrous, 10–55 cm ⫻ 1.5–3 (–4) mm, 1⁄3–1⁄2 the frond length, glabrous or with rare minute tubercles; blades ovate-deltate, 3–5-pinnate proximally, 18–15 ⫻ (15–)20–40 cm wide; rachises atropurpureous, glabrous, somewhat flexuous toward tips; pinnae 5–9 compound pairs, alternate, stalked to 4 cm, gradually tapering distally; pinnulets trapeziform to rhombic, or flabellate at blade and pinna apices, fertile pinnulets 1–2 cm long and wide, cuneate at bases, denticulate cleft or lobed distally, sterile pinnulets somewhat larger than the fertile ones, stalked 2–4 mm, stalk color stopping abruptly at bulge at pinnulet bases, articulate; veins free, forking, ending in marginal teeth; indument absent on both blade surfaces, blades sometimes glaucous abaxially; idioblasts absent; sori to ca. 12 per pinnulet, confined to distal margins, with slight

40

ADIANTUM

yellow farina among sporangia; indusia 1–3 mm long, reniform to oblong, glabrous; 2n⫽60 (USA, Ver, CR, Jam).

ceolate, denticulate scales to 1.5 mm long; pinnae 2–5 pairs, alternate, stalked to ca. 5 mm; pinnules oblong, 1.3–2.5 ⫻ 0.4– 0.6 cm, sessile to short-stalked up 1 mm, non-articulate; veins free, forking, ending in sharp teeth; indument of numerous golden, septate hairs 1–1.5 mm long abaxially, glabrous adaxially; idioblasts present, especially prominent adaxially; sori to ca. 8–10 pairs along acroscopic margins and also distal half of basiscopic margins; indusia 1–2 mm long, oblong, with golden hairs like those of blades, but shorter, to ca. 0.5 mm long.

Distribution. Terrestrial in lowland forests of the Atlantic lowlands, woods along streams, tropical forests, evergreen rain forests, coffee plantations, calcareous cliffs; 50–1350 m. USA (Fla); Mexico; Guat, Bel, Hond, Nic, CR; Bah, Gr & L Ant; Ven, Trin. Selected Specimens Examined. Camp (Martı´nez S. 27319, XAL). Chis (Breedlove 23073, DS, NY). Hgo (Gimate 948, ENCB, NY). NL (Aguirre C. 648, ENCB). Oax (Mickel 5818, NY, UC). Pue (Knobloch 696, ENCB, US). Qro (Rzedowski 42853, IEB). QR (Torres 23, 33, CIQRO, MEXU). SLP (Pringle 3360, BR, DS, NY, P, UC, US). Tab (Cowan 3163, CAS, ENCB). Tam (Dressler 1888, UC). Ver (Copeland herb. 116, MICH, P, UC). Yuc (Calzada et al. 6702, UC, XAL). Unverified, Doubtful, or Mistaken Reports. Tres Marias (Nelson 4281, US, cited by Lenz, 1995, as A. tenerum, and Ferris 5704, DS, and Mason 1822, CAS, cited by Lenz as A. trapezoides, but none verified, all three probably misidentifications).

Adiantum tenerum most closely resembles A. amplum, from which it differs by having smaller, bicolorous rhizome scales, serrulate sterile pinnulets, and the generally more flabellate pinnulets. They also differ in general distribution, A. tenerum being an Atlantic slope species and A. amplum a Pacific slope species. Adiantum tenerum is very common in Veracruz and Chiapas, but less common elsewhere. A specimen cited by Dı´az-Barriga and Palacios-Rios (1992) from Michoaca´n (Hinton 15956, ENCB) is A. amplum.

28. ADIANTUM TERMINATUM Adiantum terminatum Kunze ex Miq., Inst. Versl. Meded. Kon. Naderl. Inst. Wetensch. 1842: 187. 1843. Type. Surinam. Focke 370 (U). Figs. 12L, M. Rhizomes short-creeping, 2–3 mm diam.; rhizome scales atropurpureous, 1–2 ⫻ 0.2–0.3 mm, entire or denticulate; fronds 2–5 mm apart, 30–50 cm long, arching; stipes atropurpureous, lustrous, to 30 cm ⫻ 1–1.5 mm, 1⁄2–2⁄3 the frond length, with scattered small scales 1–2 mm long, glabrescent; blades deltate, 2-pinnate, ca. 15 ⫻ 18 cm, each blade with a conform terminal pinna; rachises castaneous to atropurpureous, with linear to lan-

Distribution. Terrestrial in lowland rain forests; 350 m. Mexico; Guat, Bel, CR, Pan; Col, Ven, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol. Specimen Examined. Chis (Breedlove 38988, DS).

Most of the Chiapan specimens cited by Smith (1981) as this species we now refer to other taxa: Breedlove 26534 (NY) is a mixed collection of A. tetraphyllum and A. trichochlaenum; Breedlove 26564 (NY) and Breedlove 32898 (DS, NY) are A. trichochlaenum.

29. ADIANTUM TETRAPHYLLUM Adiantum tetraphyllum Humb. & Bonpl. ex Willd., Sp. Pl., ed. 4, 5(1): 441. 1810. Lectotype (chosen by Lellinger, Proc. Biol. Soc. Wash. 89: 704. 1977). Venezuela. near Guanaguana and Caripe, Humboldt & Bonpland s.n. (B-Willd. 20082–2 sheets!, frag. NY!). Fig. 8E. For additional synonymy, see Lellinger (1989: 157).

Rhizomes short- to usually long-creeping, 3–5 mm diam.; rhizome scales golden brown, 2–3 ⫻ 0.3 mm, linear-lanceolate, sparsely denticulate; fronds clumped to distant, 35–100 cm long, arching; stipes atropurpureous, lustrous, 25–55 cm ⫻ 2–3 mm, 1⁄2–3⁄5 the frond length, with brown scales to 3 mm long; blades deltate, 2-pinnate, ca. 25–35 cm wide, each blade with a large conform terminal pinna; rachises atropurpureous, with numerous linear to lanceolate, denticulate scales to 1.5 mm long; pinnae 1–4(–5) alternate pairs, terminal pinnules elongate, pinnatifid; pinnules 15–40 ⫻ 5–7 mm long, tips acute to acuminate, falcate, non-articulate; veins free, forking, ending in teeth at the margins;

ADIANTUM TRAPEZIFORME

indument adaxially absent, abaxially the surfaces with a few scattered hair-like scales; idioblasts conspicuous on both blade surfaces; sori mostly 3–6 per pinnule, confined to acroscopic margins; indusia 1–4 mm long, glabrous; 2n⫽58–60 (Jam).

Distribution. Terrestrial in lowland forests and lower montane forests, seasonal evergreen forests, montane rain forests; 150– 350(–1250) m. Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Ec, Peru, Bol, Parag. Selected Specimens Examined. Chis (Breedlove 39999, DS, 56582, CAS, ENCB, NY, 57237, CAS; Matuda 18398, DS, MEXU; Purpus 6742, UC, US). Oax (Hallberg 1617, NY; Herna´ndez G. 735, 1596, NY; Yatskievych 83-439a, NY). Ver (Copeland herb. 120, CAS, MEXU, MICH, UC; Purpus 2930, 15713, UC).

41

Adiantum trapeziforme L. var. oblongatum T. Moore, Index Fil. 40. 1857. Syntypes. Mexico. Veracruz: Cordillera de Veracruz, Galeotti 6338 (K); Guatemala, Skinner s.n. (K); Cuba. Poeppig s.n. (K). Based on the description of A. trapeziforme γ, an unnamed variant, in Hook., Sp. Fil. 2: 33. 1851. Adiantum trapeziforme L. var. plumieri T. Moore, Index Fil. 40. 1857. Syntypes. Mexico. Veracruz: near Veracruz, Linden 70 (K); Plumier, Traite´ Foug. Ame´r., pl. 95. 1705. Based on the description of A. trapeziforme β, an unnamed variant, in Hook., Sp. Fil. 2: 33. 1851.

Rhizomes short- to long-creeping, 3–5 mm diam.; rhizome scales bicolorous, light brown at bases to lustrous dark brown at tips, linear-lanceolate, denticulate, 2–3 ⫻ 0.1–0.3 mm; fronds distant, (40–)60–100 cm long, arching; stipes atropurpureous to blackish, lustrous, 35–60 cm ⫻ 4–6 mm, nearly 1⁄2 the frond length, glabrous except at bases, with minute tubercles; blades broadly ovate, 3–4-pinnate proximally, 2-pinnate distally, each blade with a conform, 1-pinnate terminal pinna; rachises atropurpureous, glabrous; pinnae 3–4 compound pairs, the proximal pair the longest, 25–35 ⫻ ca. 15 cm; pinnulets mostly trapeziform, inequilateral, (20–)35–50 ⫻ 10–20 cm, acute to acuminate at tips, sterile margins incised-lobate, fertile margins only shallowly so, stalked 1–5 mm, articulate, stalk color ending abruptly at pinnulet bases, articulate; veins free, forking, ending in teeth at the margins; indument absent on both blade surfaces, blades sometimes glaucous abaxially; idioblasts absent; sori 10–15 along acroscopic margins of pinnules, 1–2 per lobe, also 3–5 along distal margins; indusia 1–3 mm long; oblong, glabrous; 2n⫽60, 90 (cult).

Unverified or Mistaken Reports. Nay (reported by Mickel & Beitel, 1988; not treated in the range by Mickel, 1992; Gonza´lez O. 6657, identified as this in CAS, is A. mcvaughii). Tab (reported by Magan˜a, 1992, possibly based on a misidentification).

Among the bipinnate, idioblastic species with large pinnae, this taxon is distinctive in its combination of usually longcreeping rhizomes (0.5–2 cm between internodes), abaxial blade surfaces with scattered hair-like scales, numerous sori along the acroscopic pinnule margins, and pinnules that tend to curve toward the pinna tips. The indusia may be either glabrous (usually) or bear a few short hairs 0.1–0.3 mm long (as in Breedlove 37681, 42002, DS, from Chiapas). As commonly construed, A. tetraphyllum represents a species complex (see Kramer, Acta Bot. Neerl. 3: 481–486. 1954) that is especially diverse in South America. There is a very wide variety of material passing under this name in herbaria, some of it belonging to closely related species. We now regard all the specimens identified as A. fructuosum by Smith (1981) as A. tetraphyllum. Breedlove 48595 (CAS, NY), cited by Mickel and Beitel (1988) as A. trichochlaenum, has only a few weak hairs on a few indusia, and we now regard this also as A. tetraphyllum.

30. ADIANTUM TRAPEZIFORME Adiantum trapeziforme L., Sp. Pl. 2: 1097. 1753. Lectotype (chosen by Lellinger, Proc. Biol. Soc. Wash. 89: 704. 1977). Sloane, Voy. Jamaica 1: t. 59. 1707. Figs. 7K–N.

Distribution. Terrestrial in wet forests at lower elevations; 50– 900 m. Mexico; Guat, Bel, Hond, Salv, Nic, CR; Cuba, Jam, L Ant; Trin, Bol, Parag; widely naturalized in Asia and Pacific Islands. Selected Specimens Examined. Chis (Davidse et al. 30135, MO, NY). Gro (Palmer 519, B, NY). Hgo (Gimate 833, CAS, ENCB). Jal (Dieterle 4056, CAS, ENCB, IEB, MICH). Me´x (Hinton 7644, NY). Mich (Hinton 12277, NY). Nay (McVaugh 18799, IEB, NY). NL (Rodrı´guez s.n., 22 Apr 1979, ANSM, ENCB, UNL). Oax (Hallberg 1263, ENCB, NY). Pue (Segura A. 16, FCME). Qro (Rzedowski 42804, ENCB, IEB). SLP (Pringle 3960, B, BR, CAS, ENCB, NY, P). Tam (Palmer 303, B, NY). Ver (Calzada 917, ENCB, MEXU, MO, XAL). Unverified, Doubtful, or Mistaken Reports. Tab (reported by Magan˜a, 1992, but not verified).

42

ADIANTUM

Among the tripinnate species with a conform terminal pinna, A. trapeziforme is distinctive in its glabrous stipes, rachises and pinnulets, articulate pinnulets, and lack of epidermal idioblasts. South American material has been ascribed to this species; some of these specimens, especially from Peru, Bolivia, and Brazil, have been named A. mathewsianum Hook., which is certainly very closely related to A. trapeziforme. Additionally, true A. trapeziforme may be naturalized in South America, as it seems to be in Paraguay and in areas of Asia and the Pacific. Further study is needed.

31. ADIANTUM TRICHOCHLAENUM Adiantum trichochlaenum Mickel & Beitel, Mem. New York Bot. Gard. 46: 29–30. 1988. Type. Mexico. Oaxaca: Distrito Putla, 15 km S of Putla, 12 km N of Mezones, Mickel 6975 (NY!; isotype UC!). Figs. 12N–P. Rhizomes short-creeping, 5–8 mm diam.; rhizome scales light brown with thick, dark brown cell walls, 1–1.5 ⫻ 0.3–0.5 mm, with recurved teeth along margins; fronds clumped, 55–70 cm long, arching; stipes atropurpureous, 32–40 cm ⫻ 2–4 mm, 1⁄2– 2 ⁄3 the frond length, with scattered pectinate scales to 2 mm long, glabrescent except for scales at bases; blades deltate to oblong, 2pinnate, 20–30(–50) cm wide, each blade with a conform terminal pinna; rachises atropurpureous, with persistent lanceolate to linear scales 1–2 mm long; pinnae 2–5 lateral pairs, terminal pinnules gradually reduced toward the pinna tips; pinnules 10– 25 ⫻ 4–9 mm, rounded to acute at tips, reduced to equilateral triangular pinnules at pinna bases, non-articulate; veins free, forking, ending in teeth at the margins; indument adaxially absent, abaxially the surfaces with scattered pectinate scales 0.8 mm long; idioblasts conspicuous on both blade surfaces; sori 3–9 per pinnule, along acroscopic and sometimes also along distal pinnule margins; indusia 1–4 mm long, oblong, with castaneous hairs 0.5–0.8 mm long.

Distribution. Terrestrial on wet slopes by streams, Pacific slopes; 100–1000 m. Mexico; Guat, Salv, Nic, CR. Guatemalan and Costa Rican distribution is attributable to Rojas (2001).

Selected Specimens Examined. Chis (Breedlove 32898, DS, NY, 49098, 49552, 57390, CAS; Martı´nez S. 10633, 13638, UAMIZ, 16955, XAL; Narave F. et al. 1243, XAL). Gro (Hinton 14678, NY). Oax (Mickel 1312, UC, 6975, NY, UC, 7015, NY; Morton & Makrinius 2335, US).

Adiantum trichochlaenum is distinctive among the shortpinnuled, idioblastic species by having numerous castaneous hairs on the indusia. Adiantum fructuosum Spreng. of Cuba (see further discussion under Excluded Species) has glabrous indusia. See A. villosum for an account of a putative hybrid involving A. trichochlaenum.

32. ADIANTUM TRICHOLEPIS Adiantum tricholepis Fe´e, Me´m. Foug. 8: 72. 1857. Based on Adiantum fragile Sw. var. pubescens M. Martens & Galeotti, Me´m. Foug. Mexique 72. 1842. Type. Mexico. [Jalisco:] “sur les rochers au bord du Rio Grande de Lerma pre`s de Guadalaxara,” Galeotti 6445 (BR!; isotypes BR!, P!, UC!, US! photo ex BR). Figs. 6C, D. Adiantum chilense Kaulf. var. pilosulum Liebm., Mexic. Bregn. 267 (reprint 115). 1849. Type. Mexico. Veracruz: Papantla, Liebmann s.n. [Pl. Mex. 2248, Fl. Mex. 440, 441] (C!; isotype US!).

Rhizomes short-creeping to ascending or suberect, ca. 4–6 mm diam.; rhizome scales orange-brown, becoming brown, 5–6 ⫻ 0.8–1.5 mm, lanceolate, apices each with an ephemeral, twisted hair, margins ciliate-denticulate, especially in distal half; fronds clumped, mostly 25–70 cm long, laxly arching; stipes castaneous to atropurpureous, 10–42 cm ⫻ 1–3 mm, ca. 1⁄3–3⁄5 the frond length, glabrous except at bases; blades ovate, 3–4-times pinnate, 15–45 ⫻ 15–35 cm, gradually tapering distally; rachises atropurpureous, glabrous; pinnae (4–)6–12 pairs, to 7–20 ⫻ 4–9 cm, alternate; pinnulets 4–15 ⫻ 4–18 mm, flabellate to almost round, margins entire to denticulate, sometimes shallowly incised, stalked 2–6 mm, stalk color stopping Ⳳ abruptly at bases of pinnulets, ultimately articulate; veins free, forking, ending at the entire margins or sometimes in tiny teeth in sterile pinnulets; indument of numerous stiff or curved acicular hyaline hairs 0.3– 0.5 mm long on both sides of pinnulets; idioblasts absent; sori 3–10 per pinnulet, without farina; indusia 1–7 mm long, oblong to slightly lunate, with hairs ca. 0.3 mm long.

ADIANTUM WILESIANUM

43

acroscopic and distal part of basiscopic margins; indusia 1–4 mm long, oblong or slightly arcuate, glabrous.

Distribution. Terrestrial on dry cliffs near streams, tropical deciduous forests; 50–1450 m. Mexico; Guat, Bel, Hond. Selected Specimens Examined. Camp (Sanders et al. 9819, UC, UCR). Chih (Pringle 962, NY, UC). Chis (Breedlove 29049, DS, NY). Coah (Wynd 691, MO, NY, US). Col (McVaugh & Koelz 1547, CAS, IEB, NY). Dgo (Gentry 5271, NY). Gro (Orcutt 2563, DS). Jal (Pringle 1843, NY, UC, US). Me´x (Matuda 31674, UC, US). Mich (del Llano 34, IEB). Mor (Lyonnet 762, CAS, MEXU, MO, NY). Nay (Te´llez 11631, ENCB). NL (Pringle 1981, B, BR, CAS, DS, MO, UC, US). Oax (Mickel 5192, NY, UC). Qro (Carranza 1218, ENCB, IEB). QR (Torres 10, MEXU, UAMIZ). SLP (Gonza´lez Quintero 120, ENCB). Sin (Mexia 308, CAS, MO). Son (Sanders et al. 9355, UC). Tam (Palmer 180, B, NY, UC). Ver (Purpus 6062, GH, MO, NY, UC). Yuc (Gaumer 386, B, BR, CAS, F, K, MEXU, MO, NY, UC, US).

Adiantum tricholepis is widespread and common in much of eastern and western Mexico, but is uncommon in Oaxaca and Chiapas. It is easily recognized by the presence of stiff, whitish hairs on both blade surfaces.

33. ADIANTUM VILLOSUM Adiantum villosum L., Syst. Nat., ed 10, 2: 1328. 1759. Type. Neotype (designated by Proctor, Fl. L. Antill. 2: 185. 1977). Jamaica. Near Spanish Town, Sloane Herb. 1: 127 (BM). Figs. 11L, M. Rhizomes short-creeping, 5–8 mm diam.; rhizome scales dark brown, denticulate, 1.5–2 ⫻ 0.1–0.3 mm; fronds clumped, 50– 100+ cm long, arching; stipes atropurpureous, lustrous, 40–55 cm ⫻ 2–4 mm, ca. 2⁄3 of the frond length, with numerous brown hair-like and lanceolate scales 1–2 mm long; blades ovate, 2pinnate, 25–45 ⫻ 25–40 cm wide, with conform terminal pinna; rachises atropurpureous, with lanceolate to hair-like, denticulate scales; pinnae 3–6 pairs, 15–25 ⫻ 4–6 cm, alternate; pinnules 20– 35 ⫻ 8–12 mm, sides of proximal and median pinnules forming right angle at pinnule bases, distal pinnules on each pinna about 1 ⁄2 the length of the longest pinnules, terminal pinnules hastate, non-articulate; veins free, forking, ending in small teeth (sterile pinnules); indument adaxially absent, abaxially the surfaces with scattered tan pectinate scales to 1 mm long; idioblasts conspicuous on both blade surfaces; sori 10–12 per pinnule, along both

Distribution. Terrestrial in wet forests, tropical rain forests, seasonal evergreen forests, wooded slopes along streams, lower montane rain forests; 0–800 m. Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Tob, Ec, Peru, Braz. Selected Specimens Examined. Camp (Zamora C. & Me´ndez D. 5247, XAL). Chis (Breedlove 32577, DS, NY; Fisher 35400, NY, US). Gro (Vela´zquez 616, FCME). Oax (Corte´s 6, ENCB, UAMIZ; Mickel 5773, NY, UC). QR (Torres et al. 66, UAMIZ). Tab (Matuda 3205, US). Ver (Nee 23614-b, XAL; Rzedowski 18890, ENCB, MEXU, NY).

The acute-tipped pinnules with right-angle bases distinguish this taxon among the bipinnate, idioblastic species with large pinnules and hair-like scales on the blades. Putative hybrids with A. trichochlaenum, having malformed spores and large pinnae with papillate emergences on the indusia, have been found in Chiapas (Purpus 6864, NY, UC, US). See A. pulverulentum for discussion of hybrids with that species. Specimens of A. villosum from the West Indies and some from Mesoamerica have the sori nearly continuous along the acroscopic pinna margins, while those from Mexico and most from Guatemala and Honduras have the sori interrupted into ca. 5– 7 units, each mostly 2–4 mm long.

34. ADIANTUM WILESIANUM Adiantum wilesianum Hook., Sp. Fil. 2: 50, t. 83C. 1851. Lectotype (chosen by Proctor, 1985: 245; relectotypified here). Jamaica. Cultivated at Kew, Wiles s.n. (K! isolectotype BM!, photo UC!). Proctor’s designation of the lectotype at BM is inappropriate because the Hooker herbarium is at Kew, and the type was cultivated at Kew. Figs. 8A–C. Rhizomes short-creeping, ca. 5 mm diam.; rhizome scales light brown with thickened, dark brown cell walls, margins denticulate, 1–1.5 ⫻ 0.8–1 mm; fronds clumped to subdistant, 70–120 cm long, arching; stipes atropurpureous to blackish, lustrous, to 60 cm ⫻ 3–6 mm, ca. 1⁄2 the frond length, glabrous except at bases; blades broadly ovate to deltate, 3–4-pinnate proximally,

44

ADIANTUM

with conform, 1-pinnate terminal pinna; rachises atropurpureous to blackish, short-puberulent abaxially with hairs 0.1 mm long; pinnae 3–5 pairs, alternate, stalked to 5 cm, distal pinnae 1pinnate, proximal 1–3 pinna pairs 2–3-pinnate, more distal pairs 1-pinnate; pinnulets dimidiate, subsessile or with stalks to 0.5 mm long, oblong to narrowly trapezoidal, non-articulate, acroscopic margins incised-lobate, minutely denticulate, basiscopic margins entire, concave proximally, toothed distally; veins free, forking, ending in teeth or crenulations; indument absent on both blade surfaces, the costae, costules, and minor axes abaxially with short hairs 0.1 mm long; idioblasts absent; sori 4–6 per pinnulet, confined to acroscopic margins; indusia 1–3 mm long, oblong to lunate, glabrous, doubly recurved (forming pouches); 2n⫽90 (cult).

Distribution. Terrestrial in low elevation wet forests, evergreen rain forests, tropical rain forests, seasonal evergreen forests, on slopes and along small streams; 50–1200 m. Mexico; Guat, Bel, Hond, Nic. Selected Specimens Examined. Chis (Breedlove 57969, CAS, ENCB; Martı´nez S. 6995, ENCB). Hgo (Turra 2975, ENCB). Oax (Herna´ndez G. 735, CAS, ENCB, MO, 892, CAS, ENCB, MO, NY). SLP (Kenoyer A7, MO, US). Tab (Cowan 3154, ENCB, NY; Ventura A. 20416, ENCB, MO). Ver (Copeland herb. 119, B, GH, MEXU, MICH, NY, UC, US; Rzedowski 19088, ENCB, NY).

The short, stiff hairs on the abaxial side of the rachises distinguishes A. wilesianum from all other Mexican Adiantums. The species is very similar to A. polyphyllum Willd. from northern South America and Trinidad, but that species has the axes glabrous abaxially and the color of the stalks stops abruptly at the bases of the pinnulets.

35. ADIANTUM WILSONII Adiantum wilsonii Hook., Sp. Fil. 2: 6, t. 72A. 1851. Type. Jamaica. Near Bath, Wilson s.n. (K–3 sheets, photos UC!, US). Figs. 12A–D. Rhizomes short- to long-creeping, 2–3 mm diam.; rhizome scales brown, concolorous, margins entire to minutely denticu-

late; fronds 5–10 mm apart, mostly 20–50 cm long, arching; stipes atropurpureous to blackish, 15–31 cm ⫻ 1.5–2.5 mm diam., sparsely filiform-scaly to hairy, often glabrescent; blades broadly ovate to deltate, 1-pinnate, each with a subsimilar but often slightly broader terminal pinna (juvenile fronds sometimes cordate); rachises atropurpureous, sparsely to moderately filiformscaly and hairy; pinnae 1–3(–6) pairs, alternate, entire (sterile serrulate), ovate-lanceolate, rounded or subcordate at bases, acute at tips, ca. 6–10 ⫻ 2.5–4 cm, costae medial, distinct, stalks 2–6 mm long, stalk color passing into the pinna bases, nonarticulate; veins forking, anastomosing toward pinna margins, ending in small teeth; indument absent adaxially, abaxially of scattered hair-like scales and hairs on the costae and laminar tissue near the costae; idioblasts present; sori continuous along the acroscopic and basiscopic pinna margins, pinna apices sterile; indusia to 12 cm long, linear, glabrous.

Distribution. Terrestrial in lowland and lower montane forests; 0–800 m. Mexico; Guat, Bel, Hond, Nic, CR, Pan; Gr Ant; w Col, Ec. Selected Specimens Examined. Chis (Breedlove 34185, DS, 34532, DS, MO, 57978, CAS; Martı´nez S. 14710, 15645, MEXU, XAL; Matuda 3616, F, K, LL, MEXU, MO, NY, US). Oax (Herna´ndez G. 2315, NY). Tab (Go´mez-Pompa 703, CAS, MEXU). Ver (Riba 803, K, MEXU, MO, XAL, 840, UAMIZ).

This was treated by Smith (1981) under A. dolosum Kunze [⫽ A. asperum (Poir.) Desv.], a very closely related species from the Guianas. Adiantum asperum differs primarily by the greater number of narrower lateral pinnae and the dark color of the pinna stalks not entering the costae abaxially. In Mexico, A. wilsonii is most similar to A. macrophyllum, but the resemblances are not indicative of a close relationship. Rather, A. wilsonii is related to several anastomosing-veined South American species, including A. asperum, A. phyllitidis J. Sm., from the Guianas to Colombia, and A. poeppigianum (Kuhn) Hieron., from Peru. This group comprises the segregate genus Hewardia. Excluded Species Adiantum fructuosum Kunze ex Spreng., Syst. Veg. 4: 113. 1827. Adiantum tetraphyllum var. majus E. Fourn., Mexic. Pl. 1: 129. 1872. (a renaming of A. fructuosum Kunze, and so based on

ALSOPHILA

the type of that name; see Lellinger, Proc. Biol. Soc. Wash. 89: 704. 1977). Type. Cuba. Poeppig [209] (isotypes B–photo UC!, L–photo UC!, US!– photo UC!, frag. NY!). This is very closely related to and possibly conspecific with A. tetraphyllum, with which it has been combined by several authors (e.g., Stolze, 1981; R. Tryon & Stolze, 1989). Other close relatives are A. amblyopteridium, A. oaxacanum, and A. trichochlaenum. Most likely, hybridization is occurring among species of this group of Adiantum, wherever they occur together. Mexico has been included in the range of A. fructuosum by many authors (e.g., Smith, 1981; Palacios-Rios, 1992). Aside from problems distinguishing it from A. tetraphyllum, there are questions whether Mexican material is really conspecific with the Cuban type. We now believe that most or all specimens so named from Mexico are other species. For example, the specimens cited by Smith (1981) are either A. trichochlaenum (e.g., Breedlove 38511, DS) or A. tetraphyllum (e.g., Breedlove 22184, 22276, 26354A, 31511, 38582, 39999, DS). Specimens cited from Veracruz by Palacios Rios (1992) seem to be mostly, if not entirely, A. tetraphyllum (e.g., Purpus 2930, UC; this number, in different herbaria, cited as both species by Palacios Rios). Adiantum fructuosum has been attributed a wide range by those who have dealt with it, from Mexico, Guatemala, Belize, Nicaragua, Costa Rica, Panama, Greater Antilles, Colombia, Venezuela, Trinidad, Guianas, Ecuador, Peru, Brazil, and Bolivia (see Lellinger, 1989; Jermy in Davidse et al., 1995). Specimens seen from Guatemala, Nicaragua, Costa Rica, and Panama seem to match the Cuban type well.

45

6 . A L SO P H I L A Alsophila R. Br., Prodr. 158. 1810. Type: Alsophila australis R. Br. (additional synonymy in R. Tryon, 1970). Nephelea R. M. Tryon, Contr. Gray Herb. 200: 37. 1970. Type: Nephelea polystichoides (Christ) R. M. Tryon [⬅ Cyathea polystichoides Christ].

Terrestrial; stems decumbent to erect, to 15 m, with or without spines; fronds to 4.2 m, short-sessile to long-stipitate; stipes brown to black, smooth or spiny; stipe base scales concolorous or bicolorous, marginate, with an apical seta or with apical and lateral setae, also with squamules; blades pinnate-pinnatifid to quadripinnate; rachises with hairs, scales and squamules; veins free, mostly simple; indusia globose (sphaeropteroid) or absent; sporangia 16-spored, 64-spored in A. salvinii; x⫽69. Alsophila is a genus of ca. 235 species in the New and Old World tropics, three in Mexico. Recent treatments (e.g., Riba in Davidse et al., 1995) have submerged Nephelea under Alsophila. The genus is distinct from other tree ferns by its frond scales bearing black acicular setae. References Conant, D. S. 1983. A revision of the genus Alsophila (Cyatheaceae) in the Americas. J. Arnold Arbor. 64: 333–382; Conant, D. S., L. A. Raubeson, D. K. Attwood & D. B. Stein. 1995 [1996]. The relationships of Papuasian Cyatheaceae to New World tree ferns. Amer. Fern J. 85: 328–340; Conant, D. S., L. A. Raubeson, D. K. Attwood, S. Perera, E. A. Zimmer, J. A. Sweere & D. B. Stein. 1996. Phylogenetic and evolutionary implications of combined analysis of DNA and morphology in the Cyatheaceae. Pp. 231–248 in J. M. Camus, M. Gibby & R. J. Johns (eds.), Pteridology in Perspective. Royal Botanic Gardens, Kew; Conant, D. S. & B. D. Stein. 1999. Floristic relationships in Southeast Asia based on DNA and morphological data from Cyathea (Alsophila), with emphasis on diversification of the genus on Mount Kinabalu and in New Guinea. XVI International Botanical Congress, Abstracts p. 73; Gastony, G. J. 1973. A revision of the fern genus Nephelea. Contr. Gray Herb. 203: 81–148.

Key to the Mexican Species of Alsophila 1. Stipes with 6–10 pairs of reduced, skeletonized pinnae (aphlebiae); stipes tuberculate, lacking spines; unexpanded crosiers unarmed; indusia absent; apical setae of stipe scales ephemeral, short; fronds 3-pinnate to 3-pinnate-pinnatifid; ultimate segments toothed and drying flat; rachises and costae atropurpureous to dark brown with regular marginate scales and bullate scales. ......... 2. A. salvinii. 1. Stipes lacking aphlebiae; stipes with dark, curved spines; unexpanded crosiers with stout black spines; indusia sphaeropteroid or absent; apical setae of stipe scales persistent, long; fronds 2-pinnate-pinnatifid; ultimate segments entire or subdentate with teeth often hidden by inrolled margins (when dry); rachises and costae tan to light brown. 2. Indusia sphaeropteroid; costae abaxially with small black-tipped scales but lacking bullate scales. .................................................... 1. A. firma. 2. Indusia absent, costae abaxially with bullate scales. .............................................................................................................................. 3. A. tryoniana.

Alsophila firma (Baker) D. S. Conant, J. Arnold Arbor. 64: 372. 1983. Figs. 13F–H, J-L.

Type. Mexico. [Veracruz:] Jalapa, Schiede & Deppe 802 (B!, frags. NY!, US!; isotypes BM, K!, LE!). Cyathea trejoi Christ, Bull. Herb. Boissier, se´r. 2, 5: 733. 1905. Type. Mexico. Chiapas: San Pablo, 1903, Mu¨nch s.n. (P?; frag. NY!). Cyathea articulata Fe´e, Me´m. Foug. 8: 111. 1857. Type. Mexico. Oaxaca: Villa Alta and Talea, Galeotti 6531 (P; isotypes B, frag. NY!, BR!–9 sheets).

Cyathea firma (Baker) Domin, Pterid. Dominica 264. 1929, non Mett. ex Kuhn, 1869. Hemitelia firma Baker, J. Bot. 15: 161. 1877. Type. Ecuador. Andes of Quito, Aug 1875, Sodiro s.n. (K; isotype P, frag. US). Cyathea mexicana Schltdl. & Cham., Linnaea 5: 616. 1830. Nephelea mexicana (Schltdl. & Cham.) R. M. Tryon, Contr. Gray Herb. 200: 40. 1970.

Trunks to 10.5 m tall, 30 cm diam.; stipe spines stout, to 12 mm long, black, curved; stipe scales very narrow, dark brown with pale erose margins and long dark persistent setae tip; blades to 3 m long, 2-pinnate-pinnatifid, very firm to subcoriaceous;

1. ALSOPHILA FIRMA

46

ALSOPHILA

pinnae stalked to 1 cm; veins mostly simple; pinnule apices acute, slightly ascending, crenulate, but margins often recurved; hairs on costules and segment midveins abaxially, costules also with small black-tipped scales, lacking bullate scales; segment midveins glabrous adaxially or with a single hair near the tip; costules hairy adaxially; sori medial; indusia globose, sphaeropteroid; 2n⫽138 (CR).

forked, basal lobes with veins 2-forked to pinnate; sori medial, naked; indusia absent.

Distribution. Wet montane forests of Atlantic slope; 900–2500 m. Mexico; Guat, Bel (Holst 5854, UC), Hond, Salv, Nic; Peru.

Distribution. Wet montane forests; 600–2500 m. Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Ec. Selected Specimens Examined. Chis (Breedlove 23321, NY, 32194, DS). Hgo (Arreguı´n 499, NY). Me´x (Hinton 3629, US). Oax (McVaugh 22455, MICH; Mickel 1089, NY). Pue (Arreguı´n 536, NY). Qro (Rubio 311, ENCB, IEB, MEXU). SLP (Mickel 581, NY). Ver (Ventura A. 4518, NY). Unverified, Doubtful, or Mistaken Reports. Gro (Lorea 2366, ENCB, cited by Lorea-Herna´ndez & Vela´zquez M., 1998, but not verified).

Among the indusiate species of Mexican Alsophila, A. firma can be distinguished most readily by the stout black spines on the stipes.

2. ALSOPHILA SALVINII Alsophila salvinii Hook. in Hooker & Baker, Syn. Fil. 36. 1866. Figs. 13A–E. Cyathea salvinii (Hook.) Domin, Pterid. Dominica 263. 1929. Type. Guatemala. Chilasco, Salvin & Godman s.n. (K!–2 sheets). Alsophila muenchii Christ, Bull. Herb. Boissier, se´r. 2, 5: 734. 1905 [“munchii”]. Type. Mexico. Chiapas: San Pablo, Mu¨nch 139 (P!, as “isotype”; isotypes DS!, US!).

Trunks 3–6 m tall; stipes atropurpureous, tuberculate, with 6– 10 pairs of reduced skeletonized pinnae (aphlebiae); stipe spines absent; stipe scales castaneous, 2–3 mm wide, with pale erose margins and short ephemeral apical setae, reduced squamulae intermixed; blades to 3.5 m long, 3-pinnate to 3-pinnatepinnatifid, pinnae subsessile to stalked 0.5–4 cm, pinnules sessile to short-stalked 3 mm long at bases; segments 1.2–2.4 cm wide, cut shallowly to halfway to the midvein, glabrous; rachises and costae atropurpureous to dark brown, with dark brown marginate scales and bullate scales, costae hairy adaxailly; veins 1–2-

Selected Specimens Examined. Chis (Breedlove 21465, DS, ENCB; Heath & Long 934, XAL; Matuda 38280, MEXU). Oax (Mickel 4318, MEXU, NY, 5580, NY, US; Rzedowski 33582, MEXU; Torres C. 9531, 11308, MEXU). Ver (Calzada 10961, XAL; Sousa 3604, MEXU).

Alsophila salvinii is distinct in its dark frond axes and the presence of reduced, skeletonized pinnae (aphlebiae). As old trunks fall, buds develop on the trunk, take root, and develop into independent trees. This species is most closely related to species in Madagascar and Borneo (Conant, 1983).

3. ALSOPHILA TRYONIANA Alsophila tryoniana (Gastony) D. S. Conant, J. Arnold Arbor. 64: 371. 1983. Figs. 13M–R. Nephelea tryoniana Gastony, Contr. Gray Herb. 203: 118. 1973. Type. Guatemala. Dept. Zacapa: canyons bordering Rı´o Lima, Sierra de las Minas, below Finca Alejandria, Steyermark 30009 (F–5 sheets; isotypes GH!, US!).

Trunks to 4.5 m tall; stipe spines ca. 1 cm, black; stipe scales castaneous, with pale margins and each with a dark persistent apical seta and occasionally dark lateral setae; blades to 1.7 m, 2pinnate-pinnatifid, rachises pale brown; pinnae to 65 cm, abaxial pinna rachis scales linear, whitish with black setae at apex and laterally; pinnules with whitish bullate scales with black apical setae, also usually marginally setulose, distally the scales bullate but without black setae, segments also with stiff hairs ca. 0.5 mm; sori medial; indusia absent.

ANANTHACORUS

47 References

Crane, E. H., D. R. Farrar & J. F. Wendel. 1995. Convergent simplification leads to a polyphyletic Vittaria. Amer. Fern J. 85: 283–305; Crane, E. H. 1997 [1998]. A revised circumscription of the genera of the fern family Vittariaceae. Syst. Bot. 22: 509–517; Maxon, W. R. 1908. Studies of tropical American ferns– No. 1. A new genus allied to Vittaria. Contr. U.S. Natl. Herb. 10: 486–487.

1. ANANTHACORUS ANGUSTIFOLIUS Ananthacorus angustifolius (Sw.) Underw. & Maxon in Maxon, Contr. U.S. Natl. Herb. 10: 487. 1908. Figs. 14A–D.

Distribution. Cloud forests and evergreen rain forests; 1100– 2500 m. Mexico; Guat, Hond, Salv, Nic. Selected Specimens Examined. Ver (Lira 140, 160, MEXU; Nee 24992, XAL; Ramirez 830, 1026, 1175, 1184, XAL; Riba et al. 1087, 1094, UAMIZ; Sousa 3604, MEXU).

Alsophila tryoniana is distinct by the exindusiate sori and bullate scales on the segment midveins abaxially. Conant (1983) has suggested that this species arose as a cross between A. firma and A. salvinii.

7. ANANTHACORUS

Pteris angustifolia Sw., Prodr. 129. 1788. Taenitis angustifolia (Sw.) Spreng., Syst. Veg., ed. 16, 4: 42. 1827. Vittaria angustifolia (Sw.) Baker in Martius, Fl. Bras. 1(2): 544. 1870, non Blume, 1828. Type. “Jamaica and Hispaniola,” probably collected by Swartz in Jamaica (Proctor, Fl. L. Antill. 2: 376. 1977). Vittaria costata Kunze, Linnaea 9: 77. 1834. Type. Peru. Poeppig s.n. (holotype LZ, destroyed; isotype W; drawing of probable holotype at B).

Rhizomes compact to short-creeping; roots with abundant soft hairs; rhizome scales linear to lanceolate, 2–6 ⫻ 0.3–0.7(–1) mm, 4–10 cells wide at the base; fronds 2–7 mm distant, with distinct stipes absent or very short, flattened, and narrowly alate; blades 15–45 ⫻ 0.9–1.5 cm at widest part, 1.5–2 mm wide at base, attenuate at both ends, slightly falcate, midribs evident, concolorous; vein areoles hexagonal or sometimes pentagonal; sporangia near outer edge of the ultimate series of areoles, 1–1.5 mm from the blade margins, sometimes interrupted; 2n⫽240 (Jam).

Ananthacorus Underw. & Maxon in Maxon, Contr. U.S. Natl. Herb. 10: 487. 1908. Type: Ananthacorus angustifolius (Sw.) Underw. & Maxon. [⬅ Pteris angustifolia Sw.] Epiphytic or epipetric; rhizomes short-creeping, dorsiventral, solenostelic, at apices with clathrate scales; fronds distichous, pendent or arching, clumped to somewhat remote, stipes absent or very short and flattened; blades linear-elliptic, entire, costate, glabrous; veins obscure, anastomosing and forming areoles in 2 (–4) series on each side of midrib (visible in transmitted light), areoles without included veinlets; sori of two submarginal Ⳳ continuous lines, one on each side of the midrib, sunken in grooves; indusia absent; paraphyses abundant, filiform, not enlarged at tips; spores bilateral, hyaline, smooth; gametophytes lacking gemmae; x⫽120. Ananthacorus is monotypic, the single species widespread from Mexico to Bolivia. It has often been subsumed within Vittaria, a genus shown by Crane (1997) to be polyphyletic. Ananthacorus appears to be most closely related to Scoliosorus and Vittaria s.str., according to phylogenetic reconstruction by Crane. It differs from Vittaria in having vein areoles in usually two or three (sometimes four or five) series between the midrib and blade margins and in the absence of gemmae on the gametophytes. From Scoliosorus, Ananthacorus differs in having only two submarginal linear sori per blade and in having filiform paraphyses that lack a globose apical cell.

Distribution. Epiphytic or epipetric in lower montane and montane forests, semi-deciduous tropical forests; 50–1500 m. Mexico; Guat, Bel, Hond, CR, Pan; Gr & L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol. Selected Specimens Examined. Chis (Breedlove 48289, CAS, ENCB; Martı´nez S. 7687, MEXU, NY). Col (Lott et al. 2978, UC, UCR). Gro (Lorea 2852, IEB, MEXU, FCME; Storer 112, UC, US). Jal (Iltis et al. 313, NY, WIS). Nay (McVaugh 19221, CAS, IEB, MICH, MO, NY). Oax (Conzatti et al. 3038, ENCB, MEXU, US; Mickel 6116, NY, UC). Ver (Purpus 6186, B, MO, NY, UC; Va´zquez T. et al. 2585, CHAPA, NY).

This is relatively common in rain forest areas of Guerrero, Oaxaca, Veracruz, and Chiapas, but uncommon or rare elsewhere in Mexico. It is superficially similar to some species of Radiovittaria, especially R. remota (Fe´e) E. H. Crane in tropical

48

ANEMIA

America, but the latter has distinct, darkened stipes, marginal commissural veins, and lacks hexagonal areoles. In blade shape, Ananthacorus is also similar to Polytaenium lineatum, which differs in having 4–6 linear sori per frond.

8. ANEMIA Anemia Sw., Syn. Fil. 6, 155. 1806. Type: Anemia phyllitidis (L.) Sw. [⬅ Osmunda phyllitidis L.]. Rhizomes short-creeping, horizontal or ascending, clothed with dark or orange hairs; fronds erect, rarely forming a flat rosette, 1–3-pinnate, generally anadromous, rarely catadromous, papyraceous to coriaceous; veins free or netted; fertile pinnae held erect, rarely horizontally or at an angle; sporangia restricted to the erect, dissected, lowermost pair of pinnae just below the sterile part of the blade (hemidimorphic), rarely fronds wholly dimorphic (holodimorphic) with sporangia borne on entire blade; sporangia in two rows on ultimate segments of fertile pinnae, sessile, subglobose to oval; annuli subapical; spores tetrahedral-globose, striate with parallel smooth or spiny ridges; x⫽38.

Anemia includes about 120 species, mostly in Latin America, ten in Africa, one in southern India. The genus is most abundant in Brazil (70 species), with a secondary center in Mexico (20 species). Anemia includes subg. Anemiorrhiza (calciphiles of the Caribbean basin), and the more widely distributed subg. Anemia. The latter includes subg. Coptophyllum, formerly treated as distinct, but molecular data (Skog et al., 2002) have shown it to be inseparable. Some species of Anemia hybridize freely and frequently. The genus Anemia is most closely related to the African genus Mohria and more distantly to Schizaea, Actinostachys, and Lygodium, together comprising the family Schizaeaceae (Skog, 2001; Wikstrom et al., 2002). References Mickel, J. T. 1962. Monographic study of the fern genus Anemia, subgenus Coptophyllum. Iowa State Coll. J. Sci. 36: 349–482; Mickel, J. T. 1967. The phylogenetic position of Anemia colimensis. Amer. J. Bot. 54: 432–437; Mickel, J. T. 1981. Revision of Anemia subgenus Anemiorrhiza (Schizaeaceae). Brittonia 33: 413–429; Mickel, J. T. 1982. The genus Anemia (Schizaeaceae) in Mexico. Brittonia 34: 388–413; Skog, J. E., E. A. Zimmer & J. T. Mickel. 2002. Additional support for two subgenera of Anemia (Schizaeaceae) from data for the chloroplast intergenic spacer region trnL-F and morphology. Amer. Fern J. 92: 119–130; Walker, T. G. 1962. The Anemia adiantifolia complex in Jamaica. New Phytol. 61: 291–298.

Key to the Mexican Species of Anemia 1. Fronds holodimorphic; plants small, 5–12(–20) cm tall; Yucatan Peninsula. ........................................................................................... 4. A. cicutaria. 1. Fronds hemidimorphic; plants small to large, 2.5–80 cm tall; if on Yucatan Peninsula, then over 20 cm tall. 2. Fronds pinnate-pinnatifid to tripinnate. 3. Fronds tripinnate to tripinnate-pinnatifid. 4. Fertile pinnae erect, well differentiated; segments oblanceolate, serrulate; blades anadromous. ..................................... 1. A. adiantifolia. 4. Fertile pinnae nearly horizontal, only slightly differentiated; segments rhomboid, entire; blades catadromous. ............ 5. A. colimensis. 3. Fronds pinnate-pinnatifid to bipinnate-pinnatifid. 5. Fertile pinnae spreading, held nearly horizontally or at an angle, not erect; fronds 2.5–16 cm tall. 6. Fronds 2.5–5.5 cm tall; Sinaloa. ............................................................................................................................................ 3. A. brandegeei. 6. Fronds 9–16 cm tall; Nayarit. ............................................................................................................................................... 8. A. intermedia. 5. Fertile pinnae fully erect; fronds (9–)20–60 cm tall. 7. Fronds bipinnate (to bipinnate-pinnatifid); segments ovate-obtuse, entire; blades catadromous; spore angles rounded. ............................................................................................................................................................................ 10. A. karwinskyana. 7. Fronds bipinnate-pinnatifid (uncommonly bipinnate); segments acute, toothed; blades anadromous; spore angles with slight protuberances. .............................................................................................................................. 20. A. tomentosa var. mexicana. 2. Fronds 1-pinnate. 8. Fertile pinnae remote from the sterile. 9. Pinnae 1–2(–4) pairs; Oax, Chis. .................................................................................................................................................. 19. A. speciosa. 9. Pinnae 5–9 pairs; Coah and Tam to Ver and Oax. .................................................................................................................. 11. A. mexicana. 8. Fertile pinnae approximate to the sterile. 10. Veins anastomosing, at least 10 anastomoses per pinna. 11. Anastomoses (50–)80–130 per pinna, uniform throughout, involving every vein; pinnae acuminate. ..................... 16. A. phyllitidis. 11. Anastomoses mostly 10–20 per pinna, often with some free veins; pinnae acute to obtuse. ...................................... 12. A. muenchii. 10. Veins free or with 1–2(–6) anastomoses per pinna in A. semihirsuta. 12. Pinnae not at all incised. 13. Blades oblanceolate to oblong, not tapering to apices; terminal pinnae broadly obdeltate; stipes of sterile fronds short, 1–3(–6) cm long; pinna margins entire to minutely denticulate; Sin to Chis, Me´x, and Ver. ............... 14. A. oblongifolia. 13. Blades narrowly oblong to narrowly deltate, tapering to pinnatifid apices or each with narrowly oblanceolate terminal pinnae; stipes of sterile fronds 6–15 cm long; pinna margins denticulate; Son, Sin, Dgo, Nay, Jal. ......... 2. A. affinis. 12. Pinnae (at least the proximal ones) at least slightly incised or lobed. 14. Pinnae deeply incised, hirsute (Anemia recondita cannot be distinguished in the key from its parents, A. hirsuta and A. jaliscana.). .................................................................................................................................................................. 7. A. hirsuta.

ANEMIA ADIANTIFOLIA

49

14. Pinnae shallowly incised to slightly lobed, hirsute to glabrous. 15. Blades narrowly deltate to oblong; proximal pinnae broadly and shallowly crenate; blade apices pinnatifid. 16. Pinnae acute; spores tuberculate; southern and western Mexico. ........................................................... 15. A. pastinacaria. 16. Pinnae broadly rounded; spores with smooth ridges, malformed; Nay, Jal, Col. .................................... 13. A. multiplex. 15. Blades narrowly oblong, not deltate; proximal pinnae subentire to incised; blade apices often with obdeltate terminal pinnae. 17. Pinnae broadly oblong, proximal pinnae sparingly incised, hirsute on both surfaces, often with 1–2(–6) vein anastomoses per pinna; spore ridges rugulose; Ver, Oax, Chis. ....................................................... 18. A. semihirsuta. 17. Pinnae rhomboid or narrowly oblong, denticulate to incised, hirsute to glabrous adaxially, glabrous or nearly so abaxially, the veins entirely free; spore ridges smooth. 18. Fronds (4–)14–38 cm long; pinnae hirsute adaxially, only very sparsely hairy abaxially; pinnae oblong to rhomboid-ovate, broadly cuneate at bases, entire to denticulate, rarely incised; central and western Mexico. .................................................................................................................................... 9. A. jaliscana. 18. Fronds 7–14 cm long; pinnae completely glabrous on both surfaces; pinnae narrowly oblong to oblanceolate, narrowly cuneate at bases, denticulate, rarely incised; Oax, Chis. .............................. 6. A. familiaris.

1. ANEMIA ADIANTIFOLIA Anemia adiantifolia (L.) Sw., Syn. Fil. 157. 1806. Figs. 15C, D. Osmunda adiantifolia L., Sp. Pl. 2: 1065. 1753. Ornithopteris adiantifolia (L.) Bernh., Neues J. Bot. 1(2): 40. 1806 [1805]. Lectotype (chosen by Proctor, Fl. L. Antill. 2: 53. 1977). Plumier, Traite´ Foug. Ame´r., pl. 158, 1705, based on material from Haiti, near Petit Goave. Osmunda asplenifolia Sav. in Lamarck, Encycl. 4: 652. 1797. Anemia asplenifolia (Sav.) Sw., Syn. Fil. 157. 1806. Anemia adiantifolia var. asplenifolia (Sav.) Hook. & Grev. Icon. Filic. 1: t.16. 1829. Type. Hispaniola. Martin s.n. (P; isotype B-Willd. 19499). Anemia carvifolia C. Presl, Reliq. Haenk. 1(1): 74. 1825. Type. Mexico. Haenke s.n. (PRC!). Anemia adiantifolia (L.) Sw. var. pumila Christ, Bull. Herb. Boissier, se´r. 2, 7: 794. 1907. Type. Puerto Rico. Sintenis 2194 (P!; isotype US!). Anemia adiantifolia (L.) Sw. var. subaurita Christ, Bull. Herb. Boissier, se´r. 2, 7: 794. 1907. Type. Puerto Rico. Utuado “in rupibus ad Los Angeles,” Sintenis 5948 (P!; isotype B!).

Rhizomes horizontal, creeping, ca. 2 mm diam.; rhizome hairs dark brown; fronds erect, 13–60 cm long, 1–20 mm apart; stipes 1 ⁄2–2⁄3 the frond length, 1–1.4 mm diam., stramineous to castaneous, hirsute to glabrous; blades deltate, 3-pinnate, 9–24 cm wide, coriaceous; pinnae 12–18 pairs, alternate to subopposite, segments oblanceolate, bases cuneate, apices obtuse, margins minutely denticulate, pilose with stiff white hairs; veins free; fertile pinnae remote from the sterile pinnae, usually shorter than the sterile blade; spores striate, ridges smooth; 2n⫽76, 152⫹ sterile triploid (Jam); 2n⫽76 (Col, SLP, Jam).

Distribution. Open to lightly shaded, rocky slopes, usually on limestone or calcareous soils, in mesic regions; 80–1720 m. USA (Fla); Mexico; Guat, Bel, Hond, CR, Pan; Bah, Gr Ant; Col, Ven, Trin, Ec, Braz. Selected Specimens Examined. Camp (Martı´nez S. 3017, MEXU). Chis (Stoutamire 3616, NY). Col (Mickel 1354, NY). Gro (Hinton 7258, NY). Hgo (Espinosa S. 222, MEXU). Jal (Mendoza 3685, MEXU). Me´x (Tejero-Dı´ez 2824, IZTA). Mich (Hinton 15909, NY). Mor (Pringle s.n., 29 Oct 1903, CAS, but at US is adiantifolia ⫻ mexicana var. mexicana; Lyonnet 1287, US). NL (Pringle 1977, NY). Oax (Conzatti 3791, NY). Pue (Tenorio L. 14062, MEXU). Qro (Servı´n 1233, IEB). QR (Cabrera 313, NY). SLP (Haynes 5675, NY). Tab (Cowan 3452, NY). Tam (Chase 7572, NY). Ver (Purpus 2848, NY). Yuc (Gaumer 372, NY).

This species and other members of subg. Anemiorrhiza in Mexico (A. cicutaria, A. colimensis, A. mexicana, and A. speciosa) are confined to calcareous rocks and soils. Anemia adiantifolia seems to be diploid in Mexico, but Walker (1962) found 2n, 3n, and 4n plants in Jamaica. In light of the varied numbers in several species of Anemia, other ploidy levels of A. adiantifolia might be expected in Mexico. This is one of the more common species of Anemia in Mexico. It is characterized by its fine dissection and coarse texture. It hybridizes with both varieties of A. mexicana: Anemia adiantifolia ⫻ mexicana var. mexicana (Fig. 15F) in Morelos, Nuevo Leo´n, Tamaulipas, and San Luis Potosı´; 2n⫽152 univalents (SLP); Anemia adiantifolia ⫻ mexicana var. makrinii (Fig. 15E)

50

ANEMIA

in Colima; n⫽76 univalents (Col). Anemia adiantifolia has also been found growing with A. colimensis (also subg. Anemiorrhiza), but no hybrids with that species have been found.

2. ANEMIA AFFINIS Anemia affinis Hook. & Baker, Syn. Fil., ed. 2, 525. 1874. Type. “North Mexico,” Sierra Madre [i.e., Sin, Dgo, or Nay], Seemann 1951 (K!). Figs. 19A–C. Rhizomes horizontal, compact, 2–4 mm diam.; rhizome hairs orange; fronds erect, 13–20 cm long; stipes 1⁄3–1⁄2 the frond length, 0.8–1.3 mm diam., stramineous, sparsely hirsute; blades oblong to narrowly deltate, 1-pinnate, 4–9 cm wide, each tapering to a pinnatifid apex or with narrowly obdeltate terminal pinna; pinnae 2–6 pairs, opposite to subopposite, oblong, dimidiate, bases truncate, apices obtuse, margins denticulate; blade surfaces very short-hairy adaxially, glabrous abaxially; veins free; fertile fronds to 35 cm long, stipe 2⁄3–3⁄4 the frond length; fertile pinnae approximate to the sterile pinnae, far surpassing the sterile blade in height; spores striate, ridges smooth; 2n⫽152 (Sin).

to be a fertile (or apomictic) hybrid with A. pastinacaria, judging from the frond form and chromosome number, 2n⫽ca. 380 (Sin). Gonza´lez Tamayo 368 (MICH) (Fig. 21E) from Jalisco is a very small fertile plant (7–10 cm) but consistent with A. affinis in being hairy adaxially, glabrous abaxially.

3. ANEMIA BRANDEGEEI Anemia brandegeei Davenp., Fern Bull. 13: 20, t. XVII, Fig. A. 1905. Figs. 21A–C. Hemianemia brandegeei (Davenp.) C. F. Reed, Bol. Soc. Brot., ser. 2, 21: 157. 1948. Type. Mexico. Sinaloa: vicinity of Culiaca´n, Cerro Colorado, 5 Nov 1904, Brandegee s.n. (GH!; isotypes UC!, US!).

Rhizomes horizontal, ca. 0.4 cm diam.; rhizome hairs orange; fronds clumped, 2.5–5.5 cm long; stipes ca. 1⁄2 the frond length, terete, less than 0.5 mm diam., stramineous, hirsute; blades deltate-ovate, pinnate-pinnatifid, chartaceous, 1.5–3 cm long; pinnae 3–5 pairs, lobes rounded, hirsute on both surfaces; fertile pinnae sessile, horizontal, remote from the sterile pinnae; spores 61–75 ␮m, the angle protuberances conspicuous, the ridges smooth, solid.

Distribution. Rocky, grassy slopes in open oak and pine woods; 350–1200 m. Mexico. Specimens Examined. Dgo (Ortega 4329, US). Jal (Gonza´lez T. 368, MICH). Nay (Feddema 794, IEB; Mason 3089, ARIZ; McVaugh 18694, NY). Sin (Gentry 6502, NY; Ownbey & Ownbey 1891, NY; Reveal 4048, NY). Son (Gentry et al. 19338, NY; Van Devender 93-1169, ASU).

Anemia affinis is most often confused with A. jaliscana, which it resembles in general blade dissection. Anemia jaliscana in Nayarit closely approaches A. affinis in the denticulate blade margins and size of fronds, but in A. affinis the proximal pinnae are never incised, and the pinnae are more rounded and more sharply denticulate. Anemia jaliscana is more hirsute, being short-hairy on both surfaces whereas A. affinis is glabrous abaxially. Gentry 6502 (NY) from Sinaloa consists of one frond of A. affinis, one frond of A. jaliscana, and two fronds of the presumed hybrid, with malformed spores. Anemia affinis rarely crosses with A. hirsuta in Nayarit (Keil & Canne 8962-1, NY), Jalisco (Schugun 16159, GUADA), and Sinaloa (2n⫽304) to form what appears to be a fertile hybrid (Mickel 1718, NY). Mickel 1719 (NY) appears

Distribution. Face of perpendicular cliffs. Known only from the type collection. The extremely small size of this plant is quite distinctive, but it is the only character we have found to separate it from A. intermedia. Other differences, such as degree of dissection, are probably only a reflection of the difference in size. Since the two species are known from a total of only three specimens, we hesitate combining them until their variation is better known.

4. ANEMIA CICUTARIA Anemia cicutaria Poepp. ex Spreng., Syst. Veg. 4: 31. 1827. Figs. 17C, D. Coptophyllum cicutarium (Poepp.) Underw., Mem. Torrey Bot. Club 2: 15. 1902. Type. Cuba. Ad Embarcadero del Canimar, Poeppig 837 (B!; isotype P!)

ANEMIA FAMILIARIS Anemia bipinnata T. Moore, Index Fil. 116. 1859, nom. nud. Mohria intermedia J. Sm., London J. Bot. 2: 387. 1843, nom. nud.

Rhizomes horizontal, short-creeping, ca. 1 mm diam.; rhizome hairs dark brown; fronds erect, holodimorphic, 5–12(–20) cm long, approximate; stipes 1⁄2–3⁄5 the frond length, 0.6 mm diam., stramineous, with sparse hairs to 1.5 mm; blades deltate, 2–3pinnate, 2–3(–8) cm wide, chartaceous; pinnae 3–6 pairs, alternate to subopposite, segments oblanceolate, bases cuneate, apices obtuse, margins denticulate; blade surfaces pilose with stiff hairs; veins free; fertile fronds far surpassing the sterile, often twice the length; spores reticulate.

51

blade surfaces with stiff unicellular hairs adaxially and multicellular hairs abaxially on veins; veins free; fertile pinnae remote from the sterile pinnae, slightly longer than the lowermost sterile pinnae and held in the same plane as the sterile portion of the blades, occasionally two pairs to wholly dimophic fronds; spores striate, the ridges broad, smooth, approximate to one another; 2n⫽76 (Col).

Distribution. Lightly shaded limestone or gypsum hills; 300– 1870 m. Mexico.

Distribution. Open to lightly shaded limestone, limestone crevices; near sea level. Mexico; Guat; Cuba, Bah. Specimens Examined. QR (Swallen 2877, US). Yuc (Baker 53104, DS; Schott 677, NY).

This species and A. wrightii Baker in Hooker & Baker, of Cuba, Bahamas, and Florida, form a distinct species group characterized by small, dimorphic, thin-textured fronds and spores with reticulate ridges. Anemia cicutaria is distinct by the hairiness and segment shape of its blades, the fertile fronds being more open, and the sterile fronds more dissected. The spore ridges are finely anastomosing, making the spores appear coarsely pitted rather than striate.

5. ANEMIA COLIMENSIS Anemia colimensis Mickel, Iowa State Coll. J. Sci. 36: 434. 1962 (for more complete description based on fertile material, see also Mickel, 1967). Type. Mexico. Colima: mountains 10 mi SSW of Colima on Manzanillo road, McVaugh 15537 (MICH!; isotype US!; original material entirely sterile; fertile topotype material, Mickel 1357, NY!). Figs. 22A–H. Rhizomes horizontal, short-creeping, 4–6 mm diam.; rhizome hairs orange to dark brown; fronds erect, 20–42 cm long; stipes 1 ⁄3–1⁄2 the frond length, 1–1.9 mm diam., stramineous, lightly hirsute; blades ovate-deltate, tripinnate to tripinnate-pinnatifid, 10–18 cm wide, mostly catadromous, subcoriaceous; pinnae 11– 14 pairs, opposite to subopposite, segments rhomboid to ovate, short-petiolate to narrowly adnate, apices obtuse, margins entire;

Selected Specimens Examined. Col (McVaugh 26194, CAS, IEB, MICH; Mickel 1357, 1692, NY; Pray 1940, LAM). Gro (Calonico Soto 1362, MEXU; Calzada 16358, FCME, NY). Oax (Calzada 18195, 18255, MEXU, NY, 20439, MEXU).

This species has the most unspecialized foliar morphology of any species in the genus with its only slightly differentiated basal pair of fertile pinnae held in the same plane as the rest of the frond. This may indicate a basal position of the species in Anemia. It grows in Colima on gypsum rocks with A. adiantifolia and A. mexicana var. makrinii, and recently has been discovered in eastern Guerrero and western Oaxaca. These recent collections are distinct in having uniformly two or more pairs of fertile pinnae, rather than one.

6. ANEMIA FAMILIARIS Anemia familiaris Mickel, Brittonia 34: 399. 1982. Type. Mexico. Oaxaca: Distrito Juchita´n, 14 km E of Tapanatepec on Rte 190, 6 km W of Chiapas border, Mickel 5984 (NY!). Fig. 18C. Rhizomes horizontal, compact, short-creeping, 5–6 mm diam.; rhizome hairs orange; fronds erect, 7–14 cm long; stipes 1⁄3–1⁄2 the frond length, 0.3–0.7 mm diam., stramineous, subglabrous to very lightly hairy; blades narrowly oblong, 1-pinnate, 2.5–3.2 cm wide, chartaceous; pinnae 3–5(–7) pairs, mostly opposite, cuneate-obovate to rhomboid or subflabellate, often excavate on basiscopic margin, 1.1–1.8 cm ⫻ 6–10 mm, eroseserrulate, glabrous; veins free; fertile fronds similar in size and form to sterile, or slightly longer-stiped; fertile pinnae approxi-

52

ANEMIA

mate to the sterile pinnae, shorter than or equal to the sterile blades in height; spores with smooth, narrow, spaced ridges.

laminae hirsute; veins free; fertile pinnae approximate to the sterile pinnae, usually far surpassing the sterile blades in height; spores striate, ridges smooth, distant; 2n⫽152 (Col, Gro, Jal, Me´x, Nay, Jam, Peru); 2n⫽76–86, 76–92, 88–98 (Oax); see below.

Distribution. Terrestrial, on shrubby stream banks and in forests; 150–400 m. Mexico. Specimens Examined. Chis (Breedlove 28271a, DS). Oax (Mickel 5969, NY).

This species is closely allied to A. hirsuta but has short fertile pinnae and only erose-serrulate, glabrous, sterile pinnae. These characters suggest an affinity to A. affinis of western Mexico, but the shorter fertile pinnae and small size of A. familiaris are distinctive. Anemia familiaris also resembles A. obovata (Underw.) Maxon of Cuba, which has glabrous, entire pinnae. However, the latter has only 2–4 pinna pairs that are broader, more obovate to oblong, and with pinna bases that are not as narrowly cuneate. An odd specimen from Jalisco (Gonza´lez 368, MICH) closely resembles A. familiaris, but is probably a dwarf specimen (7–10 cm) of A. affinis.

7. ANEMIA HIRSUTA Anemia hirsuta (L.) Sw., Syn. Fil. 156. 1806.

Fig. 18A.

Osmunda hirsuta L., Sp. Pl. 2: 1064. 1753. Ornithopteris hirsuta (L.) Bernh., Neues J. Bot. 1(2): 50, t. 3, f. 15b. 1806 [1805]. Lectotype (chosen by Proctor 1989: 47). Plumier, Traite´ Foug. Ame´r., pl. 162, 1705, based on a plant collected at Le Fond de Baudin, near Le´ogane, Haiti. Anemia hirsuta var. achillaeifolia M. Martens & Galeotti., Me´m. Foug. Mexique 20. 1842. Type. Mexico. Oaxaca: Distrito Juquila, Zacatepec & Juquila, Galeotti 6363 (BR!; isotypes BR!, GH!, LE!). Anemia opaca Fe´e, Me´m. Foug. 9: 41. 1857. Type. Mexico. Oaxaca: Talea, Galeotti 6567 (BR!–4 sheets). Several other synonyms and segregate taxa have been described from South America.

Rhizomes horizontal, compact, short-creeping, 2–6 mm diam.; rhizome hairs orange; fronds erect, (6–)12–32 cm long; stipes 1⁄3–2⁄3 the frond length, 0.5–0.9 mm diam., stramineous, hirsute; blades narrowly oblong, occasionally deltate, 1.8–5.6 cm wide, 1-pinnate, the proximal pinnae deeply incised with 1 to several segments, chartaceous; pinnae 6–10 pairs, opposite to subopposite, oblong, cuneate at bases, often excavate basiscopically, apices obtuse, margins minutely erose-denticulate,

Distribution. Open to mostly shaded grassy slopes and trail banks; 40–2000 m. Mexico; Guat, Salv, Hond, Nic, CR, Pan; Gr Ant; Col, Ven, Ec, Peru, Braz, Bol. Selected Specimens Examined. Chis (Breedlove 26938, NY). Col (McVaugh 16077A, MICH). Gro (Rzedowski 26865, NY). Jal (McVaugh 19843, NY). Me´x (Hinton 1640, NY). Mich (Hinton 15214, NY). Mor (Pringle 8792, NY). Nay (Feddema 698, MICH). Oax (Mickel 6263, NY). Pue (Sharp 45926, MEXU). Tam (Patterson 7300, TEX). Ver (Stoutamire 3611, NY).

Anemia hirsuta is one of the most common and widespread species in the genus. It hybridizes with several other species in Mexico, including A. affinis, A. jaliscana, A. karwinskyana, A. pastinacaria, A. phyllitidis, and A. tomentosa. This species is extremely plastic in its morphology, ranging from small to large, pinnae nearly undivided to extremely lacerate, and blades ranging from narrowly oblong to deltate. Some of its variants are indistinguishable externally from some of the hybrids, such as A. hirsuta ⫻ jaliscana. Such hybrids can be distinguished by their abortive or larger spores and by their intermediate dissection within the same population. Anemia hirsuta has a genome in common with A. jaliscana (38II ⫹ 114I in the hybrids). Mickel 6944 (NY) and Breedlove 37735 (DS, ENCB) have blades that are deeply dissected and exhibit significant spore malformation. Quite possibly there are different cytological races within A. hirsuta. Although most chromosome counts for A. hirsuta have been 2n⫽152, occasional counts deviate from the base of x⫽38 (see Mickel, 1982). The cause of these irregularities is not known, and further studies of A. hirsuta—morphological, cytological, and molecular—are needed not only in Mexico, but throughout this species’ range.

8. ANEMIA INTERMEDIA Anemia intermedia Copel. in M. E. Jones, Contr. W. Bot. 15: 123. 1929 (Pl. XVII, Fig. B). Fig. 21D.

ANEMIA JALISCANA

53

Hemianemia intermedia (Copel.) C. F. Reed, Bol. Soc. Brot., ser. 2, 21: 157. 1948. Type. Mexico. Nayarit: Acaponeta, El Tigre Mine, Jones 23472 (POM!; isotype UC!).

Distribution. Rocky, grassy slopes in pine or oak woods; 700– 2200 m. Mexico.

Rhizomes horizontal, less than 1 cm diam.; rhizome hairs orange; fronds 9–16 cm long, caespitose; stipes terete, 4–8 cm long, ca. 1 mm diam., light to dark brown, densely hirsute; blades deltate-ovate, bipinnate to bipinnate-pinnatifid, 5–12 ⫻ 4.5–6 cm, chartaceous; pinnae 7–10 pairs; pinnules narrowly adnate, oblong, crenate to pinnatifid, hirsute on both surfaces; veins free; fertile pinnae sessile, horizontal or slightly ascending, remote from the sterile pinnae; spores 56–81 ␮m, the angle protuberances conspicuous, the ridges smooth, solid.

Key to the Mexican Varieties of Anemia jaliscana 1. Pinnae incised to denticulate; terminal pinnae with base angle of 20–40(–50)⬚; Gro to Son & BCS. .............. 9a. var. jaliscana. 1. Pinnae subentire; terminal pinnae with base angle of 50– 85⬚; Me´x, w. Gro. ....................................................... 9b. var. integrifolia.

9a. Anemia jaliscana var. jaliscana

Fig. 19F.

Distinct from var. integrifolia by its incised to denticulate pinnae margins and narrowly cuneate bases of terminal pinnae; 2n⫽228 (Jal, Nay).

Distribution. Oak forests in humid canyon; 780 m. Mexico. Specimens Examined. Nay (Te´llez 12036, GH, MEXU).

Anemia intermedia is intermediate between A. brandegeei of Sinaloa and A. clinata Mickel (Panama to Bolivia) in size, the medium brown stipe color, and slightly ascending fertile pinnae. If the three species are distinct, A. intermedia may either represent an intermediate stage in the divergent evolution of the group or be of hybrid origin from the other two species. Quite possibly A. brandegeei is a diminutive form of A. intermedia, and the former has priority.

9. ANEMIA JALISCANA Anemia jaliscana Maxon, N. Amer. Fl. 16: 44. 1909. Type. Mexico. Jalisco: near Guadalajara, Pringle 3850 (US!; isotypes BR p.p.!, DS!, GH!, MEXU!, NY!). Rhizomes horizontal, compact, 2–5 mm diam.; rhizome hairs orange; sterile fronds erect, (4–)14–38 cm long; stipes 1⁄4–1⁄2 the frond length, 0.5-1.4 mm diam., stramineous, sparsely hirsute; blades oblong to narrowly deltate, 1-pinnate, 3–5 cm wide, thin; pinnae 5–8 pairs, opposite or subopposite, oblong to rhomboidovate, bases truncate, apices obtuse, margins irregularly incised to crenately lobed, denticulate; veins free; blade surfaces shorthairy adaxially, hairs sparse abaxially; fertile fronds to 28 cm tall, stipe 2⁄3 to 3⁄4 of the frond length; fertile pinnae approximate to the sterile pinnae, far surpassing the sterile blades in height; spores striate, ridges smooth.

Distribution. Among rocks on grassy slopes in open pine or oak woods; 700–1700 m. Mexico. Selected Specimens Examined. BCS (Carter 3484, MEXU, UC). Col (Reko 4849, US). Dgo (Ibana Garcı´a 797, US). Gro (Lorea 2571, NY). Jal (Pringle 1833, NY). Me´x (Mickel 712a, NY). Mich (Hinton 12212, NY). Nay (McVaugh 16416, NY). Sin (Gonza´lez Ortega 1504, MEXU). Son (Tenorio L. 10245, MEXU).

9b. Anemia jaliscana var. integrifolia Mickel, var. nov. Type. Mexico. Me´xico: Dist. Temascaltepec, Tejupilco, 1340 m, 30 Sep 1932, Hinton 1935 (holotype NY!; isotype GH!). Fig. 21F. Ab Anemia jaliscana Maxon var. jaliscana pinnis subintegris pinnisque apicalibus late cuneatis diversa. (L, integer, entire, ⫹ folium, leaf, alluding to the pinna margins being more nearly entire than those of the type variety.).

Distinct from var. jaliscana by its subentire pinna margins and broadly cuneate bases of terminal pinnae; 2n⫽228 (Gro, Me´x).

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ANEMIA Anemia rosei Maxon, N. Amer. Fl. 16: 46. 1909. Type. Mexico. Jalisco: Cerro de San Esteban, Pringle 11254 (US!; isotype NY!).

Rhizomes horizontal, compact, short-creeping, 6–9 mm diam.; rhizome hairs orange; fronds erect, (14–)20–60 cm long; stipes 1⁄2–3⁄4 the frond length, 1–1.5 mm diam., stramineous, hirsute; blades deltate to deltate-ovate or narrowly oblong, 2pinnate, rarely bipinnate-pinnatifid, 6–15 cm wide, subcoriaceous, mostly catadromous; pinnae 8–12 pairs, subopposite to alternate, pinnules narrowly adnate, subcordate at base, segments ovate-obtuse, lobes rounded, hirsute; veins free; fertile pinnae remote from the sterile pinnae, shorter than or usually equal to the sterile blades in height; spores striate, ridges subverruculate. Distribution. Rocky hillsides in pine or oak woods; 1120–2200 m. Mexico. Specimens Examined. Gro (Hinton 9472, GH, NY; Lorea 801, 2552, NY). Me´x (Hinton 8221, GH, NY; Matuda 31340, MEXU; Mickel 712a, NY).

Hinton 8221 (ARIZ) with malformed spores is mixed with A. hirsuta and is probably the hybrid between that and var. integrifolia, thus representing a counterpart to A. recognita, the sometimes fertile hybrid between A. hirsuta and A. jaliscana var. jaliscana. The range of var. integrifolia overlaps that of var. jaliscana in western Guerrero. Anemia jaliscana is perhaps the most difficult species of Anemia to distinguish in Mexico. The pinnae are somewhat trapezoidal in the most distinctive form, but vary to more elliptic or suborbicular. Occasionally, the proximal pinnae are slightly incised. The pinna margins range from entire to finely denticulate, even on the same plant; pinnae from sterile fronds tend to be more nearly entire than those from fertile ones. When pinna margins are incised, A. jaliscana may look more like A. hirsuta, with which it frequently hybridizes to form A. recondita. Hybrids are often mistaken for A. hirsuta or abnormal A. jaliscana. In fact, one of the NY isotype sheets of A. jaliscana has one plant of A. jaliscana and two of A. recondita. The two species (A. hirsuta and A. jaliscana) apparently share a common ancestor since there is regular pairing of approximately one genome (n⫽ca. 38II ⫹ 114I). In the northern part of its range, especially in Sinaloa, plants of A. jaliscana may look much like A. affinis in having larger, more denticulate pinnae, more broadly oblong than in plants farther south. However, A. affinis can generally be distinguished by the pinnae being suborbicular and glabrous beneath, and being tetraploid (vs. hexaploid).

10. ANEMIA KARWINSKYANA Anemia karwinskyana (C. Presl) Prantl, Unters. Morph. Gefa¨sskrypt. 2: 9. 1881. Figs. 16A, B. Anemia villosa Humb. & Bonpl. ex Willd. ζ?[sic] karwinskyana C. Presl, Suppl. Tent. Pterid. 83. 1845. Type. Mexico. “Christo,” Karwinski “exs. n. 8” (B!, frag. NY!; isotype US!).

Distribution. Rare on moist banks in pine-oak woods; 600– 1900 m. Mexico. Selected Specimens Examined. Gro (Hinton 11338, NY). Hgo (Durata V. s.n., May 1978, FCME). Jal (Mickel 1368, NY). Me´x (Mickel 710, NY; Rzedowski 20706, NY). Mich (King & Soderstrom 4863, NY). Mor (Lyonnet [520900005], MEXU). Nay (McVaugh 16509, NY). Oax (Conzatti 4369, US; Mickel 3993, NY). Unverified, Doubtful, or Mistaken Reports. Hond (Meyer 9924, MO, cited by Moran & Mickel in Davidse et al., 1995, is A. guatemalensis Maxon).

This species is recognized by its catadromous bipinnate fronds and rounded segment tips. It appears uniform morphologically, but apparently has cytological races. In Jalisco, it is presumably diploid, tetraploid, and hexaploid, judging from its usual 2n⫽76II and two sterile variants with 38II⫹38I and 38II⫹114I. Spore size is quite variable and may be correlated with different ploidy levels. Whereas most specimens have average spore diameters of 76–84 ␮m, perhaps representing the tetraploid, one specimen averaged only 70.1 ␮m (possibly the diploid) and three had spores averaging 87–93 ␮m (possibly the hexaploid). Anemia karwinskyana also crosses with A. hirsuta (meiotic count shows 152 univalents) in Jalisco.

11. ANEMIA MEXICANA Anemia mexicana Klotzsch, Linnaea 18: 526. 1844.

ANEMIA MUENCHII Ornithopteris mexicana (Klotzsch) Underw., Native Ferns, ed. 6, 76. 1900. Type. Mexico. Aschenborn 575 (B!).

Rhizomes horizontal, creeping, ca. 2 mm diam.; rhizome hairs stiff, dark brown; fronds erect, (15–)30–60 cm long; stipes ca. 1⁄2 the frond length, ca. 1 mm diam., stramineous, glabrate; blades deltate-lanceolate, 1-pinnate, 9–18 cm wide, subcoriaceous; pinnae 5–9 pairs, lanceolate-deltate to narrowly deltate to rounded or cuneate at bases, often slightly excavate basiscopically, apices acuminate, margins minutely serrulate; veins free; blade surfaces hirsute with minute white hairs abaxially, adaxially glabrous, green or bluish green and iridescent; fertile pinnae remote from the sterile pinnae, surpassing or occasionally equal to the sterile blades; spores striate, ridges smooth, close.

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Anemia makrinii Maxon, J. Wash. Acad. Sci. 8: 199. 1918. Type. Mexico. Oaxaca: Distrito Pochutla, Cafetal Nueva Esperanza, Conzatti, Reko & Makrinius 3087 (US!; isotype NY!).

Distinct by its narrowly deltate pinnae, 1–1.7 cm wide; pinna bases cuneate to rounded or truncate; laminae blue-green adaxially; 2n⫽76 (Col).

Distribution. Lightly shaded limestone outcrops; 70–1900 m. USA (Tex); Mexico. Key to the Mexican Varieties of Anemia mexicana 1. Pinnae lanceolate-deltate, 1.8–3 cm wide; pinna bases truncate, green adaxially; n., c. and e. Mexico. ..... 11a. var. mexicana. 1. Pinnae narrowly deltate, 1–1.7 cm wide; pinna bases cuneate to rounded or truncate; blue-green adaxially; sw. Mexico. ......................................................................... 11b. var. makrinii.

11a. Anemia mexicana var. mexicana

Fig. 17E.

Distinct by its pinnae lanceolate-deltate, 1.8–3 cm wide; pinna bases truncate, green above; 2n⫽76 (SLP).

Distribution. Rare on lightly shaded limestone outcrops at lower elevations near Pacific Coast; 400–1100 m. Mexico. Specimens Examined. Col (McVaugh 18059, NY). Gro (McVaugh 22168, MICH). Mich (Guerrero C. 1230, XAL; Koch & Fryxell 83243, NY). Oax (Makrinius 51, GH, NY). Unverified, Doubtful, or Mistaken Reports. Tab (reported by Magan˜a, 1992, but is probably var. mexicana). Bel (Davidse & Brandt 32127, MO, cited by Moran & Mickel in Davidse et al., 1995, is A. speciosa).

Var. makrinii is distinct from var. mexicana, which is found more to the north and east, by having pinnae that are narrower, cuneate at bases rather than truncate, and iridescent bluish green adaxially. Both varieties are diploid and both cross with A. adiantifolia (var. makrinii in Colima, var. mexicana in San Luis Potosı´) with the hybrids having 78I in meiosis.

12. ANEMIA MUENCHII Distribution. Lightly shaded limestone outcrops; 70–1900 m. USA (Tex); Mexico. Selected Specimens Examined. Coah (Wynd & Mueller 229, NY). Gro (Orcutt 4165, DS). Gto (Ventura & Lo´pez 7728, IEB, MEXU). Hgo (Clark 6891, NY). Me´x (Matuda 37304, MEXU). Mor (Pringle 8695, NY). NL (Pringle 1976, NY). Qro (Rzedowski 27777, NY). SLP (King 3913, NY). Tam (Mickel 579, NY). Ver (Duran E. et al. 279, MEXU). Unverified or Mistaken Reports. Tab (reported by Magan˜a, 1992, as var. makrinii, but from the photo and range it appears to be var. mexicana).

Guerrero material is mostly intermediate between the two varieties. 11b. Anemia mexicana var. makrinii (Maxon) Mickel, Brittonia 33: 421. 1981. Figs. 17A, B.

Anemia muenchii Christ, Bull. Herb. Boissier, se´r 2, 7: 792. 1907 [“munchii”]. Type. Mexico. Chiapas: San Cristo´bal, Mu¨nch 30 (P!; isotype DS!).Figs. 15A, B. Rhizomes horizontal to somewhat ascending, 3–5 mm diam.; rhizome hairs reddish brown; fronds erect, 7–23 cm tall; stipes 1⁄3– 1 ⁄2 the frond length, 1–1.5 mm diam., stramineous, pilose; blades deltate-oblong, 1-pinnate, 6–8 cm wide, chartaceous, glabrous; pinnae (2–)4–7 pairs, opposite to subopposite, proximal pinnae short-petiolate 1–2 mm, distal pinnae narrowly adnate, elliptic to ovate, rounded at bases, slightly excavate basiscopically, apices obtuse to acute, margins entire to minutely serrulate, glabrous; veins casually anastomosing, 10–20 anastomoses per pinna; fertile fronds taller (11–37 cm long) and longer-stiped (1⁄2–2⁄3 of the frond length) than the sterile fronds; fertile pinnae approximate to the sterile blade, far exceeding the sterile blades in height;

56

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spores with few narrow ridges bearing short, blunt, peg-like spines; 2n⫽ca. 228 (Oax).

bases mostly narrowly cuneate, apices obtuse, minutely denticulate to entire or somewhat crenulate, the proximal pinnae often slightly incised, 2.5–3 times as long as broad; blade surfaces glabrous abaxially, short-hirsute adaxially; veins free; fertile fronds taller and longer-stiped than the sterile; fertile pinnae approximate to the sterile blade, far exceeding the sterile blades in height; spores ca. 90 ␮m diam., with broad smooth ridges, furrows between ridges narrow, many malformed; 2n⫽532 (Jal, Nay).

Distribution. Locally common on moist wooded slopes by streams at low elevations; 200 m. Mexico. Specimens Examined. Chis (only the type). Oax (Antonio B. & Heinrich GUI 232, MEXU; Mickel 6874, NY). Ver (Beaman 5237, MEXU p.p.; Calzada 8109, XAL; Copeland herb. s.n., 9 Mar 1938, MICH; Medina A. & Va´zquez E. 496, MEXU, XAL; Purpus 2229, GH, NY, UC, US).

This species is much like A. phyllitidis in having netted veins and spiny spores, but the veins are mostly free (only 10–20 anastomoses per pinna, especially along the margins), and the pinnae are more rounded and somewhat adnate toward the blade apices. A clear chromosome count has not been made, the best one showing 90–100 bivalents at meiosis. If the chromosome number for A. muenchii is indeed n⫽114, it is different from the number known for A. phyllitidis (2n⫽152, and presumably also 2n⫽76). The type material of A. muenchii agrees with Mickel 6847 (Oaxaca) in the number of anastomoses per pinna and general form, but the pinnae are more numerous and smaller. Unfortunately, the fertile pinnae of the type had been cut off just below the sporangia, so the spores are unknown. Anemia muenchii more closely resembles A. ⫻paraphyllitidis in the greater number and smaller size of the pinnae, but the latter has malformed spores, lanceolate pinnae, and generally 25–32 anastomoses per pinna.

13. ANEMIA MULTIPLEX Anemia multiplex Mickel, Brittonia 34: 405. 1982. Type. Mexico. Nayarit: road from Tepic to Compostela, just N of Km 25, Mickel 1709 (NY!). Fig. 19E. Rhizomes horizontal, compact, short-creeping, ca. 3 mm diam.; rhizome hairs orange; fronds erect, sterile ones to 23 cm long, fertile to 33 cm long; stipes stramineous, subglabrous to lightly hairy, 1⁄3–1⁄2 the sterile frond length, 1⁄2–3⁄4 of the fertile frond length, 0.8–2 mm diam.; blades narrowly deltate, 1pinnate, 3.6–5.6 cm wide, chartaceous; pinnae 7–11 pairs, oblong,

Distribution. Disturbed, lightly wooded, grassy banks and pastures; 900–1400 m. Mexico. Specimens Examined. Col (McVaugh 16077, MICH). Jal (Cochrane et al. 13052, NY; Mickel 1362, 1701, NY). Nay (Mickel 1384, 1713, NY).

This species is remarkable in that it is tetrakaidecaploid, 2n⫽532. There is considerable spore malformation, but there are well formed spores as well. In some squashes there appear to be some univalents. Anemia multiplex is most easily confused with A. jaliscana, which can be distinguished by the well formed spores (seen under a dissecting scope) and narrow, widely set spore ridges. Anemia multiplex is distinct from A. oblongifolia in its larger size, deltate blades with narrow apices, longer pinnae, and longer stipes. It is also similar to A. pastinacaria in the deltate blades and long pinnae, but the pinnae are more obtuse, the plants are larger, the spores have broad ridges, and the chromosome number is high with at least partial spore abortion. There is considerable variation in the size of the plants, even within the same population, but the general form, spore morphology, and chromosome number seem to be the same.

14. ANEMIA OBLONGIFOLIA Anemia oblongifolia (Cav.) Sw., Syn. Fil. 156. 1806. Fig. 18B. Osmunda oblongifolia Cav., Icon. 6: 69, t. 592, f. 2. 1801. Type. Panama. Ne´e s.n. (MA). Osmunda humilis Cav., Icon. 6: 69, t. 592, f. 3. 1801. Anemia humilis (Cav.) Sw., Syn. Fil. 156. 1806. Anemia oblongifolia (Cav.) Sw. β humilis (Cav.) Hook. & Baker, Syn. Fil., ed. 2, 431. 1874. Type. “Panama.” Taboga, Ne´e s.n. (MA).

ANEMIA PASTINACARIA Anemia pilosa M. Martens & Galeotti, Me´m. Foug. Mexique 19, pl. 2, f. l. 1842. Anemia pilosa M. Martens & Galeotti α brevistipes Liebm., Mexic. Bregn. 301 (reprint 149). 1849. Type. Mexico. Oaxaca: Zacatepec, Galeotti 6353 (BR!; isotypes B, BR!, frag. NY!, photo NY!, LE!). Anemia cornea Prantl, Unters. Morph. Gefa¨sskrypt. 2: 104. 1881. Type. Mexico. Oaxaca: Trapiche de la Concepcio´n, Liebmann s.n. (C! p.p.; isotype US!). Anemia seemannii Hook., London J. Bot. 7: 564, t. 16. 1848. Type. “Taboga, near Panama.” Seemann s.n. (K!, frag. NY!).

Rhizomes horizontal, compact, short-creeping, 5–10 mm diam.; rhizome hairs orange; sterile fronds spreading to prostrate, somewhat rosette-like, 2.5–15 cm long; stipes generally less than 1 cm long, 0.3–1.5 mm diam., stramineous, hirsute; blades narrowly oblong to narrowly oblanceolate, 1-pinnate, 2–3.8 cm wide, papyraceous to coriaceous; pinnae (3–)6–10 pairs, opposite to subopposite, oblong, dimidiate, cuneate at bases, apices obtuse, margins entire to minutely denticulate, terminal pinnae usually obdeltate; blade surfaces hirsute to tomentose; veins free; fertile fronds spreading to erect, stipes nearly lacking to 1⁄2 the frond length; fertile pinnae approximate to the sterile pinnae, far surpassing the sterile blades in height; spores striate, ridges smooth, broad, closely spaced; 2n⫽ca. 380 (Oax).

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15. ANEMIA PASTINACARIA Anemia pastinacaria Moritz ex Prantl, Unters. Morph. Gefa¨sskrypt. 2: 110. 1881. Lectotype (chosen by Lellinger, Proc. Biol. Soc. Wash. 98: 367. 1985). “New-Grenada [Colombia]. In convalli del Tigre.” Moritz 26 (B). Fig. 18D. Anemia pilosa M. Martens & Galeotti β longistipes Liebm., Mexic. Bregn. 301 (reprint 149). 1849. Anemia longistipes (Liebm.) C. Chr., Index Filic. 53. 1905. Type. Mexico. Oaxaca: Between Trapiche de la Concepcio´n and Tepitongo, Liebmann s.n. (C!). Anemia helveola Fe´e, Me´m. Foug. 9: 41. 1857. Type. Mexico. Oaxaca: Galeotti 6586bis (G?, not seen). The only specimen we have seen labelled as A. helveola is Galeotti 6541 (BR), which is A. pastinacaria. If indeed the type of A. helveola is that species, then A. helveola has priority. In that case, conservation of A. pastinacaria would be appropriate.

Rhizomes horizontal, compact, short-creeping, 4–8 mm diam.; rhizome hairs orange; sterile fronds erect, 8–20 cm long; stipes 1⁄4–1⁄3 the frond length, 0.4–1.9 mm diam., stramineous, hirsute; blades oblong to deltate, 1-pinnate, 2.5–9 cm wide, chartaceous to subcoriaceous; pinnae 5–9 pairs, opposite to subopposite, narrowly oblong, dimidiate, truncate at base, tapering toward apices, acute at tips, margins entire to minutely denticulate or erose, occasionally shallowly incised and forming 1–3 lobes on the acroscopic side of the lower pinnae; blade surfaces lightly hirsute to glabrous; veins free; fertile fronds to 25 cm tall, stipes 3⁄5–4⁄5 the frond length; fertile pinnae approximate to the sterile pinnae, far surpassing the sterile blades in height; spores striate, ridges spiny; 2n⫽228 (Jal, Nay, Oax, Trin).

Distribution. Frequent on open to partially shaded, disturbed, rocky slopes; 90–1900 m. Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Col, Ven, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol, Arg. Selected Specimens Examined. Chis (Breedlove 27100, NY; Pe´rez Farrera 700, MEXU). Gro (Lorea 2819, FCME, NY; Paxon et al. 17M817, MEXU). Jal (McVaugh 21068, MICH). Me´x (Hinton 2166, MEXU). Nay (Feddema 2638, MICH). Oax (Mickel 4203, NY). Sin (Ripley & Barneby 14031, NY). Ver (Te´llez 4731, MEXU).

There is variation in number of pinnae, stipe length, hairiness, texture, marginal teeth, and plant size. There are several species of this complex, however, in the Planalto of Brazil. The exact chromosome number has not been determined, but the number seems high (perhaps 2n⫽380 (decaploid)). A few plants from eastern Oaxaca and Chiapas (Mickel 1204, 1207, 5962, 5985, NY; Breedlove 36813, DS) have longer stipes, more rounded pinnae, more denticulate margins, and thinner texture, but do not appear distinct enough to be treated as a separate taxon.

Distribution. Open to lightly shaded rocky banks, grassy slopes, disturbed areas, in orange clay soils; 90–1150 m. Mexico; Bel, Guat, Hond, Salv, Nic, CR, Pan; Cuba; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol. Selected Specimens Examined. Chis (Breedlove 20544, NY, 22553, MEXU, NY). Gro (Hinton 14650, NY; Lorea 2820, NY). Jal (Diaz 21277, NY; Mickel 1342, NY). Nay (Feddema 906, MICH). Oax (Herna´ndez G. 448, NY; Solheim et al. 1165, NY). Ver (Bourgeau 3617, NY).

This species is distinct with its narrowly deltate fronds, pinnatifid apices, and oblong, acute pinnae. The spores are often a mixture of well formed (having narrow ridges and being short-

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to long-tuberculate) and malformed, suggesting this species is a triploid apomict, at least in Mexico.

16. ANEMIA PHYLLITIDIS Anemia phyllitidis (L.) Sw., Syn. Fil. 155. 1806. Figs. 20C, D. Osmunda phyllitidis L., Sp. Pl. 2: 1064. 1753. Anemidictyon phyllitidis (L.) J. Sm. in Hooker, Gen. Fil., t. 103. 1842. Type. Plumier, Traite´ Foug. Ame´r., pl. 156, 1705, based on a plant from Hispaniola.

Rhizomes ascending, 5–10 mm diam.; rhizome hairs orange; fronds erect, 20–70 cm long; stipes 1⁄2–2⁄3 the frond length, 0.7– 1.7 mm diam., stramineous, hirsute; blades deltate, 1-pinnate, (8– )12–26 cm wide, with conform terminal pinna; pinnae 2–5 pairs, opposite to subopposite, lanceolate, rounded at bases, shortpetiolate 1–4 mm, apices acuminate, margins finely serrulate; blade surfaces hirsute on costae but mostly glabrous; veins netted; fertile pinnae approximate to the sterile pinnae, slightly to far surpassing the sterile blades in height; spores striate, ridges with blunt spines; 2n⫽152 (Chis, Oax, and elsewhere), also probably 76 (Oax).

ploid (or triploid apomict), A. semihirsuta. Although the diploid form has not been found, cytological evidence suggests its presence now or in the past. See A. semihirsuta for more details (also Mickel, 1982). The presumed backcross of A. semihirsuta to A. phyllitidis (Anemia ⫻paraphyllitidis Mickel, Brittonia 34: 407. 1982. Type. Mexico. Oaxaca: Distrito Villa Alta, trail from Villa Alta to airstrip, Mickel 1118, NY!; Anemia collina sensu M. Martens & Galeotti, Me´m. Foug. Mexique 20. 1842, non Raddi, 1819; Figs. 20A, B) resembles A. phyllitidis, but has only casually anastomosing veins (mostly 25–32 anastomoses per pinna), malformed spores, and meiotic counts of ca. 38II ⫹ 76I and ca. 38II ⫹ 114I (Oax). The pinnae of the young fronds are oval but more elongate on mature fronds. It is most often confused with A. muenchii, which see for further discussion. Anemia ⫻paraphyllitidis is known from Mexico (Hgo, Oax, Chis), El Salvador, Costa Rica, Hispaniola, and Venezuela. Go´mez (Brenesia 18: 156. 1980) described Anemia ⫻didicusana as a hybrid between A. hirsuta and A. phyllitidis in Costa Rica, but the hybridization between these two species is very complex, involving fertile and/or sterile crosses with multiple ploidal levels of at least one of the parents. Examination of holotype, isotype, and paratype material of A. ⫻didicusana has shown it to be very close to A. ⫻paraphyllitidis. Plants of A. ⫻didicusana have 3–6 pairs of incised pinnae, whereas A. ⫻paraphyllitidis has entire pinnae or only a single incision in the basal pinnae-pair; there seems to be a difference in the number of anastomoses per pinna between the two entities. The complexity of the hybridization and the uniformity of other material of A. ⫻paraphyllitidis causes us to maintain A. ⫻didicusana as distinct until living material from Costa Rica can be studied.

17. ANEMIA RECONDITA

Distribution. Moist slopes of scrubby roadbanks or in woods along streams, widespread; (300–)800–1900 m. Mexico; Guat, Hond, Salv, Nic, CR, Pan; Gr Ant; Col, Ven, Trin, Guy, Sur, Ec, Peru, Braz, Bol, Parag, Arg, Uru. Selected Specimens Examined. Chis (Breedlove 21792, NY). Col (McVaugh 16085, NY). Gro (Hinton 14722, NY). Hgo (Koch 8773, NY). Jal (Gonza´lez T. 482, MICH). Me´x (Hinton 2185, NY). Mich (Hinton 12316, NY). Nay (McVaugh 18900, MICH). Oax (Hallberg 1485, NY). Qro (Ferna´ndez y Acosta 2107, NY). SLP (Pringle 3361, NY). Tam (Sharp et al. 50179, US). Ver (Purpus 1981, NY).

Anemia phyllitidis is widespread in tropical America, but uncommon in Mexico. It is distinct in its netted veins and pinna shape. It resembles A. mexicana superficially but differs in its netted veins, thin texture, and fertile pinnae approximate to the sterile blades. It is distinct from A. muenchii in the acuminate pinnae and more extensively netted veins. It apparently has crossed with A. hirsuta to form the hexa-

Anemia recondita Mickel, Brittonia 34: 409. 1982. Type. Mexico. Nayarit: SE of Tepic on Rte 15, Km 859–860, Mickel 1703 (NY!). Fig. 19D. Rhizomes horizontal, compact, short-creeping, 3–6 mm diam.; rhizome hairs orange; fronds erect, sterile fronds 12–19 cm tall, fertile fronds 15–42 cm tall; stipes 0.7–1 mm diam., stramineous, hirsute, ca. 1⁄3 the sterile frond height, ca. 2⁄3 the fertile frond height; blades narrowly oblong to narrowly deltate, 1pinnate, 2.5–5 cm wide, papyraceous, apices pinnatifid; pinnae 6–9 pairs, apices pinnatifid, pinnae opposite to subopposite, oblong, truncate to cuneate at bases, often excavate basiscopically, apices acute to obtuse, margins minutely denticulate and shallowly to deeply incised; veins free; blade surfaces hirsute; fertile pinnae approximate to the sterile pinnae, usually surpassing the sterile blades in height; spores generally malformed, rarely well formed, striate, the striate smooth, narrow; 2n⫽38II ⫹ 114I (Jal, Me´x, Nay), rarely n⫽190II (Nay), in meiosis.

ANEMIA SEMIHIRSUTA

59

cana ca. 71 (68–74) ␮m, whereas those of A. recondita average 100 (89–114) ␮m in diam. The sculpturing, however, is similar in all three–narrowly ridged with broad grooves between ridges.

18. ANEMIA SEMIHIRSUTA Anemia semihirsuta Mickel, Brittonia 34: 410. 1982. Type. Mexico. Oaxaca: Distrito Villa Alta, trail from Villa Alta to airstrip, Mickel 1120 (NY!). Figs. 20E, F, 21G.

Distribution. Found nearly everywhere A. hirsuta and A. jaliscana occur together, in disturbed areas, such as roadbanks and lightly shaded pastures; 900–2200 m. Mexico. Selected Specimens Examined. Gro (Toledo 98, XAL). Jal (Mickel 1685, NY; Pringle 3850, MEXU, NY). Me´x (Mickel 1336, NY). Mich (Hinton 12212, NY; del Llano 119, IEB). Mor (Lyonnet 510900001, MEXU). Nay (Mickel 1706a, 1708b (malformed spores), 1708c (well formed spores), NY).

Anemia recondita has apparently arisen through hybridization between A. hirsuta and A. jaliscana. Both putative parents are variable in size and dissection. Anemia hirsuta has subentire to highly lacerate, hirsute pinnae, usually pinnatifid blade apices, and spores with smooth ridges. Anemia jaliscana has nearly entire pinnae in the southern part of its range (Me´xico, and to a lesser extent, Jalisco), with distinctly subrhomboid or trapezoid rounded pinnae, but in northern Jalisco and Nayarit, pinnae can be denticulate to slightly lacerate, and A. jaliscana then resembles A. affinis. Anemia recondita also varies on the same plant, with the pinnae of sterile fronds generally more nearly entire and those of the fertile fronds more incised. Anemia recondita, consequently, may take various forms. Its degree of dissection depends largely on the dissection of A. hirsuta in that specific locality. Thus, hybrids may have barely incised pinnae (as little as some plants of A. jaliscana) to deeply incised pinnae, and although A. recondita never reaches the extremes of dissection found in A. hirsuta, at some localities with highly dissected A. hirsuta as a parent, it may be more dissected than plants of A. hirsuta at other localities. Identification can best be made when both parents from a locality are seen together with the hybrid, since A. recondita can then be spotted as intermediate. An individual plant taken out of context, however, might be taken for either A. hirsuta or A. jaliscana. Generally, any plant looking like either of these two species and having malformed spores (seen with 30⫻ magnification) can be identified as A. recondita. In one locality (Tepic-Compostela road, Nayarit), A. recondita grows with A. hirsuta, A. jaliscana, A. pastinacaria, and A. multiplex, and has a doubled chromosome number (decaploid, 2n⫽380) and well formed spores (Mickel 1708c). The spores are much larger than those of the parents. Spores of A. hirsuta at this locality average 72 (66–77) ␮m in diam., those of A. jalis-

Rhizomes horizontal, compact, short-creeping, 4–7 mm diam.; rhizome hairs orange; fronds erect, 10–30 cm tall; stipes 6– 18 cm long, ca. 1⁄2 the frond length, ca. 1 mm diam., stramineous, hirsute; blades narrowly oblong or narrowly deltate, 1-pinnate, the proximal 1–5 pairs of pinnae incised, chartaceous, hirsute; pinnae 5–10 pairs, opposite to subopposite, broadly oblong, cuneate at bases, often excavate basiscopically on lower margin, apices obtuse, margins minutely denticulate; veins mostly free, with 0–2(–6) anastomoses per pinna; fertile pinnae approximate to the sterile pinnae, usually somewhat surpassing the sterile blades in height; spores with rugulose, narrow, distant ridges; 2n⫽228 (Oax).

Distribution. Uncommon on disturbed, partially shaded banks in mesic forests; 900–1600 m. Mexico; Nic. Selected Specimens Examined. Chis (Breedlove 22268, 32536, DS; Matuda 4802, LL, MEXU, NY). Oax (Mickel 887, 1120, 1299, 4713, 4758, NY). Ver (Calzada et al. 10180, XAL; Calzada & Va´zquez B. 10236, XAL).

This species is apparently of hybrid origin between A. hirsuta and a diploid race of A. phyllitidis. See Mickel (1982) for details. It may well be a triploid apomict rather than a sexual hexaploid, but at this time we are uncertain. The backcross of A. semihirsuta to A hirsuta shows 2n⫽ca. 76II and 38I at meiosis (Oax). A collection with narrower pinnae, but with well formed spores, is probably of this species (Chis: Matuda 17901, DS, F, MEXU). A Chiapas collection (Matuda 1767, MICH, NY) that has malformed spores and casually anastomosing veins differs from typical A. semihirsuta in having larger pinnae and resembles A. ⫻didicusana (see discussion under A. phyllitidis).

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ANETIUM

19. ANEMIA SPECIOSA

to the sterile blades in height; spores striate, ridges smooth, close; 2n⫽152 (Jal), 2n⫽228 (Oax).

Anemia speciosa C. Presl, Suppl. Tent. Pterid. 89. 1845. Fig. 17F. Anemia mexicana Klotzsch var. paucifolia Hook., Sec. Cent. Ferns, t. 65. 1861. Ornithopteris speciosa (C. Presl) C. F. Reed, Bol. Soc. Brot., ser. 2, 21: 153. 1947. Type. Mexico. Ne´e s.n. (PR).

This species is similar to A. mexicana var. mexicana by its plant size, 1-pinnate blades, coriaceous texture, truncate pinna bases, and spore morphology (large, close, parallel ridges), but distinct by its 1–2(–4) pairs of sterile pinnae per blade, short fertile pinnae, and more southern and Antillean distribution.

Distribution. Infrequent on open to slightly shaded grassy slopes, often grazed, generally disturbed; 390–1850 m, down to 60 m in Isthmus of Tehuantepec. Mexico; Hisp; Col, Ven. Three other varieties are known from southern South America. Selected Specimens Examined. Chih (Palmer 91, NY). Dgo (Ortega 4294, US). Gto (Ventura V. & Lo´pez P. 9888, IEB). Jal (Pringle 1834, NY). Me´x (Kimnach & Moran 139, UC). Nay (Jones 23473, UC). Oax (Mickel 761, NY). Pue (Purpus 4045, UC). Sin (Breedlove 18050, NY). SLP (Schaffner 26, NY). Son (Gentry et al. 19290, NY). Tam (Bartlett 10866, MICH). Ver (Purpus 8460, NY). Zac (Anderson 12631, NY).

Distribution. Shaded limestone rocks in lowland forests; 200– 400 m. Mexico; Guat, Bel, Nic; Cuba. Selected Specimens Examined. Chis (Breedlove 21564, 32950, MEXU, NY, 48871, ENCB; Davidse et al. 29707, UC; Dressler 1623, GH, MEXU, NY). Oax (Calzada 10404, XAL, 10352, ENCB, UC, XAL; Sharp s.n., 29 Oct 1960, NY). Unverified, Doubtful, or Mistaken Reports. Ver (Nee 22363, F, XAL, cited by Palacios-Rios, 1992, but not verified).

20. ANEMIA TOMENTOSA VAR. MEXICANA Anemia tomentosa (Sav.) Sw. var. mexicana (C. Presl) Mickel, Iowa State Coll. J. Sci. 36: 427. 1962. Figs. 16C, D.

Unverified, Doubtful, or Mistaken Reports. Ags (Gonza´lezAdame 612, HUAA, cited by Siqueiros-Delgado & Gonza´lez-Adame, 2004, but not verified). BCN (reported by Knobloch & Correll, 1962, and Wiggins, 1980, as Anemia anthriscifolia).

The two cytotypes of A. tomentosa in Mexico apparently are allopatric and can be distinguished by spore size. The hexaploid, which occurs in Oaxaca, has spores 76–94 (avg. ca. 81) ␮m in diameter, whereas material from Jalisco (tetraploid) and Sonora has spores 69–79 (avg. ca. 74) ␮m in diameter. Anemia tomentosa is distinguished in Mexico by its thin texture and bipinnate-pinnatifid blades. In Mexico it most closely resembles A. karwinskyana, which is distinct in its catadromous architecture and has segments that are more obtuse and rounded rather than toothed.

Anemia fulva (Cav.) Sw. γ mexicana C. Presl, Suppl. Tent. Pterid. 84. 1845, based on A. flexuosa var.? [sic] anthriscifolia sensu Kunze, Linnaea 18: 308. 1844. Type. “In regione subtropica Mexico orientalis, Leibold pl. mex. exs. sine num.” (LZ–destroyed). Anemia distans Fe´e, Me´m. Foug. 9: 41. 1857. Type. Mexico. Oaxaca: Talea, Galeotti 6567bis (BR!).

Questionable Species Anemia guatemalensis Maxon, Chis (Pe´rez-Farrera 1452, UAMIZ, UNICACH, cited by Pe´rez-Farrera, Garcı´a, Riba & Lo´pez-Molina, Amer. Fern J. 93: 152, 2003, but not verified.).

Rhizomes horizontal, compact, short-creeping, ca. 7 mm diam.; rhizome hairs orange; fronds erect, 9–39 cm tall; stipes 1⁄2– 2 ⁄3 the frond length, 0.9–2.2 mm diam., stramineous to light brown, hirsute; blades deltate-ovate, bipinnate-pinnatifid, 4.6– 10.7 cm wide, chartaceous to subcoriaceous; pinnae 6–13 pairs, opposite to subopposite, pinnules narrowly adnate, ovate, segment lobes acute; blade surfaces hirsute; veins free; fertile pinnae approximate to or remote from the sterile pinnae, about equal

9. ANETIUM Anetium Splitg., Tijdschr. Natuurl. Gesch. Physiol. 7: 395. 1840. Pteridanetium Copel., Gen. Fil. 224. 1947, based on Anetium Splitg., not Anetia Endl. (1839); a superfluous new name according to Morton (Amer. Fern J. 43: 71. 1953). Type: Anetium citrifolium (L.) Splitg. [⬅ Acrostichum citrifolium L.].

ANOGRAMMA

61

Epiphytic; rhizomes slender, long-creeping, dorsiventral, dictyostelic, densely scaly; rhizome scales clathrate, iridescent; fronds remote; blades simple, entire (edges of larger fronds repand), elliptic, glabrous, each with a midrib; veins anastomosing, forming many series of elongate polygonal (often hexagonal or pentagonal) areoles on both sides of midribs, the long axis of the areoles oblique to the midrib, areoles lacking included veinlets, marginal veinlets short, free; sporangia borne mostly on the veins, but some also between veins in small clusters or larger, irregular patches, not organized into discrete sori, easily abraded or caducous when mature; indusia absent; paraphyses absent; spores tetrahedral-globose, surface of minute, dense, rod-like particles that form a partially reticulate layer, hyaline; gametophytes with paired gemmae; x ⫽ 60. Anetium is monotypic and widespread in the Neotropics. According to the phylogenetic reconstruction by Crane (1997), it is sister to Polytaenium, and differs from that by the sporangia being scattered on the veins and often between the veins on laminar tissue; also, the fronds are borne more distantly on the long-creeping rhizomes. See treatment of Radiovittaria for additional comments on relationships of the vittarioid ferns, of which Anetium is a member. Reference Crane, E. H. 1997 [1998]. A revised circumscription of the genera of the fern family Vittariaceae. Syst. Bot. 22: 509–517.

1. ANETIUM CITRIFOLIUM Anetium citrifolium (L.) Splitg., Tijdschr. Natuurl. Gesch. Physiol. 7: 395. 1840. Figs. 23A–D Acrostichum citrifolium L., Sp. Pl. 2: 1067. 1753. Hemionitis citrifolia (L.) Hook., Sp. Fil. 5: 193. 1864. Pteridanetium citrifolium (L.) Copel., Gen. Fil. 224. 1947. Lectotype (chosen by R. Tryon, 1964: 225). Plumier, Traite´ Foug. Ame´r., pl. 116, 1705, illustrating a collection from Martinique.

Rhizomes 1.5–4 mm diam.; rhizome scales brownish, lanceolate to ovate-attenuate, 5–10 ⫻ 1.5–2.5 mm; stipes absent or very short in large fronds; blades pendent and flaccid, herbaceous-fleshy when living, papyraceous when dried, elliptic-oblong or slightly oblanceolate, mostly 10–30(–100) ⫻ 2–8(–12) cm, decurrent at the base, apices obtuse to acute to abruptly acuminate; midribs extending 3⁄4 the length of the blades; 2n⫽120 (Trin).

Distribution. Epiphytic in lowland rain forests, on limestone ridges; 100–300 m. Mexico; Guat, Bel, Hond, Nic, CR, Pan; Jam, Hisp, PR; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol. Specimens Examined. Chis (Breedlove 33940, DS, NY; also cited by Rovirosa, 1909: 248, on the basis of Martı´nez 1093, but specimen not located). Ver (Wendt et al. 5802, CHAPA, MEXU, NY).

This rather rare species is easily distinguished from similarly shaped species of Polytaenium by the scattered, superficial sporangia. Sterile fronds can be discerned by the longer creeping rhizomes and large, papyraceous blades.

1 0 . A N O GR A M M A Anogramma Link, Fil. Spec. 137. 1841. Type: Anogramma leptophylla (L.) Link [⬅ Polypodium leptophyllum L.]. Terrestrial, or less commonly epipetric; rhizomes erect, sparsely scaly to hairy; fronds small, clumped; stipes slender, green to stramineous; blades 2–4-times pinnate, glabrous (except at bases); veins free; sporangia along the veins on ultimate and penultimate terminal segments; indusia and paraphyses absent; spores tetrahedral-globose, with equatorial ridges tan to dark brown; gametophytes said to be perennial, at least in A. leptophylla, with annual sporophytes; x ⫽ 29. There are perhaps seven species in the genus, but their taxonomy is still not entirely resolved. The type species, A. leptophylla, is subcosmopolitan, being widespread in Latin America, Africa, southern Europe, and southern Asia, but the others are rather restricted in range. Five are found in America and one is African; one on Ascension Island is extinct. The boundaries between A. leptophylla, A. guatemalensis, and A. chaerophylla are obscure. It is curious that the rare A. chaerophylla grows with A. leptophylla in Chiapas (Smith, 1981) and the only collection of A. chaerophylla in Oaxaca is mixed collection with A. leptophylla. Further study is needed on the intraspecific and intrapopulational variation. Anogramma is closely related to Pityrogramma and more distantly to Eriosorus (Tryon & Tryon, 1982), and differs from both relatives in size, texture, and lack of farina.

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ANOGRAMMA References

Unverified, Doubtful, or Mistaken Reports. Mor (Ordorica 161, ENCB, cited by Riba et al., 1986, but not verified).

Baroutsis, J. G. & G. J. Gastony. 1978. Chromosome numbers in the fern genus Anogramma. 2. Amer. Fern J. 68: 3–6; Gastony, G. J. & J. G. Baroutsis. 1975. Chromosome numbers in the genus Anogramma. Amer. Fern J. 65: 71–75; Nakazato, T. & G. J. Gastony. 2001. Molecular phylogenetics of Anogramma species and related genera (Pteridaceae: Taenitidoideae). Botany 2001, Albquerque, New Mexico, Abstract 344; Tryon, R. M. 1962. Taxonomic fern notes. II. Pityrogramma (including Trismeria) and Anogramma. Contr. Gray Herb. 189: 52–76.

The pale spores and the presence of rhizome scales distinguish this species from the more common A. leptophylla. Anogramma chaerophylla apparently is not contained within a monophyletic Anogramma, but is sister to Pityrogramma (Nakazato & Gastony, 2001).

Key to the Mexican Species of Anogramma

2. ANOGRAMMA LEPTOPHYLLA

1. Spores dark brown; rhizomes and stipe bases with hairs 1 mm long; blade segments rounded at apices. ......... 2. A. leptophylla. 1. Spores whitish to tan; rhizomes scaly or glabrous; blade segments obtuse to acute at apices. 2. Rhizomes with scales 1–1.5 ⫻ 0.1 mm. ............... 1. A. chaerophylla. 2. Rhizome scales absent. ........................................ 3. A. novogaliciana.

1. ANOGRAMMA CHAEROPHYLLA Anogramma chaerophylla (Desv.) Link, Fil. Spec. 138. 1841. Figs. 24F–J Gymnogramma chaerophylla Desv., Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck. Gesammten Naturk. 5: 305. 1811. Type. “Habitat in America Australis (Paraguay),” Commerson s.n. (isotype P-JU, photo MO).

Rhizomes short-creeping, ca. 1 mm diam.; rhizome scales few, whitish to tan, 1–1.5 ⫻ 0.1 mm; fronds 4–25 cm long, clumped; stipes 1⁄2–2⁄3 frond length, stramineous to reddish brown, especially at bases, glabrous, grooved; blades deltate, 3–4-pinnatepinnatifid, 2–10 cm wide; pinnae alternate, 4–6 pairs; segments linear-oblong, obtuse to acute at apices, thin, coalescing to ca. 2 mm wide at points, glabrous; spores whitish to tan; 2n⫽58 (Jam), 116 (Braz).

Anogramma leptophylla (L.) Link, Fil. Spec. 137. 1841. Figs. 24A–C. Polypodium leptophyllum L., Sp. Pl. 2: 1092. 1753. Gymnogramma leptophylla (L.) Desv., Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck. Gesammten Naturk. 5: 305. 1811. Lectotype (following Morton, Amer. Fern J. 60: 101–103. 1970). LINN 1251.56, photo A. Gymnogramme leptophylla (L.) Desv. var. mexicana Kunze, Linnaea 18: 310. 1844. Type. Mexico. Leibold 25 (B!). Pityrogramma guatemalensis Domin, Spisy Prˇir. Fak. Karlovy Univ. 88: 9. 1928. Anogramma guatemalensis (Domin) C. Chr., Index Filic., Suppl. 3: 26. 1934. Type. Guatemala. “Fuego,” Salvin & Godman 172 (K, photo UC!, frag. NY!).

Rhizomes short-creeping, 1 mm diam., with hairs 1 mm long; fronds 2–25 cm long, clumped; stipes 1⁄2–2⁄3 the frond length, stramineous to reddish brown at bases, glabrous except for scattered hairs 1 mm long at bases; blades lanceolate, 2–3-pinnate; pinnae 4–6 pairs; segments obovate, cuneate at bases, apices obtuse, glabrous; spores dark brown; 2n⫽54 (Nay), 58 (Oax, NZ), ca. 116 (India).

Distribution. Terrestrial on mossy banks in oak and other deciduous forests; 800–2800 m. Mexico; Guat, Salv, Nic, CR, Pan; Ven, Ec, Peru, Braz; Eurasia, Africa, NZ.

Distribution. Terrestrial in wet oak woods; 2000–2150 m. Mexico; Salv; Gr Ant; Galapagos, Braz, Parag, Arg, Uru. Selected Specimens Examined. Chis (Breedlove 22842, 40251, DS, 58297, 66115, CAS). Me´x (Hinton 7335, NY). Mich (Dı´az B. 6476, IEB). Oax (Mickel 4710b, 9164, NY). Ver (Castillo C. 3106, XAL; Purpus 2937, NY).

Selected Specimens Examined. Chih (Pringle 1446, NY). Chis (Breedlove 22937, NY). Col (McVaugh 16110, NY). Gro (Reveal et al. 4263, NY). Gto (Dı´az B. & Pe´rez C. 7393, IEB). Jal (McVaugh 22051, NY). Me´x (Hinton 5350, NY). Mich (McVaugh 22803, MICH). Mor (Lyonnet 777, MEXU, US). Nay (McVaugh 18977, MICH, NY). Oax (Mickel 5032, NY). Sin (Correll & Correll 28868, US). Son (Fishbein 1038, ARIZ). Ver (Mu¨ller 3050, NY). Unverified, Doubtful, or Mistaken Reports. Ags (Gonza´lezAdame 618, HUAA, cited by Siqueiros-Delgado & Gonza´lez-Adame, 2004, but not verified).

ARACHNIODES

Anogramma leptophylla is a variable species as to size and nature of ultimate segments. Frond segments vary from flabellate to long and finely dissected (variant known as A. guatemalensis; see Smith, 1981; R. Tryon, 1962; Tryon & Tryon, 1982). Nakazato and Gastony (2001) have shown A. guatemalensis to be nested within A. leptophylla.

3.

ANOGRAMMA NOVOGALICIANA

Anogramma novogaliciana Mickel in McVaugh, Fl. Nov.-Gal. 17: 169. 1992. Type. Mexico. Jalisco: San Sebastia´n, E of Arroyo del Cura, Mexia 1398 (NY!). Figs. 24D, E. Rhizomes short-creeping, ca. 1 mm in diameter, naked; fronds 3–8 cm long, clumped; stipes ca. 1⁄2 the length of the frond, stramineous to reddish brown especially at bases, glabrous; blades deltate, 2–3-pinnate-pinnatifid, 1.5–3.5 cm wide; pinnae alternate, 2–4 pairs; segments linear-oblong, obtuse to acute at apices, thin, glabrous; spores whitish to tan.

63

(ours), scaly; fronds monomorphic, medium-sized, erect, not articulate; stipes with 3 or more vascular bundles in cross section, scaly throughout or only at bases; blades subcoriaceous, 2–4(– 6)-pinnate, deltate to pentagonal, less often ovate, anadromous, proximal pinnae enlarged basiscopically, basiscopic edges of costular channels continuous with costal channels, with basiscopic edges continuous with rachis channels; rachises grooved adaxially, grooves narrow, continuous with those of costae, glabrous (ours) to hairy; segments often with acute or cuspidate teeth; veins free; indument absent or of minute hairs, glands and scales lacking; sori round; indusia round-reniform, each attached at a sinus, often caducous; paraphyses absent; spores bilateral with a winged or folded perispore; x⫽41. According to Tindale (1961), Arachniodes is a genus of about 50 species, mostly of Southeast Asia, a few in Africa, and only four in the New World. Our species is the only widespread one in the Neotropics. The relationships of Arachniodes seem to be generally with Dryopteris and Polystichum, but perhaps more closely with Polystichopsis (primarily Antilles) or Cyclodium (primarily northern South America and Amazonia). Morton (1960) allied it with Ctenitis, but that differs in lacking pronounced rachis grooves and having minute septate hairs along the midribs adaxially. Tryon and Tryon (1982) included Arachniodes (and also Polystichopsis) in a broadly circumscribed Dryopteris. It is uncertain whether the neotropical species of Arachniodes are really congeneric with Old World species. Preliminary molecular data support inclusion of Arachniodes in the dryopteroid clade, but distinct from Dryopteris; however, sampling is insufficent to be more specific (Cranfill, unpubl. data). References

Distribution. On rocks in humid pine-oak or pine-fir forests or tropical deciduous forests; 1300–2000 m. Mexico. Specimens Examined. Jal (Cervera 5460, UC; Gonza´lez T. 78, MICH; Gonza´lez & Palafox 45, ENCB, NY; Mexia 1690a, MICH; Pringle 3123, MEXU, UC; Velarde 17720, GUADA). Mich (Dı´az Luna 9304, GUADA).

Anogramma novogaliciana resembles A. chaerophylla in the pale spores and acute blade segments. It differs from A. chaerophylla in its lack of rhizome scales and the occurrence of sori on the penultimate as well as ultimate segments.

11. ARACHNIODES Arachniodes Blume, Enum. Pl. Javae 241. 1828. Type: Arachniodes aspidioides Blume. Byrsopteris C. V. Morton, Amer. Fern J. 50: 149. 1960. Type: Byrsopteris aristata (G. Forst.) C. V. Morton [⬅ Polypodium aristatum G. Forst] ⬅ Arachniodes aristata (G. Forst.) Tindale.

Commonly terrestrial (ours), less often epipetric or hemiepiphytic; rhizomes long- or short-creeping, rarely suberect

Christensen, C. 1920. A monograph of the genus Dryopteris. Part. II. The tropical American bipinnate-decompound species. Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 8, 6: 1–132; Morton, C. V. 1960. Observations on cultivated ferns. VI. The ferns currently known as Rumohra. Amer. Fern J. 50: 145–155; Tindale, M. D. 1961. Pteridophyta of southeastern Australia. Aspidiaceae. Contr. New South Wales Natl. Herb., Fl. Ser. 211: 47–78.

1. ARACHNIODES DENTICULATA Arachniodes denticulata (Sw.) Ching, Acta Bot. Sin. 10: 260. 1962. Figs. 25A, B. Polypodium denticulatum Sw., Prodr. 134. 1788. Aspidium denticulatum (Sw.) Sw., J. Bot. (Schrader) 1800(2): 40. 1801. Polystichum denticulatum (Sw.) J. Sm., J. Bot. 4: 195. 1841. Dryopteris denticulata (Sw.) Kuntze, Revis. Gen. Pl. 2: 812. 1891. Rumohra denticulata (Sw.) Copel., Gen. Fil. 114. 1947. Byrsopteris denticulata (Sw.) C. V. Morton, Amer. Fern J. 50: 152. 1960. Type. Jamaica. Swartz s.n. (S; isotype C). Aspidium jucundum Fe´e, Me´m. Foug. 10: 41. 1865. Dryopteris denticulata f. jucunda (Fe´e) C. Chr., Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 8, 6: 115. 1920. Lectotype (chosen by Christensen, 1920: 115). Mexico. [Oaxaca:] Galeotti 6563 (P; isotypes BR–photos UC!, US!, RB).

Rhizomes ascending to suberect, short, compact; rhizome scales brown, lustrous, 10–20 ⫻ 1–2 mm, linear-lanceolate, entire; fronds clustered, mostly (30–)50–90 cm long; stipes casta-

64

ARGYROCHOSMA

neous at base, often stramineous distally, 12–32 cm ⫻ (1–)2–4 mm, 1⁄3–1⁄2(–2⁄3) the frond length, glabrous except for linear brown scales at bases; blades dark green adaxially, lighter green abaxially, chartaceous to subcoriaceous, 12–30 ⫻ 8–26 cm, deltate to pentagonal, 4-pinnate at bases, the proximal pair of pinnae strongly inequilateral, greatly exaggerated basiscopically, anadromous; rachises glabrous; pinnae alternate; segments oblanceolate to obovate, 5–8 ⫻ 3–4 mm, bases cuneate, margins strongly dentate with spinulose tips; indument of costae and costules of sparse, hair-like, fibrillose scales; indusia tan to red-brown, glabrous, persistent, 1–2(–3) mm diam.; 2n⫽82 (Oax, Chis), 164 (Jam).

Distribution. Terrestrial in wet montane forests; 1000–2900 m. Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Cuba, Jam; Col, Ven, Guy, Ec, Peru, Braz, Bol. Selected Specimens Examined. Chis (Breedlove 53600, CAS; Calzada et al. 8728, UC, XAL); Gro (Lorea 3290, FCME). Hgo (Gimate L. 797, NY, XAL). Oax (Croat & Hannon 65634, UC; Wendt 5058, NY). Pue (Ventura A. 366, ENCB, 385, ARIZ, ENCB, LL, NY). Ver (Va´zquez B. 489, UC, XAL; Ventura A. 12741, MEXU).

Christensen (1920) recognized eight named forms and varieties and several unnamed ones of this polymorphic species. These variants differ chiefly in size and dissection of the blades, blade texture, and other rather nebulous characteristics. Chromosome counts indicate that polyploidy is a part of this varia-

bility. Christensen’s forms seem to be connected by numerous intermediates, so we choose not to apply infraspecific names to Mexican material.

1 2 . A R GY R O C H O SM A Argyrochosma (J. Sm.) Windham, Amer. Fern J. 77: 38. 1987. Notholaena R. Br. sect. Argyrochosma J. Sm., J. Bot. (Hooker) 4: 50. 1841. Lectotype (chosen by Christensen, 1906, Index Filic. XL): Notholaena nivea (Poir.) Desv. [⬅ Pteris nivea Poir.] ⬅ Argyrochosma nivea (Poir.) Windham. Usually epipetric; rhizomes compact, horizontal to ascending, usually unbranched; rhizome scales tan to brown, concolorous, linear to lanceolate, margins entire; fronds monomorphic, clumped, 3–30 cm long; stipes castaneous to atropurpureous or black, terete or somewhat flattened adaxially, glabrous except for a few scales near bases; blades lanceolate, ovate, or deltate, 2–6pinnate, coriaceous to chartaceous; abaxial surfaces usually covered by whitish farina, less commonly glabrous; adaxial surfaces glabrous or sparsely glandular, dull; rachises straight or flexuous; ultimate blade segments stalked or subsessile, usually ecostate, elliptic to ovate or deltate, usually less than 4 mm wide; bases subcordate, stalks lustrous and dark-colored; segments often articulate, deciduous; segment margins plane or recurved; veins of ultimate segments free, usually obscure; false indusia absent, though the unmodified or poorly differentiated margins somewhat recurved; sori borne along veins abaxially, often submarginal; sporangia containing 64 or 32 spores, usually intermixed with farina; spores tetrahedral-globose, brown, with cristate or rugose surfaces; x⫽27. Argyrochosma is a genus of about 20 species, all New World. Twelve species occur in Mexico. Molecular data (Gastony & Rollo, 1995) show that Argyrochosma is monophyletic and sister to Pellaea and Astrolepis. References Tryon, R. M. 1956. A revision of the American species of Notholaena. Contr. Gray Herb. 179: 1–106; Windham, M. D. 1987. Argyrochosma, a new genus of cheilanthoid ferns. Amer. Fern J. 77: 37–41.

Key to the Mexican Species of Argyrochosma 1. Blade indument absent. 2. Ultimate segments not articulate, the dark color of the stalks passing into at least the basal part of the segments; segments plane to slightly recurved. 3. Stipes and rachises castaneous to dark brown; blades 2–3-pinnate. .................................................................................................. 5. A. jonesii. 3. Stipes and rachises blackish; blades 2-pinnate. ............................................................................................................................. 7. A. lumholtzii. 2. Ultimate segments articulate, the dark color of the stalks stopping abruptly in a transverse line at their apices; segments often folded lengthwise. 4. Segments 1–3 mm wide; rachises adaxially shallowly grooved or flat, castaneous, flexuous; Son, Coah, Chih, NL, Zac. ...................................................................................................................................................................................................... 8. A. microphylla. 4. Segments 3–4 mm wide; rachises terete, atropurpureous to black, straight; bases of ultimate segments mostly inequilaterally cordate; widespread. ............................................................................................................................................................................... 3. A. formosa. 1. Blades farinose, with or without hairs or scales. 5. Stipes 1⁄10–1⁄3 of the frond length; blades narrowly oblong, 0.8–1.5(–2) cm wide. 6. Stipes and rachises castaneous; rachises with a few small, linear to linear-lanceolate, castaneous scales; adaxial segment surfaces sparsely farinose. ....................................................................................................................................................................... 9. A. pallens.

ARGYROCHOSMA FENDLERI

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6. Stipes and rachises blackish; rachises without scales; adaxial segment surfaces glabrous or sparsely farinose near margins. ................................................................................................................................................................................................. 10. A. palmeri. 5. Stipes 1⁄3–1⁄2 of the frond length; blades deltate, lanceolate, or ovate, rarely oblong, (2.5–)3.5–18 cm wide. 7. Blade indument of farina plus hairs or scales. 8. Blades with sparse hairs on rachises and adaxial segment surfaces; Mor. ............................................................................... 12. A. pilifera. 8. Blades with linear-lanceolate scales on rachises, hairs absent; BCS. ................................................................................. 11. A. peninsularis. 7. Blade indument of farina only, hairs and scales absent. 9. Rachises strongly flexuous. .............................................................................................................................................................. 2. A. fendleri. 9. Rachises straight or only weakly flexuous. 10. Color of segment stalks clearly passing into the segment bases. ................................................................................... 6. A. limitanea. 10. Color of segment stalks stopping abruptly at segment bases. 11. Stipes and rachises blackish, rarely castaneous or atropurpureous; segments (2.5–)3–5 mm wide, usually glabrous adaxially, occasionally farinose; abaxial farina white; widespread. ............................................................... 4. A. incana. 11. Stipes and rachises castaneous to atropurpureous; segments 1–2(–3) mm wide, sparsely farinose adaxially; abaxial farina pale yellow; Chih, Coah, NL. .............................................................................................................. 1. A. delicatula.

1. ARGYROCHOSMA DELICATULA Argyrochosma delicatula (Maxon & Weath.) Windham, Amer. Fern J. 77: 40. 1987. Figs. 27D–G. Notholaena delicatula Maxon & Weath., Contr. Gray Herb. 127: 7. 1939. Type. Mexico. Coahuila: Lerios, 15 leagues east of Saltillo, Palmer 1387 (US!; isotypes GH!, K!, P!, US!, YU!).

Rhizomes compact, horizontal or ascending; rhizome scales 4– 6 ⫻ 0.2–0.3 mm, linear, concolorous, orange-brown to dark brown, entire; fronds (6–)10–25(–35) cm long, clumped; stipes nearly half the frond length, castaneous to atropurpureous, lustrous, glabrous, terete; blades mostly deltate, tripinnate to subquadripinnate, herbaceous; rachises terete; pinnae 4–8 pairs; segments rhombic, oblong, or orbicular, 1–2(–3) mm across, segments articulate where the dark color of the stalk abruptly terminates at the base of the segments; abaxial surfaces with pale yellow farina; adaxial surfaces sparsely farinose; sori borne on the distal 1⁄2 of the veins, margins unmodified; sporangia 64-spored; 2n⫽54 (NL).

CAS, GH, K, LL, MEXU, MICH, MO, MSC, NY, P, UC, US; Windham 482, NY).

This species is not entirely distinct from Argyrochosma incana, although from the few collections available it seems sufficiently so. Possibly it is a geographic variant of that species. R. Tryon (1956) distinguished A. delicatula from A. incana by the former having scattered adaxial farina, whereas A. incana was thought to be glabrous adaxially. Specimens of A. incana with adaxial farina occur sporadically throughout the range of the species. Argyrochomsa delicatula is best distinguished on the basis of its very small segments, thinner texture, and pale yellow farina. This species was not reported by Windham in FNA Ed. Comm., 1993, but a specimen at NY (Lemmon s.n., 23 Apr 1880, from southeastern Arizona), formerly determined as A. incana, has the small, thin segments with pale yellow farina characteristic of A. delicatula. Other records of A. incana from Arizona should be critically examined.

2. ARGYROCHOSMA FENDLERI Argyrochosma fendleri (Kunze) Windham, Amer. Fern J. 77: 40. 1987. Figs. 27K, L. Notholaena fendleri Kunze, Farrnkra¨uter 2: 87, pl. 136. 1851. Pellaea fendleri (Kunze) Prantl, Bot. Jahrb. Syst. 3: 417. 1882. Cheilanthes cancellata Mickel, Phytologia 41: 433. 1979. Type. U.S.A. New Mexico: Trapp formations of the Rio del Norte near Santa Fe, Fendler 1017a (LZ, destroyed; isotypes B, BM, GH!, K, MO).

Distribution. Exposed or shaded rocky places, limestone ledges, shady slopes, 900–2230 m. USA (se Ariz); Mexico. Selected Specimens Examined. Chih (Jones s.n., 12 Sep 1903, UC). Coah (Johnston et al. 11506, LL). NL (Hinton et al. 21025, 21597, 23260, TEX; Palmer 1386, GH, P, US; Pray 3029, LAM; Pringle 2581, B, BM,

Rhizomes compact, horizontal to ascending, 2–3 mm diam.; rhizome scales concolorous orange-brown, linear-lanceolate, 7– 10 mm; fronds 5–25 cm long, clumped; stipes 1⁄2 the frond length, castaneous, 0.75–1.5 mm diam., terete, glabrous; blades deltate, 4–6-pinnate proximally; rachises terete, glabrous, strongly flexuous; pinnae 4–5 pairs, costae of pinnae and pinnules strongly flexuous; ultimate segments not articulate, dark color of stalks continuing into segment bases abaxially, coriaceous, covered by whitish farina, adaxially glabrous or with scattered farina; segment margins plane to recurved, often partially concealing sporangia; veins usually obscure adaxially; sori submarginal, borne on distal 1 ⁄4 of veins; sporangia 64-spored; 2n⫽54 (USA).

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Distribution. Rocky slopes and cliffs, usually on granitic or volcanic substrates; elevation unknown in Mexico, 1700–3000 m in North America. USA (Colo, N Mex, Wyo); Mexico. Specimen Examined. Son (Mason 3202, ARIZ).

Argyrochosma fendleri is distinct by its strongly flexuous rachises and rachillae. Argyrochosma limitanea has slightly flexuous rachises and costae, but the frond axes are not so strongly spreading, and this species is more widespread in Mexico. All specimens of A. limitanea with slightly flexuous rachises and costae have 32 spores per sporangium, whereas specimens of A. fendleri consistently have 64.

Distribution. Dry, rocky, wooded slopes, limestone slopes, steep rocky ravines, thorn scrub, in mortar at Monte Alban; 1250–2700 m. Mexico; Guat. Selected Specimens Examined. Ags (Rzedowski 25048, NY). Chis (Ghiesbreght 227, GH, NY, US). Gro (Anderson 12874, NY). Gto (Ventura & Lo´pez 8900, CAS). Hgo (Frye & Frye 2558, GH, MO, NY). Me´x (Bravo 78, MEXU). NL (Pennell 17177, US). Oax (Mickel 4092, NY). Pue (Purpus 1151, NY). Qro (Rzedowski 47536, IEB). SLP (Pringle 3401, GH, LL, MO, NY). Tam (Stanford et al. 882, DS, NY). Ver (Bourgeau 2894, GH, NY, US).

Argyrochosma formosa resembles A. incana in its black frond axes, stalked segments and being widespread, but lacks farina. The other non-farinose species of Argyrochosma in Mexico, A. microphylla, is distinct by its smaller segments (1–3 mm vs. 3–5 mm) and castaneous (vs. black) frond axes.

3. ARGYROCHOSMA FORMOSA Argyrochosma formosa (Liebm.) Windham, Amer. Fern J. 77: 40. 1987. Figs. 26A–C. Allosorus pulchellus M. Martens & Galeotti, Me´m. Foug. Mexique 47, pl. 10, f. 1. 1842, hom. illeg., non C. Presl, 1836. Allosorus formosus Liebm., Mexic. Bregn. 220 (reprint 68). 1849. Pellaea pulchella (M. Martens & Galeotti) Fe´e, Me´m. Foug. 5: 129. 1852. Pellaea formosa (Liebm.) Maxon, Contr. U.S. Natl. Herb. 24: 61. 1922. Notholaena formosa (Liebm.) R. M. Tryon, Contr. Gray Herb. 179: 99. 1956. Cheilanthes formosa (Liebm.) Mickel & Beitel, Mem. New York Bot. Gard. 46: 113. 1988. Type. Mexico. Oaxaca: S of Sola, Galeotti 6352 (BR!; isotype K!). For additional synonymy, see R. Tryon (1956).

Rhizomes compact, horizontal; rhizome scales 6–9 mm ⫻ 0.3 mm, tan to dark orange, linear; fronds to ca. 40 cm tall, clumped; stipes 1⁄4–1⁄2 the frond length, atropurpureous to black, glabrous, terete; blades deltate to ovate-lanceolate, 3–4-pinnate; rachises terete; pinnae 6–9 pairs, alternate to subopposite, glabrous above, gray-green, abaxial surfaces glabrous to glaucous; segments orbicular to inequilaterally cordate, appearing sagittate when folded, articulate, 3–4 mm long and wide, dark axis color stopping abruptly at bases of segments; sori elongate along vein ends, forming a marginal band 0.5–1 mm wide, laminar margins curved, unmodified; fertile segments often folded in half lengthwise; n⫽2n⫽81, apogamous (Oax).

4. ARGYROCHOSMA INCANA Argyrochosma incana (C. Presl) Windham, Amer. Fern J. 77: 40. 1987. Figs. 27A–C, H, J, 28E–H. Notholaena incana C. Presl, Reliq. Haenk. 1(1): 19, t. 1, f. 2. 1825. Cheilanthes incana (C. Presl) Mickel & Beitel, Mem. New York Bot. Gard. 46: 115. 1988. Lectotype (chosen by Maxon & Weatherby, Contr. Gray Herb. 127: 5. 1939). Mexico. Haenke s.n. (PR!, photo GH!). Gymnogramma candida Mett., Abh. Senckenberg. Naturf. Ges. 3: 50. 1859. Pellaea candida (Mett.) Prantl, Bot. Jahrb. Syst. 3: 417. 1882. Lectotype (chosen by R. Tryon, 1956: 92). Mexico. Valle de Me´xico, Schmitz 231 (B, photos BM, GH!, US!).

Rhizomes short, stout, compact, horizontal or ascending; rhizome scales 5–6 ⫻ 0.8 mm, linear-lanceolate, concolorous, dark lustrous brown, entire; fronds to 30 cm long, clumped; stipes ca. 1 ⁄2 the frond length, atropurpureous to black, rarely castaneous, lustrous, glabrous, terete; blades deltate, (2.5–)3.5–8 cm wide at bases, 3–4-pinnate at bases, pinnae 6–9 pairs, opposite; segments (2.5–)3–5 mm long, mostly elliptic, gray-green adaxially with sparse white farina, dark color of pinna rachis ending abruptly at base of segments; abaxial surfaces densely white-farinose; segment margins not curved nor modified into false indusia; sori elongate along the veins, forming a band 0.5–1.5 mm wide; 2n⫽54 (USA).

ARGYROCHOSMA LIMITANEA

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Notholaena jonesii Maxon, Amer. Fern J. 7: 108. 1917. Pellaea jonesii (Maxon) C. V. Morton, Amer. Fern J. 40: 251. 1950. Cheilanthes jonesii (Maxon) Munz, Aliso 4: 87. 1958. Type. U.S.A. California: Inyo Co., Panamint Canyon, 4 May 1897, Jones s.n. (US!; photo GH).

Rhizomes short, compact, horizontal; rhizome scales 5–7 ⫻ 0.3 mm, dark brown, linear; fronds 4–15 cm long, clumped; stipes 1⁄2 to 2⁄3 the frond length, castaneous to dark brown, terete; blades ovate-lanceolate, 2–3-pinnate; abaxial and adaxial surfaces glabrous; rachises straight, terete or adaxially flattened to shallowly grooved; segments orbicular to ovate, not articulate, the dark color of stalks continuing into segment bases abaxially; segment margins plane to slightly recurved, not concealing sporangia; sori submarginal, on distal 1⁄2 of secondary veins; sporangia 64spored; 2n⫽54, 108 (USA). Distribution. Wooded, rocky slopes, steep banks and ledges, shaded rocks, pine-oak forests; 800–2950 m. USA (Ariz); Mexico; Guat; Hisp. Selected Specimens Examined. Ags (Rzedowski 25048a, NY). Chih (Palmer 215, BM, K, NY, US). Chis (Ghiesbreght 226, G, GH, K, YU). Coah (Wendt & Riskind 1701, NY). Dgo (Miller et al. 346, UC). DF (Langman 2616, US). Gro (Mexia 9014, US). Gto (Kenoyer 1969, US). Hgo (Novoa 28, MEXU). Jal (McVaugh 13093, US). Me´x (Pringle 11267, NY, US). Mich (Arse`ne 5756, US). Mor (Pringle 15698, US). Nay (Te´llez 10546, MEXU). NL (Aguirre C. 1093, ENCB). Oax (Mickel 6478, NY). Pue (Clausen 7492, NY). Qro (Dı´az-Barriga & Carranza 7359, IEB). SLP (Schaffner 961e, G, GH, K). Sin (Breedlove 18479, NY). Son (Drouet et al. 3667, US). Tam (Stanford 2569, NY). Tlax (Arse`ne 1739, MEXU). Zac (Breedlove 63973, CAS). Unverified, Doubtful, or Mistaken Reports. Ver (Ortega & Pattison 2183, XAL, cited by Palacios-Rios, 1992, but not verified).

Argyrochosma incana is the most widespread and variable species of Argyrochosma in Mexico. It is distinct by its deltate, 3–4pinnate blades with white farina usually limited to the abaxial surfaces. The species is highly variable in occasionally having (1) smaller segments as little 2 mm wide; (2) farina abaxially; (3) axes somewhat flexuous; (4) axes atropurpureous rather than blackish; and (5) segments more strongly articulate. There is a particularly distinct collection from Coahuila (Clark 6746, NY) that has strongly ascending pinnae and small articulate segments. A group of specimens from Chihuahua (Knobloch 5960, MSC; Pringle s.n., 21 Sep 1888, MEXU, UC), Coahuila (Dorr 2509, MEXU, TEX, 2519, TEX; Henrickson 11909, 11990a, LL), Guanajuato (Correll & Correll 28813, LL), Tamaulipas (Stanford 684, MEXU, 2569, NY), and Zacatecas (Henrickson 13332, 13475, LL) have especially small segments (though the Correll specimen has small and large segments on the same plant), somewhat flexuous rachises, and atropurpureous axes. This may be as distinct as A. delicatula, but more work is needed on this complex.

5. ARGYROCHOSMA JONESII Argyrochosma jonesii (Maxon) Windham, Amer. Fern J. 77: 40. 1987. Figs. 26D, E.

Distribution. Calcareous cliffs and ledges; 800 m, 600–1900 m in USA. USA (Ariz, Calif, Nev, Utah); Mexico. Specimens Examined. Son (Burgess 5759, ARIZ; Fischer & Yatskievych 82–12, 82–23, ARIZ). Unverified, Doubtful, or Mistaken Reports. BCN (reported by Wiggins, 1980, but not verified).

Windham (1993) reported two sexually reproducing cytotypes in the southwestern United States. The diploid is known from a few localities in the Sonoran and Mojave deserts; the tetraploid is found throughout the Mojave Desert and cismontane southern California. Argyrochosma jonesii most closely resembles A. lumholtzii, but the former can be readily distinguished by the castaneous to brownish stipes and rachises.

6. ARGYROCHOSMA LIMITANEA Argyrochosma limitanea (Maxon) Windham, Amer. Fern J. 77: 40. 1987. Notholaena limitanea Maxon, Amer. Fern J. 9: 70. 1919. Pellaea limitanea (Maxon) C. V. Morton, Amer. Fern J. 40: 251. 1950. Cheilanthes limitanea (Maxon) Mickel, Phytologia 41: 434. 1979. Type. U.S.A. New Mexico: Dona Ana Co., Tortugas Mt, SE of Las Cruces, 14 Sep 1902, Wooton s.n. (US!, photo GH).

Rhizomes short, compact, horizontal; rhizome scales brown; fronds 5–30 cm, linear; stipes 1⁄3–1⁄2 the frond length, reddish

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brown to black, terete, glabrous; blades narrowly to broadly deltate, 3–5-pinnate proximally, coriaceous, abaxially covered by dense white farina, adaxially glabrous or sparsely glandular; rachises terete, glabrous adaxially, straight to somewhat flexuous; costae straight to slightly flexuous, branches not arising from prominent angles; ultimate segments not articulate to stalks, dark color of stalks passing into segment bases abaxially; segment margins recurved, often concealing sporangia; sporangia submarginal, borne on distal 1⁄2 of secondary veins, 32-spored.

Blades lanceolate to oblong, 3–4-pinnate, proximal pinnae ⁄ – ⁄ the blade length; abaxially covered by white (rarely pale yellow) farina; n⫽2n⫽81, apogamous (USA). 14 13

Distribution. Calcareous or non-calcareous cliffs and ledges; 800–2800 m. Sw USA; Mexico. This species is distinct from A. incana and A. delicatula by the axis color passing into the segment bases. For the most part, these taxa have different geographic ranges, and genetic interaction between them is precluded because both are asexual triploids. Key to the Mexican Subspecies of Argyrochosma limitanea 1. Blades broadly deltate, 4–5-pinnate, with basal pinnae at least 1⁄2 as long as blades; Chih, Son. ................... 6a. subsp. limitanea. 1. Blades lanceolate to oblong, 3–4-pinnate, with basal pinnae 1⁄4–1⁄3 as long as blades; widespread. ....... 6b. subsp. mexicana.

6a. Argyrochosma limitanea subsp. limitanea

Figs. 28A–C.

Blades broadly deltate, 4–5-pinnate, with proximal pinnae at least 1⁄2 as long as blades; n⫽2n⫽81, apogamous (Coah).

Distribution. Rocky cliffs and slopes, usually calcareous; 1500– 2800 m. USA (Ariz, N Mex, Tex); Mexico. Selected Specimens Examined. Chih (Correll 23290, LL, NY). Coah (Stanford et al. 107, NY, US). Dgo (Palmer 465, NY). Hgo (Clark 7022, NY). NL (Favela 143, ENCB). Qro (Zamudio R. 3396, IEB). Sin (Breedlove 18521, NY). SLP (Lundell 5793, ARIZ, MICH). Tam (Stanford et al. 684, NY, UC, US). Zac (Dorado 1596, MEXU).

There are some specimens in Chihuahua that are intermediate between the two subspecies.

7. ARGYROCHOSMA LUMHOLTZII Argyrochosma lumholtzii (Maxon & Weath.) Windham, Amer. Fern J. 77: 40. 1987. Figs. 26F, G. Notholaena lumholtzii Maxon & Weath., Contr. Gray Herb. 127: 16. 1939. Type. Mexico. Sonora: Huehuerachi, Hartman (Lumholtz Exped.) 298 (US!, frag. NY!; isotypes GH!, PH!).

Distribution. Calcareous or non-calcareous cliffs and ledges; 800–2300 m. USA (Ariz, Calif, N Mex, Utah); Mexico. Selected Specimens Examined. Chih (Bye & Weber 8321, MEXU, MICH; Correll 22634, US; Knobloch 1146, 8027, MSC; LeSueur 1127, GH, MICH, UC, US; Pringle 1583, MO). Son (Phillips 738, US; Shreve 6735, US; Wiggins 7127, DS; Wiggins & Rollins 443, DS).

6b. Argyrochosma limitanea subsp. mexicana (Maxon) Windham, Contr. Univ. Michigan Herb. 19: 32. 1993. Fig. 28D. Notholaena limitanea Maxon subsp. mexicana Maxon, Amer. Fern J. 9: 72. 1919. Notholaena nivea (Poir.) Desv. var. mexicana (Maxon) Farwell, Amer. Midl. Naturalist 12: 283. 1931. Cheilanthes limitanea (Maxon) Mickel var. mexicana (Maxon) Mickel, Phytologia 41: 435. 1979. Type. Mexico. Chihuahua: Santa Eulalia Mts., Pringle 451 (US!; isotypes GH!, MICH!, NY!, P!).

Rhizomes compact, erect; rhizome scales linear, 4–5 mm long, entire, somewhat contorted, orange-brown; fronds 7–15 cm long, clumped; stipes ca. 2⁄5 the frond length, terete, black, glabrous; blades deltate-lanceolate or deltate-ovate, bipinnate, herbaceous or subcoriaceous, glabrous, gray-green; rachises straight, terete, glabrous; pinnae 6–8 pairs; segments remote, orbicular or slightly oblong to subcordate, 3–4 mm diam., entire, the terminal deltate, non-articulate, the dark color of the stalks passing into the segment bases; segment margins unmodified, plane to slightly recurved; sporangia 64-spored, borne on the distal 1⁄2–1⁄3 of veins.

ARGYROCHOSMA PALLENS

Distribution. Habitat data absent; 1300 m. Mexico. Specimens Examined. Son (Fishbein 4458, MO; Lloyd (Lumholtz Exped.) 489, GH).

Argyrochosma lumholtzii differs from A. jonesii by the blackish axes and the bipinnate blades (vs. castaneous to brownish axes and the blades 2–3-pinnate).

8. ARGYROCHOSMA MICROPHYLLA Argyrochosma microphylla (Mett. ex Kuhn) Windham, Amer. Fern J. 77: 40. 1987. Figs. 26H, J, K. Pellaea microphylla Mett. ex Kuhn, Linnaea 36: 86. 1869. Notholaena parvifolia R. M. Tryon, Contr. Gray Herb. 179: 98. 1956. Cheilanthes parvifolia (R. M. Tryon) Mickel, Phytologia 41: 435. 1979. Type. U.S.A. New Mexico: Wright 825 (B, photo GH).

Rhizomes short, compact, horizontal; rhizome scales brown to dark orange, 4–7 mm long; fronds to 25 cm long, clumped; stipes 1⁄3–1⁄2 the frond length, castaneous, terete to somewhat flattened adaxially, glabrous; blades deltate to ovate, 3–4-pinnate; pinnae 5–8 pairs, alternate; rachises slightly to strongly flexuous, flattened or shallowly grooved adaxially; segments orbicular to cordate, 1–3 mm, articulate, fertile often folded, dark color of stalks stopping abruptly at segment bases; adaxial and abaxial surfaces glabrous, gray-green adaxially; sori submarginal, elongate along vein ends and forming marginal bands 1 mm wide, laminar margin curved, undifferentiated; fertile segments often folded in half; 2n⫽54 (Coah, USA).

69

Distribution. Rocky limestone hillsides and cliffs, open talus slopes; 1000–2300 m. USA (N Mex, Tex.); Mexico. Selected Specimens Examined. Chih (Harvey 1512, MICH; Pringle 440, MICH, NY, 458, NY; Worthington 12521, NY). Coah (Harvey 1196, MICH; Hinton 16510, NY; Muller 3079, MICH, NY). NL (Pennell 17188, MEXU, NY, US). Son (Fishbein 2156, ARIZ). Zac (Stanford et al. 47, NY).

Argyrochosma microphylla is distinct by the small segments and shallowly grooved rachises. Some plants have ascending pinnae like A. limitanea, some zig-zag rachises like A. fendleri. Dorr et al. 2095 (NY), has strongly flexuous rachises like A. fendleri, but the rachillae are less flexuous, and it lacks farina. This species may include diploid and tetraploid cytotypes (Knobloch et al., 1973).

9. ARGYROCHOSMA PALLENS Argyrochosma pallens (Weath. ex R. M. Tryon) Windham, Amer. Fern J. 77: 41. 1987. Figs. 29K–M. Notholaena pallens Weath. ex R. M. Tryon, Contr. Gray Herb. 179: 78, f. 45. 1956. Cheilanthes pallens (Weath. ex R. M. Tryon) Mickel & Beitel, Mem. New York Bot. Gard. 46: 123. 1988, non (Sw.) Desv., 1827. Type. Mexico. Durango: Sandia Station, Pringle 10149 (GH!; isotypes P!, US!).

Rhizomes erect or decumbent; rhizome scales linear-lanceolate and ending in capillary apices, 2–5 mm long, concolorous, reddish brown, entire; fronds to 20 cm long, clumped; stipes 1⁄5–1⁄3 the frond length, stipes and rachises castaneous, terete, farinose and with sparse, linear-lanceolate to linear, castaneous scales 1– 2 mm long; blades narrowly oblong, 1–2 cm wide, bipinnatepinnatifid to tripinnate; pinnae 9–12 pairs; segments oblong to broadly deltate-ovate, obtuse, not articulate, the dark color of costae passing into bases of segments, but obscurely so; adaxial surfaces sparsely farinose; abaxial surfaces densely white-farinose; sori elongate along veins, forming a band 0.5–1 mm wide, segment margins plane to slightly curved, not modified; 2n⫽54 (Oax).

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ARGYROCHOSMA

Distribution. Cliffs and ledges, usually on limestone, “matorral espinoso”; 1500–2920 m. Mexico. Selected Specimens Examined. Chih (Correll & Gentry 22813, LL, NY, US; Pringle 1440, GH, LL, MEXU, MICH, NY, P, US). Dgo (Corral Dı´az 75 (⫽Worthington 9417), NY, TEX). Hgo (Pringle 13643, MICH, UC, US). Jal (Pringle 11899, MICH, US; Palmer 541, NY, US). Oax (Lorence 4801, ENCB; Windham 527, NY). Pue (Purpus 3147, ARIZ, NY, US). SLP (Johnston et al. 12262, LL). Unverified, Doubtful, or Mistaken Reports. Ags (Siqueiros-Da´vila 4113, 4124, HUAA, cited by Siqueiros-Delgado & Gonza´lez-Adame, 2004 [as Cheilanthes pallens], but not verified). Gto (Rzedowski 4114, IEB, cited by Dı´az- Barriga & Palacios-Rios, 1992, but number is 41146 and specimen is A. incana). Qro (Zamudio 3396, IEB, cited by Dı´az-Barriga & Palacios-Rios, 1992, also cited as A. incana, but specimen is A. limitanea subsp. mexicana).

Argyrochosma pallens is very close to A. incana, but the blades in the former are narrowly oblong (vs. deltate), and the rachises and stipes castaneous, farinose, and bearing sparse scales (vs. black, glabrous, and lacking scales).

10. ARGYROCHOSMA PALMERI Argyrochosma palmeri (Baker) Windham, Amer. Fern J. 77: 41. 1987. Figs. 29G, H, J. Notholaena palmeri Baker in Hooker, Icon. Pl. 17: t. 1678 & text. 1887. Type. Mexico. San Luis Potosı´: mts of San Luis Potosı´, Parry & Palmer 991 (K!; isotypes GH!, NY!, US!).

Rhizomes short, compact, horizontal; rhizome scales lanceolate to linear-lanceolate, concolorous light brown, 3 mm long, entire; fronds to 15 cm long, clumped; stipes 1⁄10–1⁄6 the frond length; stipes and rachises black, terete, glabrous or with only very sparse farina; blades narrowly oblong, mostly bipinnate, to tripinnate at bases; pinnae 8–12 pairs, the proximal somewhat reduced; segments not articulate, mostly oblong, obtuse, the distal sessile, the proximal stalked and with the dark sclerotic color of stalks entering their bases; adaxial surfaces naked to sparsely whitefarinose, especially near margins; abaxial surfaces densely farinose; sori on nearly the whole length of the veins, segment margins plane to slightly recurved, not modified; sporangia 64spored.

Distribution. Rocky outcrops in oak forests; 2450 m. Mexico. Specimens Examined. Dgo (Correll & Correll 20085, LL, NY). Gto (Correll & Correll 28817, LL, MEXU, MICH, US; Dı´az B. & Garcı´a 7181, IEB; Pray 3078, LAM; Zamudio 2523, IEB). SLP (only the type collection).

This species closely resembles A. pallens in the oblong blades and short stipes, but differs in the blackish stipes and rachises and lack of scales on the rachises (vs. castaneous to dark castaneous stipes and rachises and the farinose-glandular rachises with a few small, reddish scales).

11. ARGYROCHOSMA PENINSULARIS Argyrochosma peninsularis (Maxon & Weath.) Windham, Amer. Fern J. 77: 41. 1987. Figs. 29A–C. Notholaena peninsularis Maxon & Weath., Contr. Gray Herb. 127: 15. 1939. Type. Mexico. Baja California Sur: Sierra de la Laguna, from San Bernardo to El Sanz, Nelson & Goldman 7430 (US!, photo US).

Rhizomes short, compact, horizontal or ascending; rhizome scales linear to lanceolate, orange-brown, 3–5 mm long, entire; fronds 18–37 cm long, clumped; stipes ca. 2⁄5 the frond length, atropurpureous, lustrous, terete, more or less scaly proximally; blades deltate to oblong-lanceolate, bipinnate-pinnatifid; rachises sparingly farinose and with sparse linear-lanceolate scales ca. 1 mm long; pinnae 7–12 pairs, narrowly oblong, short-petiolulate, the basal pair often slightly reduced; segments mostly oblong to ovate, not articulate, the dark color of the stalks passing into the base of segments, rarely stopping abruptly at bases, lacking hairs; sori on the distal 1⁄2–1⁄3 of the veins; segment margins plane, unmodified; sporangia 64-spored.

ASPIDOTIS

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Distribution. Dry rocky places, granite rocks; 1200–2150 m. Mexico.

Distribution. Shaded cliffs and rock outcrops; ca. 1900 m. Mexico.

Selected Specimens Examined. BCS (Brandegee 650, GH, NY, UC; Breedlove 43297, CAS, ENCB; Carter 2329, UC, 2408, MEXU, UC, US, 2684, UC; Jones 24154, ARIZ, MICH, NY, UC, US, 24561, US; Moran 18956, UC; Nelson & Goldman 7430, US; Pray 1771, LAM).

Specimens Examined. Mor (Lyonnet 2570, US; Ramı´rez s.n., 10 Aug 1947, MEXU; Sa´nchez 316, MEXU).

Argyrochosma peninsularis is distinct by its non-articulate segments, black stipes and rachises, rachises farinose and bearing a few scales, and most primary pinnules sessile or short-petiolulate.

12. ARGYROCHOSMA PILIFERA Argyrochosma pilifera (R. M. Tryon) Windham, Amer. Fern J. 77: 41. 1987. Figs. 29D–F. Notholaena pilifera R. M. Tryon, Contr. Gray Herb. 179: 79. 1956. Type. Mexico. Morelos: Pueblo del Tepozteco, Sa´nchez S. 98 (US!; isotypes GH!, US!).

Rhizomes short, compact, horizontal to erect; rhizome scales linear, orange-brown to dark brown, 4 mm long, entire; fronds to 25 cm long, clumped; stipes ca. 2⁄5 the frond length, atropurpureous, terete, with sparse lax hairs; blades lanceolate, bipinnate to bipinnate-pinnatifid; pinnae 8–10 pairs, opposite or subopposite; segments oblong, lobed, 6 ⫻ 4 mm long, dark stalk color entering segment bases; adaxial surfaces glabrous; abaxial surfaces white-farinose, with scarce to scattered multicellular hairs 1–2 mm long on rachises and abaxial surfaces; sori along distal 1⁄2 or more of vein length, segment margins plane, not recurved, not modified.

This species is related to A. palmeri, A. pallens, and A. peninsularis, but is readily distinguished by the presence of hairs on the stipes, rachises, and abaxial blade surfaces.

1 3. A SP I D O T I S Aspidotis (Nutt. ex Hook. & Baker) Copel., Gen. Fil. 68. 1947. Hypolepis Bernh. sect. Aspidotis Nutt. ex Hook. & Baker, Syn. Fil. 4: 131. 1867. Type: Hypolepis californica Hook. ⬅ Aspidotis californica (Hook.) Nutt. ex Copel. Terrestrial or epipetric; rhizomes compact, ascending, branched; rhizome scales linear-lanceolate, mostly concolorous dark brown, margins entire, often narrow, pale; fronds 8–35 cm long, monomorphic to slightly dimorphic, clumped; stipes usually dark reddish brown, grooved adaxially, glabrous; blades deltate or pentagonal, 3–5-pinnate, thick to thin, abaxially glabrous, adaxially lustrous, glabrous; ultimate segments short-stalked or with base narrowed and decurrent onto costae or costules, linear to lanceolate, acuminate, mostly 0.5–1.3 mm wide; veins of ultimate segments obscure, free, unbranched; sori marginal; indusia scarious, whitish, broad, round, discontinuous (in ours) or continuous; sporangia 64-spored, lacking paraphyses and glands; spores dark brown, reticulate; x⫽30. Aspidotis is a genus of four species, two of them in Mexico. They occur at bases of boulders or in rock crevices, in dry to moist, montane areas, woodlands, or chaparral, sometimes on ultramafic rocks. Lellinger (1968) recognized Aspidotis as separate from Cheilanthes based on its elongate, distantly dentate segments with striate lustrous surfaces and broad, scarious indusia. Molecular data (Gastony & Rollo, 1995) show that Aspidotis is sister to Aleuritopteris (e.g., Cheilanthes farinosa) and Sinopteris (e.g., Cheilanthes albofusca Baker, of China).

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ASPLENIUM References

Lellinger, D. B. 1968. A note on Aspidotis. Amer. Fern J. 58: 140-141; Smith, A. R. 1975. The California species of Aspidotis. Madron˜o 23: 15–24.

Key to the Mexican Species of Aspidotis 1. Sori lateral on ultimate segments; BCN. .................... 1. A. californica. 1. Sori terminal on ultimate segments; non-peninsular northern Mexico. ............................................................... 2. A. meifolia.

1. ASPIDOTIS CALIFORNICA Aspidotis californica (Hook.) Nutt. ex Copel., Gen. Fil. 68. 1947. Figs. 30G, H. Hypolepis californica Hook., Sp. Fil. 2: 71, t. 88a. 1852; Cheilanthes californica (Hook.) Mett., Abh. Senckenberg. Naturf. Ges. 3: 88. 1859. Syntypes. U.S.A. California: Santa Barbara, Nuttall s.n., Coulter s.n. (K).

Rhizomes compact, ascending; rhizome scales dark reddish brown to black, linear-lanceolate, 2 mm, entire; fronds 10–35 cm, clumped; stipes 1⁄2–3⁄4 the frond length; blades 3–4(–4.5)-pinnate, nearly as wide as long, chartaceous; ultimate segments oblanceolate to rhombic, toothed to entire, 1.5–4 mm long; sori short, discrete, lateral 1–3(–5) on ultimate segments; indusia semicircular, margins nearly entire; 2n⫽60, 120 (USA).

Distribution. Limestone cliffs and slopes; (320–)750–2000 m. Mexico. Selected Specimens Examined. Chih (Pringle 441, DS p.p.). Coah (Wynd & Mueller 350, NY, US). Dgo (Henrickson 12430, LL). NL (Muller 2700, US; Pennell 17171, US; Pringle 1987, NY, UC, US). Qro (Carranza 1474, ENCB). SLP (Palmer 264, NY, UC, US; Pennell 17972, US). Tam (Bartlett 10048, US; Gonza´lez Medrano 14568, MEXU; Hellwig 1206, NY; Palmer 567, US; Stanford et al. 2062, NY).

1 4 . A SP L E N I U M Asplenium L., Sp. Pl. 2: 1078. 1753. Lectotype (chosen by J. Smith, Hist. Fil. 316. 1875): Asplenium marinum L. Phyllitis Hill, Brit. Herb. 525, t. 74. 1757. Type: Phyllitis scolopendrium (L.) Newman [⬅ Asplenium scolopendrium L.]. See Pichi Sermolli, 1953, Webbia 9: 431–433, for another opinion. Scolopendrium Adans., Fam. Pl. 2: 20. 1763. Type: Scolopendrium vulgare J. Sm. [⫽ Asplenium scolopendrium L.].

Distribution. Rocky outcrops and crevices, shaded slopes and cliffs; 20–1300 m. USA (Calif); Mexico. Specimens Examined. BCN (Knobloch 2259A, MSC; McKeever 4, US; Montgomery 83-35, NY; Moran 23212, CAS, 26687, ARIZ, MICH; Schoenfeldt 3755, US; Yatskievych 82-202, ARIZ).

2. ASPIDOTIS MEIFOLIA Aspidotis meifolia (D. C. Eaton) Pic. Serm., Webbia 7: 327. 1950. Figs. 30A–F. Cheilanthes meifolia D. C. Eaton, Proc. Amer. Acad. Arts 18: 185. 1883. Type. Mexico. Nuevo Leo´n: Guajuco, 27 mi SE of Monterrey, Palmer 1377 (YU!, photo US; isotypes NY!, US!).

Similar to A. californica but with blades more dissected (4– 5-pinnate), sori terminal on the ultimate segments (vs. lateral), and broader distribution in northern Mexico; 2n⫽60 (Tam).

Terrestrial, epipetric, or epiphytic; rhizomes usually compact, very short-creeping to suberect or erect, rarely long-creeping, usually unbranched, scaly; rhizome scales usually concolorous, clathrate, lanceolate; fronds often clumped (occasionally spaced), usually monomorphic (rarely slightly dimorphic, as in A. dentatum), small to medium-sized, rarely large, erect, spreading, pendent, or flat on ground or rocks; stipes green to atropurpureous or black, dull to lustrous, often slightly winged; blades simple to 3-pinnate, most often 1-pinnate; veins free, stopping short of margins in clavate tips (as seen adaxially); indument on both sides seemingly absent in most species, but blades in most (all?) species with minute appressed whitish to brownish, clavate, multicellular (3–6 very short cells sometimes with a terminal enlarged glandular cell) hairs 0.1–0.3 mm between veins, rarely with longer hyaline septate hairs on axes (A. pumilum), sometimes with a few narrow, hair-like scales on rachises, in a few species the scales more numerous, broader, and decidedly clathrate; sori abaxial, along the veins, usually one per ultimate vein, rarely back-to-back or opposing one another (A. scolopendrium), oblong to linear, indusiate; indusia persistent, extending the length of sori, whitish when young, tan to light brown with age, thin, margins entire to erose, occasionally ciliate or fimbriate; sporangia with slender, uniseriate stalks; spores bilateral, brown

ASPLENIUM

to blackish, with distinct perispore, usually 64 per sporangium (32 and globose in apogamous spp.); x⫽36 (39 or occasionally 38 in species of sect. Hymenasplenium). Asplenium is a cosmopolitan genus of about 700 species, mostly of the tropics, 86 in Mexico. It includes some of the most difficult taxonomic problems for Mexican ferns. Hybridization is common among the temperate aspleniums and may also be common among tropical species, but there are no studies that document this in the Neotropics. In addition apogamy appears to be prevalent in Asplenium, as judged by life cycle studies on temperate and a few tropical groups, as well as spore morphology and number per sporangium, often 32 globose spores in apogamous species or specimens, instead of 64 reniform spores within a sporangium in sexually reproducing species (Braithwaite, 1986; Murakami & Moran, 1993). In studies on the A. aethiopicum group in Africa (closely related to A. praemorsum in Mexico), Braithwaite suggested that the apomictic taxa were able to tolerate more extreme environmental conditions than their sexual progenitors. We include minimal, but incomplete observational data on spore morphology and number per sporangium in the species descriptions below, as an impetus to further work. One of the characters sometimes used in descriptions of spleenworts concerns the presence or absence and color of scattered whitish to tan (or sometimes brown) hairs, on the abaxial lamina, on and between the veins, as well as sometimes on the stipes and rachises. On casual inspection, the laminae of most aspleniums appears perfectly glabrous, but a closer look reveals that they bear inconspicuous, multicellular (3–6 cells), uniseriate, usually closely appressed hairs mostly 0.1–0.3 mm long on the abaxial surfaces. We have found such hairs in nearly all species examined from Mexico, and suspect that this feature is an autapomorphy for the entire family. Consequently, it seems to us of little use in characterizing individual species, unless differences in cell number, density, presence or absence of a darkened glandular tip, or color are diagnostic. We have so far not found this to be the case. Several groups of Asplenium are well represented in the Mexican fern flora, in particular, the A. monanthes group, with 23 species. This number makes Mexico probably the center of greatest species diversity in the world in this cosmopolitan group. Other well represented groups are A. miradorense and allies, with ca. 13 species, and A. cuspidatum/auritum and allies, with 6 species. Several distinctive predominantly extraterritorial groups are also represented, e.g., A. exiguum and A. dalhousiae, both closely related to, or the same as, species in the Himalayas. In addition, there is the occurrence of several predominantly temperate species, especially in northern Mexico: A. chihuahuense, A. scolopendrium, A. septentrionale, and A. trichomanes. Phyllitis has traditionally been recognized as genus of 3 (to 5) species, largely of temperate regions, in Europe (common), Asia, North America (rare), and the Neotropics (rare in southern Mexico and Hispaniola). The species are difficult to distinguish. The genus is marginally distinct from Asplenium, and there are several intergeneric hybrids, a primary reason for its inclusion in Asplenium in this treatment. Recent molecular work by Schnei-

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der et al. (ms.) and by Van den Heede et al. (2003) indicates that Phyllitis is nested deeply within Asplenium, and allied to A. dalhousiae (often treated in the segregate and polyphyletic genus Ceterach). Phyllitis is traditionally distinguished by its simple elongate blades (also in some aspleniums) and opposing pairs (opening towards each other) of sori. Two other genera of Mexican ferns, both rare and with undivided blades, are closely allied to Asplenium. Holodictyum can be easily distinguished by its net venation, and Schaffneria by its simple, round blades with lustrous atropurpureous to black stipes and paired sori facing each other from adjacent veins. These two genera are kept separate in this treatment because their relationship to and origin from any extant species is obscure and because they have been regarded as distinct in many other taxonomic accounts, rather than through any conviction that they deserve generic rank. Phylogenetic studies on Asplenium and the Aspleniaceae are ongoing, but preliminary evidence suggests that the group diverged from a larger cluster of families and genera that include the athyrioid, thelypteroid, onocleoid, and blechnoid ferns. The family Aspleniaceae in the sense maintained by most systematists (e.g., Copeland, 1947; Tryon & Tryon, 1982; Kramer & Viane in Kubitzki, 1990) is clearly monophyletic, on the basis of both molecular and morphological evidence, but the delimitation of genera within the family is still unsettled and likely to undergo change. It appears now that the only element that can be clearly separated at generic rank, apart from Asplenium s.l., is Hymenasplenium Hayata, distinguished in part by its different chromosome base numbers (x⫽38, 39), and creeping rhizomes (see Mitui et al., 1989; Murakami & Moran, 1993; Schneider et al., 2002). References Braithwaite, A. F. 1986. The Asplenium aethiopicum complex in South Africa. Bot. J. Linn. Soc. 93: 343–378; Hinton, J. & G. S. Hinton. 1995. Checklist of Hinton’s collections of the flora of south-central Nuevo Leo´n and adjacent Coahuila. Acta Bot. Mex. 30: 41–112; Lo¨ve, A. & D. Lo¨ve. 1973. Cytotaxonomy of the boreal taxa of Phyllitis. Acta Bot. Acad. Sci. Hung. 19: 201–206; Lovis, J. D. 1964. The taxonomy of Asplenium trichomanes in Europe. Brit. Fern Gaz. 9: 147– 160; Maxon, W. R. 1913. Studies of tropical American ferns– No. 4. Asplenium trichomanes and its American allies. Contr. U.S. Natl. Herb. 17: 134–153; Mickel, J. T. 1976. Vegetative propagation in Asplenium exiguum. Amer. Fern J. 66: 81– 82; Mitui, K., N. Murakami & K. Iwatsuki. 1989. Chromosomes and systematics of Asplenium sect. Hymenasplenium (Aspleniaceae). Amer. J. Bot. 76: 1689– 1697; Morton, C. V. & D. B. Lellinger. 1966. The Polypodiaceae subfamily Asplenioideae in Venezuela. Mem. New York Bot. Gard. 15: 1–49; Murakami, N. & B. A. Schaal. 1994. Chloroplast DNA variation and the phylogeny of Asplenium sect. Hymenasplenium (Aspleniaceae) in the New World tropics. J. Plant Res. 107: 245–251; Murakami, N., S. Nogami, M. Watanabe & K. Iwatsuki. 1999. Phylogeny of Aspleniaceae inferred from rbcL nucleotide sequences. Amer. Fern J. 89: 232–243; Murakami, N. & R. C. Moran. 1993. Monograph of the neotropical species of Asplenium sect. Hymenasplenium (Aspleniaceae). Ann. Missouri Bot. Gard. 80: 1–38; Rasbach, H., T. Reichstein & R. L. L. Viane. 1994. Asplenium chihuahuense (Aspleniaceae, Pteridophyta), an allohexaploid species and the description of a simplified hybridization technique. Amer. Fern J. 84: 11–40; Reichstein, T. 1981. Hybrids in European Aspleniaceae (Pteridophyta). Bot. Helv. 91: 89–139; Rodrı´guez-Lo´pez, R. & M. L. Arreguı´nSa´nchez. 1992. Adiciones a la pteridoflora del estado de Nuevo Leo´n, Me´xico. II. Anales Esc. Nac. Ci. Biol. 37: 27–34; Schneider, H., S. J. Russell, C. J. Cox, F. Bakker, S. Henderson, M. Gibby & J. C. Vogel. 2002. Chloroplast phylog-

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ASPLENIUM

eny of asplenioid ferns based on rbcL and trnL-F spacer sequences (Polypodiidae, Aspleniaceae) and its implications for the biogeography of these ferns. ms. in prep; Smith, A. R. 1976 [1977]. Diplazium delitescens and the neotropical species of Asplenium sect. Hymenasplenium. Amer. Fern J. 66: 116–120; Stolze, R. G. 1986. 14(6). Polypodiaceae-Asplenioideae. In: G. Harling & B. Sparre (eds.) Flora of Ecuador 23: 1–83. Swedish Research Councils, Stockholm.; Van den Heede,

D. J., R. L. L. Viane & M. W. Chase. 2003. Phylogenetic analysis of Asplenium subgenus Ceterach (Pteridophyta: Aspleniaceae) based on plastid and nuclear ribosomal ITS DNA sequences. Amer. J. Bot. 90: 481–493; Viane, R. L. L. & T. Reichstein. 1991. Notes about Asplenium II: Some new names and combinations in Asplenium L. (Aspleniaceae, Pteridophyta). Dodonaea 59: 157–165; Windham, M. D. 1983. The ferns of Elden Mountain, Arizona. Amer. Fern J. 73: 85–93.

Key to the Mexican Species of Asplenium 1. Blades undivided, shallowly pinnatifid, or 1-forked near the tips. 2. Blades shallowly pinnatifid. ................................................................................................................................................................... 17. A. dalhousiae. 2. Blades undivided or 1-forked near the tips. 3. Blades, or divisions of blades, 1–2 mm wide. ........................................................................................................................... 69. A. septentrionale. 3. Blades 10 mm wide or more. 4. Blades orbicular to flabellate or obovate, about as broad as long; stipes blackish, lustrous. ...................................... Schaffneria nigripes. 4. Blades elliptic or linear-lanceolate; stipes (if present) greenish to tan or brownish, dull. 5. Veins netted. .......................................................................................................................................................... Holodictyum ghiesbreghtii. 5. Veins free. 6. Sori paired, those of a pair overlapping when young, opening toward one another; blades cordate at bases. .................................................................................................................................................... 66. A. scolopendrium var. americanum.

6.

Sori not paired, not overlapping, all opening toward blade apices; blades cuneate and attenuate at bases. ........................................................................................................................................................................................71. A. serratum.

1. Blades 1–3-pinnate, or more divided. 7. Blades with whitish catenate hairs 0.5–0.8 mm long on stipes, rachises, and veins of both surfaces; indusia erose with longciliate margins; blades deltate to tripartite, proximal pinnae basiscopically more developed. ....................................................... 59. A. pumilum. 7. Blades without long whitish catenate hairs, glabrous or with short clavate hairs 0.1–0.2(–0.3) mm on abaxial blade surfaces; indusia entire or erose, usually without cilia; blades deltate, ovate, lanceolate, linear, or elliptic. 8. Stipes and rachises lustrous, castaneous to atropurpureous or blackish. 9. Blades pinnate-pinnatifid to 3-pinnate. 10. Blades broadly deltate with proximal pinna pair basiscopically developed; indusia erose. 11. Pinnae deeply lacerate, segment tips toothed or lacerate; w. Mexico. .................................................................. 4. A. arcanum. 11. Pinnae lobed, the lobes rounded; e. and s. Mexico. ........................................................................................... 40. A. minimum. 10. Blades narrowly deltate, ovate, lanceolate, or linear, proximal pinna pair not strongly basiscopically developed; indusia erose or entire. 12. Fronds 4–7 cm long, 3–4-pinnate; pinnae 3–5 pairs; blades deltate, not proliferous. ................................. 77. A. tenerrimum. 12. Fronds over 6 cm long, pinnate-pinnatifid to 3-pinnate; pinnae 10 or more pairs per blade; blades deltate or usually linear to narrowly elliptic, or proximal pinnae greatly reduced, with proliferous buds subterminal on rachises, on flagelliform apices, or at emarginate pinna apices. 13. Blades 5–14 ⫻ 1.5–3.5 cm; pinna apices emarginate with small proliferous buds (often dormant). ...... 24. A. exiguum. 13. Blades 14–50 ⫻ 2.5–12 cm (if less than 4 cm wide, then fronds 25–50 cm long); pinna apices rounded to acute, margins not notched and without buds at apices. 14. Proliferous buds subterminal on rachises. 15. Blades 2-pinnate-pinnatifid to 3-pinnate, elliptic; ultimate segments falcate, 1 mm wide, each with a single vein. .............................................................................................................................45. A. muenchii. 15. Blades pinnate-pinnatifid, linear; ultimate segments multiveined, each pinna with a single (or two) oval or flabellate, nearly free, acroscopic basal segment(s), rest of pinna shallowly lobed. .......................................................................................................................................................... 72. A. sessilifolium.

9.

14. Proliferous buds terminating long, flagelliform blade apices. 16. Blades elliptic, 3-pinnate, with proximal pinnae greatly reduced. ............................................ 63. A. rutaceum. 16. Blades deltate, pinnatifid to 3-pinnate, with proximal pinnae not or only slightly reduced. 17. Blades deeply pinnate-pinnatifid to barely 2-pinnate, basal pinnules toothed; rhizome scales 2–3 mm long; indusia 4–5 mm long. ......................................................................... 60. A. radicans. 17. Blades 2-pinnate-pinnatifid to 3-pinnate, with basal pinnules lobed or deeply dentate, or more divided; rhizome scales 3–5 mm long; indusia 1–4 mm long. 18. Blades 2-pinnate-pinnatifid to barely 3-pinnate, with basal acroscopic pinnules lobed, each with a single free basal segment; indusia 1–2 mm long. ................ 27. A. flabellulatum. 18. Blades fully 3-pinnate, with basal acroscopic pinnules deeply lobed, resulting in linear-oblong to obovate segments ca. 1 mm wide; indusia 2–4 mm long. ........... 82. A. uniseriale. Blades 1-pinnate (rarely pinnatifid in depauperate forms of A. minimum), pinnae sometimes with strongly dentate margins. 19. Blades ternately-lobed or ternate with basiscopic pinnules of proximal pinnae more developed; indusia erose. ............................................................................................................................................................................................. 40. A. minimum.

ASPLENIUM

75

19. Blades linear, pinnate with many pinnae, not ternate; indusia erose to entire. 20. Blades 4–8(–12) cm wide. 21. Blades with flagelliform, proliferous apices; plants terrestrial. 22. Pinnae 1.5–3(–4) cm long, not to slightly auriculate, margins scarcely crenate-serrate; sori 3–9 pairs per pinna; indusia 2–4(–6) mm long. ................................................................................................... 13. A. cirrhatum. 22. Pinnae 3.5–6 cm long, strongly auriculate, margins sharply serrate with sinuses 0.5–1 mm deep; sori 7–11 pairs per pinna; indusia 6–9 mm long. .............................................................................. 86. A. yelagagense. 21. Blades with nonproliferous, pinnatifid or conform apices; plants terrestrial, epipetric, or epiphytic. 23. Blades firm, veins obscure; pinnae 4–25 pairs, with entire to crenulate margins; indusia 1 mm wide, subentire; sori 1–5 pairs per pinna; rachises adaxially with brown wings; rhizome scales bicolorous with central, thickened sclerotic portion. 24. Pinnae 12–25 pairs, 2–3(–4) ⫻ 0.7–1.5 cm; sori 1–4 pairs per pinna. ................................ 39. A. lamprocaulon. 24. Pinnae 4–9 pairs, 4–5 ⫻ 2–2.5 cm; sori 4–5 pairs per pinna. ...................................................... 48. A. obesum. 23. Blades chartaceous to membranaceous, veins evident; pinnae 10–30 pairs or more, with deeply crenately or serrately toothed margins; indusia 0.3–0.8 mm wide, entire to erose; sori 2–13 pairs per pinna; rachises adaxially with green wings; rhizome scales concolorous, clathrate. 25. Indusia entire, 2–4 mm long; abaxial blade surfaces glabrous; blades generally with over 30 pinna pairs, proximal pinnae reduced; pinna apices long-acuminate; rhizome scales 6–10 mm long, plants epiphytic. ..................................................................................................................... 32. A. harpeodes. 25. Indusia erose, 4–10 mm long; abaxial blade surfaces often with numerous white clavate hairs 0.1 mm long; blades with 12–22 pinna pairs, proximal pinnae not reduced; pinna apices acute to obtuse; rhizome scales 1–2.5 mm long; plants terrestrial or epipetric. .................................... 38. A. laetum. 20. Blades 0.5–3(–3.5) cm wide. 26. Vein endings obscure or not at all visible on adaxial blade surfaces; rachises blackish; rhizome scales bicolorous with black occluded center; indusia 0.8–1 mm long. 27. Pinnae (4–)10–20 ⫻ 2–5 mm, subcoriaceous, deflexed, especially at bases of blades; pinna margins entire to crenate, sinuses less than 0.3 mm deep; veins 1-forked; rhizome scales 3–5 mm long; fronds erect. ................................................................................................................................................. 61. A. resiliens. 27. Pinnae 3–10 ⫻ 1.5–3(–4) mm, thin-textured, spreading; pinna margins crenate to serrate; veins simple or forked; rhizome scales 1–2 mm long; fronds arching. 28. Rachises extended as flagelliform tips and ending in a bud on some fronds. ............................ 51. A. palmeri. 28. Rachises not flagelliform and bearing a bud at tips. 29. Fertile veins (some or all, exclusive of the basal ones) 1-forked; pinnae mainly 4–7 mm long, length 1–2 times width. ............................................................................................ 78. A. trichomanes. 29. Fertile veins (basal acroscopic ones excepted) usually simple; pinnae mainly 5–13 mm long, length usually 2–5 times width. 30. Stipes and rachises castaneous or distinctly brownish; pinna margins deeply lobed, especially acroscopic and distal margins, cut 1⁄3 or more of distance from margins to costae; BCN. ............................................................................................................... 84. A. vespertinum. 30. Stipes and rachises black or blackish; pinna margins crenate to serrate; ne to s Mexico. 31. Blades thin-chartaceous, the veins readily visible in transmitted light; pinnae sharply crenate-dentate. .................................................................................... 34. A. heterochroum. 31. Blades chartaceous, the veins not visible in transmitted light; pinnae shallowly crenate. .................................................................................................................... 47. A. nesioticum. 26. Veins endings evident on adaxial blade surfaces; rachises blackish, atropurpureous, castaneous, or light brown; rhizome scales clathrate or bicolorous with black occluded centers; indusia 1–4 mm long. 32. Indusia conspicuously ciliate or laciniate. 33. Veins 1-forked; indusia delicately ciliate; rachises with sparse hair-like scales. ...................... 8. A. blepharodes. 33. Veins simple; indusia deeply laciniate; rachises distinctly and noticeably fibrillose. ........... 26. A. fibrillosum. 32. Indusia entire to crenulate or somewhat erose, but not ciliate. 34. Pinnae fewer than 20 pairs per frond; rachises wingless or with wings to 0.1 mm wide, flanking adaxial groove (except sometimes A. castaneum, A. oligosorum). 35. Pinnae 2–5 ⫻ 1–3.5 mm; indusia 1–2 ⫻ 0.5–1 mm, entire; proliferous buds present, subterminal on rachises or in axils of proximal pinnae; rhizome scales with lumina occluded or not. 36. Blades rooting from buds borne at bases of terminal segments, or from buds borne on adaxial surfaces of distal pinnae; fronds less than 7 cm long; Chih. .................... 79. A. tryonii. 36. Blades rooting subterminally or proximally; fronds usually more than 7 cm long. 37. Proliferous buds subterminal on rachises, near blade apices; fronds 4–10 cm long;

76

ASPLENIUM rhizome scales 2–4 mm long, lumina not occluded; plants epipetric on limestone. .............................................................................................................. 73. A. soleirolioides. 37. Proliferous buds in axils of proximal pinnae; fronds 2–5(–7) cm long; rhizome scales 0.8–1.5 mm long, lumina occluded; plants epipetric on granite. .......... 76. A. stolonipes. 35. Pinnae 4–7(–10) ⫻ 3–4 mm; indusia 1–4 ⫻ 0.8–1 mm, undulate or erose; proliferous buds absent; rhizome scales clathrate, lumina not occluded. 38. Sori mostly 4–6(–8) per pinna. 39. Pinnae subentire to undulate to faintly crenulate at their tips and along acroscopic margins; blades dwindling into small terminal segments less than 5 ⫻ 2 mm (appearing indeterminate in many specimens). ....................................... 11. A. castaneum. 39. Pinnae dentate or strongly crenulate at their tips and along acroscopic margins; blades ending in rather large triangular or hastate terminal segments 5–10 ⫻ 4– 10 mm. ....................................................................................................................... 65. A. sanchezii. 38. Sori 1 or 2 per pinna. 40. Pinnae overlapping rachises, margins green; rhizome scales 1–1.5 ⫻ 0.1–0.3 mm. .................................................................................................................................. 49. A. oligosorum.

40. Pinnae not overlapping rachises, margins hyaline; rhizome scales 2–3 ⫻ 0.5–0.8 mm. .......................................................................................................................... 50. A. olivaceum. 34. Pinnae 20 or more pairs per blade; rachises with wings 0.1–0.3 mm wide flanking adaxial groove. 41. Rhizome scales clathrate, lumina not occluded; rachises castaneous to light brown. 42. Sori supramedial to submarginal, extending part way into prominent teeth or lobes. ............................................................................................................................................... 55. A. pringlei.

42. Sori inframedial to medial, not extending into teeth or lobes. 43. Blades nonproliferous; pinnae nearly round or rhomboid, 6–10 ⫻ 5–6 mm; indusia erose. ............................................................................................................ 11. A. castaneum. 43. Blades each with subterminal proliferous bud; pinnae oblong, 8–10(–22) ⫻ 4–6 mm; indusia subentire. ........................................................................................ 52. A. polyphyllum. 41. Rhizome scales bicolorous, with dark, sclerotic central portions; rachises black to castaneous. 44. Acroscopic margins of pinnae deeply (1–3 mm) incised or lacerate, bidentate; sori usually confined to basiscopic side of a pinna, parallel to the margin; rhizome scales 1.5–2 mm long. ................................................................................................................ 28. A. formosum. 44. Acroscopic margins of pinnae entire or shallowly (less than 1 mm) toothed, dentate; sori only along basiscopic sides of pinnae or along both acroscopic and basiscopic sides. 45. Rachis wings papillate; rhizome scales 1–2 mm long; indusia suberose to erose. ..................................................................................................................................... 31. A. hallbergii.

45. Rachis wings entire or slightly papillate; rhizome scales 5–6 mm long; indusia entire. 46. Stipes castaneous or reddish brown; sori usually on only basiscopic sides of pinnae; widespread and common throughout Mexico. .......................... 43. A. monanthes. 46. Stipes atropurpureous to blackish; sori on both basiscopic and acroscopic sides of pinnae; nw Mexico. .............................................................................. 30. A. gentryi. 8. Stipes and rachises dull, green, stramineous, or brownish, rarely atropurpureous or black, if dark and lustrous then only at stipe bases. 47. Blades 1-pinnate; pinnae entire, toothed, or lacerate to ca. half their width, each pinna sometimes with a basal acroscopic auricle. 48. Blades 1–2.5 cm wide. 49. Fronds subdimorphic, the sterile forming a rosette, spreading or arching, the fertile fronds central and erect, longer-stiped. .............................................................................................................................................................. 20. A. dentatum. 49. Fronds monomorphic, the sterile and fertile similar in shape and size. 50. Pinnae 1–5 lateral pairs; terminal segments broadly hastate-deltate. ............................................................. 85. A. virillae. 50. Pinnae more than 5 lateral pairs; terminal segments pinnatifid or lobulate, or apices flagelliform. 51. Blades with flagelliform apices; largest pinnae entire to erose. ............................................................. 33. A. harrisii. 51. Blades lacking flagelliform apices; largest pinnae crenulate. 52. Pinnae 30–40 pairs. .............................................................................................................................. 67. A. seileri. 52. Pinnae 6–25 pairs. 53. Pinnae 15–20(–25) pairs; stipes brownish or gray-brown; proximal pinnae not lobed. ... 15. A. cruegeri. 53. Pinnae 6–8 pairs; stipes green-stramineous; proximal pinnae pinnately divided or deeply lobed. ....................................................................................................................................... 42. A. modestum.

ASPLENIUM

77

48. Blades more than 2.5 cm wide. 54. Apices of blades flagelliform and proliferous; rachises broadly alate, wings mostly 1–2 mm wide, or sometimes more; plants terrestrial. ............................................................................................................................... 3. A. alatum. 54. Apices of blades pinnatifid or conform, non-flagelliform; buds terminal, subterminal, or absent; rachises not or only narrowly (⬍ 0.3 mm wide) alate; plants epiphytic, epipetric, or terrestrial. 55. Rhizomes creeping. 56. Stipes and rachises Ⳳ persistently scaly, scales hair-like; pinnae 7–15(–20) ⫻ 1.5–2(–3) cm; plants epiphytic (occasionally epipetric). ................................................................................................................ 70. A. serra. 56. Stipes and rachises Ⳳ glabrous or bearing glandular hairs, scales if present not readily seen without magnification; pinnae usually much smaller; plants terrestrial, epipetric, or epiphytic. 57. Plants epiphytic; pinnae strongly lacerate; sori nearly parallel to costae; stipes with scattered to dense glandular hairs 0.1–0.5 mm long. ....................................................................................... 22. A. dissectum. 57. Plants terrestrial or epipetric; pinnae subentire to shallowly toothed; sori not parallel to costae; stipes glabrous. 58. Blade apices subconform or hastate; spores spiny. .............................................................. 62. A. riparium. 58. Blade apices nonconform, pinnatifid; spores cristate. 59. Free pinna pairs 4–10; rachises green; blades membranaceous. ............................. 35. A. hoffmannii. 59. Free pinna pairs 6–25; rachises greenish brown to atropurpureous; blades chartaceous. 60. Pinnae mostly 7–14 ⫻ 0.8–2 cm, 7–9 pairs per blade; blades broadly deltate. .................................................................................................................................... 19. A. delitescens.

60. Pinnae 2–6 ⫻ 0.4–1.2 cm, 12–22 pairs per blade; blades lanceolate. ................... 38. A. laetum. 55. Rhizomes erect. 61. Pinnae usually 15–40⫹ pairs; indusia entire. 62. Blades deltate, broadest proximally; plants epiphytic, rarely epipetric. 63. Pinnae only slightly auriculate, margins biserrate; indusia 0.3–0.5 mm wide; spores reniform, tan to brown, 64 per sporangium; plants at 100–800 m. .................................. 6. A. auritum. 63. Pinnae each with an acroscopic auricle, often cut almost to costae, margins deeply biserrate to almost 1⁄3 of distance to midvein; indusia 0.5–0.8 mm wide; spores globose, large, dark brown to blackish, 32 per sporangium; plants at 100–1550(–2300) m. ........ 44. A. monodon. 62. Blades linear or with proximal pinnae reduced in size; plants terrestrial, epipetric, or epiphytic. 64. Stipes castaneous to black; rachises with narrow wings 0.2–0.5 mm wide; stipes nonalate. 65. Pinnae 3–7 cm long with long-acuminate tips (1⁄3–1⁄2 of pinna length); pinnae more than 30 pairs; rhizome scales 6–10 mm long; sori 8–15 pairs per pinna; plants epiphytic. ......................................................................................................................... 32. A. harpeodes. 65. Pinnae 1.5–3 cm long with acute tips; pinnae 16–55 pairs; rhizome scales 1.5–3 mm long; sori mostly 5–10 pairs per pinna; plants terrestrial or epipetric. ................. 41. A. miradorense. 64. Stipes stramineous, green, gray-green, or brownish; rachises with broad wings (0.2–)0.5– 0.8 mm wide; stipes alate to bases (wings 0.1–0.6 mm wide). 66. Acroscopic veins simple (except for basal vein); sori absent on basal veins of each pinna; stipes 1⁄6 or less of frond length; plants epiphytic, rarely epipetric. 67. Stipes 0.5–2 cm long; blades mostly 12–25 ⫻ 2.5–5 cm. ............................... 56. A. pseuderectum. 67. Stipes 2–8 cm long; blades 20–40 ⫻ 5–8 cm. ....................................................... 57. A. pteropus. 66. Acroscopic veins 1-forked (to simple near the pinna apices); sori often on basal vein; stipes 1⁄5–1⁄3(–1⁄2) of frond length; plants terrestrial, rarely epiphytic. 68. Pinnae 1.8 cm long or less; pinnae mostly 15–20 pairs; blades only slightly narrowed proximally. .................................................................................................. 15. A. cruegeri. 68. Pinnae mostly 2 cm long or more; pinnae more than 20 pairs; blades distinctly and sometimes strongly narrowed at bases. 69. Pinnae coarsely serrate to biserrate; stipe wings 0.5 mm wide; rhizome scales 3–6 mm long; blades lacking buds. .................................................... 7. A. barbaense. 69. Pinnae serrulate; stipe wings 0.1–0.3 mm wide; rhizome scales 2–3.5 mm long; blades often with a terminal or subterminal rachis bud. ............... 53. A. potosinum. 61. Pinnae (1–)3–14 pairs, if more numerous then blades more than 8 cm wide; blades nonproliferous; indusia entire or suberose. 70. Pinnae 2 cm long or less, lateral pinnae 2–5 pairs. ......................................................................... 85. A. virillae. 70. Pinnae more than 2 cm long or pinna pairs more than 5 lateral pairs, or both. 71. Blade apices conform or subconform (hastate), with a terminal pinna like the lateral ones; indusia entire. 72. Pinnae entire or nearly so; plants epiphytic. ...................................................... 37. A. juglandifolium. 72. Pinnae toothed or lobed; plants terrestrial, epipetric, or epiphytic.

78

ASPLENIUM 73. Blades 25–35(–60) cm long; stipes mostly 20–30 cm long, with numerous, curled scales 3–4 mm long; rhizome scales 5–10 mm long; indusia 3–7 ⫻ 1–2 mm, whitish. ..................................................................................................................... 25. A. feei. 73. Blades 8–20 cm long; stipes 10–18 cm long, lacking scales, sometimes with scattered, appressed, clavate hairs 0.1–0.3 mm long; rhizome scales 2–4 mm long; indusia 5–12 ⫻ 0.3–0.8 mm, whitish or brownish. ....................................... 80. A. tuerckheimii. 71. Blade apices pinnatifid; indusia entire or suberose. 74. Blades thick or fleshy, yellowish green; pinnae stalked 1.5–3 mm, with prominent auricles, often overlapping rachises, subcordate at acroscopic bases; rhizome scales 3– 7 mm long, orange-brown, clathrate, with golden hairs 0.8–2 mm long; indusia 5–6 mm long. 75. Fronds mostly 8–30 cm long; pinnae 1.8–4 cm long, acroscopic margins with 10– 20 teeth, veins simple or 1-forked; sori 4–6 pairs per pinna; indusia entire or with scarious fringe; rhizome scales 3–4 ⫻ 1–2 mm. ....................................... 5. A. auriculatum. 75. Fronds 25–60 cm long; pinnae 5.5–9 cm long, acroscopic margins with (17–)25– 30 teeth, veins 1–2-forked; sori 7–15 pairs per pinna; indusia entire; rhizome scales 4–7 ⫻ 0.4–0.8 mm. ................................................................................... 64. A. salicifolium. 74. Blades chartaceous to membranaceous, green to dark green; pinnae not prominently stalked or auriculate, sessile or nearly so, straight or cuneate at acroscopic bases; rhizome scales 0.5–2.5 mm long, dark brown or blackish, subclathrate or lumina occluded, without hairs; indusia entire or suberose, 1–10 mm long. 76. Pinnae 3–12 cm long; stipes mostly 5–25 cm long, 1⁄3–1⁄2 of frond length; sori mostly 5–14 pairs per pinna; plants terrestrial or epipetric. ................................. 1. A. abscissum. 76. Pinnae 2 cm long or less; stipes 1–6 cm long, 1⁄3 or less of frond length; sori 1–5 pairs per pinna; plants epipetric. ......................................................................... 58. A. pulchellum. 47. Blades pinnate-pinnatifid to 3-pinnate, with at least one acroscopic pinnule on most pinnae cut to costae. 77. Blades strongly narrowed proximally, sometimes to within a few cm of rhizome; stipes 1⁄10–1⁄3 (rarely to 1⁄2) of frond length. 78. Stipes and rachises with dense to scattered, lanceolate to hair-like, clathrate scales 0.8–1 mm long. 79. Buds often present in axils of distal pinnae or at blade tips; rhizome scales 5–15 mm long; rachial and costal scales most often greater than two cells wide; indusia 0.8–1 mm wide; reduced proximal pinnae 0–2 pairs, the smallest usually at least 2⁄3 the size of the longest; s. and e. Mexico. ....................... 9. A. blepharophorum. 79. Buds absent in axils of distal pinnae; rhizome scales 6–8 mm long; rachial and costal scales mostly two cells wide; indusia 1–1.2 mm wide; reduced pinnae 5 or more pairs, the smallest sometimes ca. 1–2 cm long; w. Mexico. .................................................................................................................................................... 23. A. eatonii. 78. Stipes and rachises glabrous; rhizome scales 0.8–5 mm long; indusia 0.3–0.8 mm wide. 80. Rachises not proliferous. 81. Blades 15–50 ⫻ 7–15 cm, 2-pinnate-pinnatifid to 3-pinnate; stipes 6–10(–15) cm long, gray-brown. 81.

....................................................................................................................................................................... 14. A. cristatum. Blades 8–13 ⫻ 2–4 cm, 2–3-pinnate; stipes 1–2 cm long, castaneous to brown.

82. Blades chartaceous, 2-pinnate, ca. 2 cm wide; pinnules strongly adnate, not or only slightly constricted at their bases (except the acroscopic one of each pinna); stipes brownish. ............. 21. A. dianae. 82. Blades thin-herbaceous, ca. 3-pinnate, 2.5–8 cm wide; pinnules constricted or short-stalked at their bases; stipes castaneous. .................................................................................................. 46. A. myriophyllum. 80. Rachises often producing small plants towards tips of blades. 83. Blades 2-pinnate-pinnatifid to 3-pinnate, elliptic; ultimate segments falcate, 1 mm wide with single vein. ............................................................................................................................................................45. A. muenchii. 83. Blades pinnate-pinnatifid to 2-pinnate; ultimate segments (pinnules) with veins pinnate or forked. 84. Blades 2-pinnate, broadly elliptic; pinnae each with 5 or more pairs of Ⳳ free pinnules that are cuneate proximally. ............................................................................................................ 68. A. semipinnatum. 84. Blades pinnate-pinnatifid, narrowly elliptic in outline; pinnae with single (or two) flabellate, nearly free, acroscopic basal segment(s), rest of pinna shallowly to deeply lobed. ............. 72. A. sessilifolium. 77. Blades widest at bases or only slightly narrowed proximally; stipes usually at least 1⁄2 (rarely 1⁄3) of frond length. 85. Fronds usually less than 15(–22) cm long; blades less than 4 cm wide (except sometimes in A. arcanum). 86. Pinnae fewer than 5 pairs per blade; blades broadly deltate. 87. Blades thin-textured, membranaceous; pinnules of proximal pinnae deeply lacerate. ..................... 4. A. arcanum. 87. Blades firm-herbaceous; pinnules of proximal pinnae merely toothed, or subentire. .............. 12. A. chihuahuense. 86. Pinnae 6–20 pairs per blade; blades lanceolate. 88. Proximal pinnae flabellate, pinnately parted into 3 cuneate lobes, which in turn are deeply cleft; Chih. ......................................................................................................................................................... 42. A. modestum. 88. Proximal pinnae pinnate nearly to apices, or pinnatifid; NL, s. Mexico.

ASPLENIUM ABSCISSUM

79

89. Sori more than 6 per pinna, not adjacent and parallel to costae; s. Mexico. ...................... 18. A. delicatulum. 89. Sori to 5 per pinna, all but the acroscopic basal one adjacent and parallel to costae; NL. .... 81. A. ultimum. 85. Fronds usually more than 15 cm long; blades more than 4 cm wide; proximal pinnae often at least twice as long as wide. 90. Rachises and stipes with sparse to dense hair-like clathrate scales 0.5–5 mm long. 91. Proximal several pairs of pinnae stalked 2 mm or more; costae obscure; plants epiphytic or epipetric. ................................................................................................................................................ 54. A. praemorsum. 91. Proximal several pairs of pinnae sessile or stalked less than 1.5 mm; costae evident; plants terrestrial. 92. Pinnules adnate at their bases, sometimes narrowed proximally; subterminal rachis buds absent; blades 2-pinnate, basal pinnules toothed; rachises with scattered to sparse, hair-like clathrate scales 2 cells wide at their bases; rhizome scales 1.5–5 mm long. ......................... 2. A. achilleifolium. 92. Pinnules narrowed (cuneate) at their bases; subterminal rachis buds sometimes present; blades 2-pinnate-pinnatifid to 3-pinnate, basal pinnules with 3–4 deep lobes; rachises with dense to scattered, hair-like clathrate scales 3–4 cells wide at their bases; rhizome scales 5–15 mm long. ..................................................................................................................................................... 9. A. blepharophorum.

90. Rachises and stipes glabrous or glabrescent, or with gland-tipped hairs. 93. Rhizomes creeping; blades 3-pinnate-pinnatifid or more divided; plants terrestrial. ..................... 10. A. breedlovei. 93. Rhizomes suberect to erect; blades 1-pinnate-pinnatifid to 3-pinnate; plants terrestrial, epipetric, or epiphytic. 94. Pinnae sessile, subsessile, or short-stalked to 1(–1.5) mm. 95. Blades 2-pinnate, pinnules merely toothed toward the tips. .............................................. 83. A. venturae. 95. Blades 2-pinnate-pinnatifid to nearly 3-pinnate. 96. Pinnules less than 2 times longer than wide; pinnae parallel-sided for most of their length. ................................................................................................................................ 14. A. cristatum. 96. Pinnules 2–4 times longer than wide; pinnae not distinctly parallel-sided ................ 74. A. solmsii. 94. Pinnae in at least proximal and medial positions stalked at least 2 mm. 97. Rhizomes short-creeping; rhizome scales black, clathrate with occluded lumina, 1–2 mm long; abaxial blade surfaces with scattered, brown, clavate hairs 0.1 mm long; blades 2pinnate-pinnatifid, basal pinnules deeply lobed with flabellate segments; plants terrestrial. ..................................................................................................................................................... 36. A. insolitum.

97. Rhizomes suberect; rhizome scales brown, clathrate with non-occluded lumina, usually greater than 2 mm long; abaxial blade surfaces seemingly glabrous or with rare to scattered, hair-like clathrate scales 0.5–1 mm long; blades pinnate-pinnatifid to 3-pinnate, if 2-pinnate-pinnatifid or 3-pinnate, segments linear or elliptic; plants epiphytic or epipetric, rarely terrestrial. 98. Blades 2-pinnate-pinnatifid, with basal acroscopic pinnule of proximal pinnae having a basal lobe cut almost to costule, to 3-pinnate with linear segments (1 vein per segment, 1 mm wide). ............................................................................................... 29. A. fragrans. 98. Blades pinnate-pinnatifid to 2-pinnate with segments deeply dentate. 99. Spores reniform, tan to brown, 64 per sporangium; blades deeply pinnatepinnatifid to 2-pinnate with dentate segments. ............................................... 16. A. cuspidatum. 99. Spores globose, black to dark brown, 32 per sporangium; blades pinnate and each pinna with a superior auricle, or pinnate-pinnatifid or 2-pinnate proximally. 100. Blades 1-pinnate, each pinna with a superior auricle, or pinnate-pinnatifid at bases of blades with one superior lobe cut nearly to costa. ................. 44. A. monodon. 100. Blades 2-pinnate proximally, to 12 pairs of nearly free, obovate pinnules per pinna. ......................................................................................................... 75. A. sphaerosporum.

1. ASPLENIUM ABSCISSUM Asplenium abscissum Willd., Sp. Pl., ed. 4, 5(1): 321. 1810. Lectotype (chosen by Morton & Lellinger 1966: 26; further restricted by Proctor, Fl. L. Antill. 2: 320. 1977). India Occidentale, collector not stated and not on type, but Bredemeyer s.n., fide Lellinger, pers. comm. (B-Willd. 19893-1; microfiche UC). Fig. 34D. Asplenium polymorphum M. Martens & Galeotti, Me´m. Foug. Mexique 56, pl. 15, f. 2. 1842, non Wall., 1828, nom. nud., nec Mett., 1859. Syntypes. Mex-

ico. Veracruz: Xalapa and Zacuapa´n, Galeotti 6295 (BR!–3 sheets; photo of Zacuapa´n specimen at US!; isosyntype of Xalapa specimen at US!). Additional synonymy given in Morton & Lellinger (1966).

Roots fibrous, not proliferous; rhizomes erect; rhizome scales dark brown, subclathrate, lumina Ⳳ occluded, ovate-lanceolate, ca. 1.5–2.5 ⫻ 0.8 mm, entire; fronds clumped, (5–)10–40(–50) cm long; stipes green to brownish, dull, (2–)5–25 cm ⫻ 0.5–2 mm, 1 ⁄3–1⁄2 of frond length, glabrescent or with a few hair-like scales, adaxially with narrow green wings on shoulders for much of their length; blades chartaceous, deltate, 1-pinnate, (3–)8–25 ⫻ (1.5–)

80

ASPLENIUM

4–16 cm, broadest at the bases or nearly so, apices confluent, pinnatifid, nonproliferous; rachises greenish to yellowish or brownish, dull, with sparse hair-like scales, adaxially with green lateral wings 0.5 mm wide; pinnae linear-lanceolate, (3–) 5–9(–12) pairs, (1–)3–9 ⫻ 0.5–1.8 cm, sessile or largest stalked to 1 mm, not articulate, excavate at basiscopic bases, truncate acroscopically, margins irregularly obtuse to commonly acute or attenuate at apices, margins biserrate or bicrenate, sinuses to ca. 2.5 mm deep; veins mostly 1- or 2-forked, Ⳳ evident on both sides of blades, tips prominent adaxially; indument abaxially of scattered, tan, appressed, clavate hairs 0.1–0.2 mm long; sori (2–) 5–14 pairs per pinna, at ca. 30–45⬚ from costae; indusia 3–6 ⫻ 0.4–0.8 mm, margins entire; spores reniform; 2n⫽72 (Fla, Jam, Trin), 144 (Oax), 288 (PR).

latter lacking green-winged rachises, but this character does not always seem reliable or consistent. In general, blades of A. abscissum have more rhombic, acute-tipped pinnae than in A. virillae, and the blades dry a lighter, yellow-green (vs. dark green), but the two species are closely related and may be merely ecotypes (A. abscissum on soil, A. virillae on rocks) of the same taxon. The lectotype of A. abscissum has been erroneously said to be from Caracas, Venezuela, by Morton and Lellinger (1966), Proctor (1977, 1989), and Adams (in Davidse et al., 1995). The type and presumed isotypes (19893-2, 19893-3) in the Willdenow herbarium clearly state “Habitat in India occidentale,” and sheet 19893-3 is labeled “Porto rico.”

2. ASPLENIUM ACHILLEIFOLIUM Asplenium achilleifolium (M. Martens & Galeotti) Liebm., Mexic. Bregn. 249 (reprint 97). 1849. Fig. 37D. Caenopteris achilleifolia M. Martens & Galeotti, Me´m. Foug. Mexique 63, pl. 16. 1842. Athyrium achilleifolium (M. Martens & Galeotti) Fe´e, Me´m. Foug. 5: 186. 1852. Syntypes. Mexico. Veracruz: Vicinity of Mirador, Galeotti 6279 (BR!; isosyntypes K!, P!, frags. NY!), 6293 (BR!–photo NY!; frag. NY!). Athyrium conchatum Fe´e, Me´m. Foug. 5: 188, t. 17C, f. 1. 1852. Asplenium conchatum (Fe´e) T. Moore, Index Fil. 49. 1857. Type. Mexico. “Ad craterem Cerro Sancti Martini,” Galeotti 6569 (P? not found; isotype K!) Asplenium grande Fe´e, Me´m. Foug. 8: 82. 1857, non Sw., 1806. Type. Mexico. Veracruz: “barranca de San Martin et de San Francisco, pre`s de Mirador,” Schaffner 74 (not found at P).

Distribution. Terrestrial and epipetric, sometimes on logs, in wet forests; 100–2050 m. USA (Fla); Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Ec, Peru, Braz, Bol, Uru. Adams (in Davidse et al., 1995) also reported French Guiana and Paraguay in the range. Selected Specimens Examined. Chis (Breedlove 22475, DS, NY, 57998, CAS). Hgo (Kenoyer s.n., ARIZ). Jal (McVaugh 14230, MEXU, US). Mor (Hinton 17091, ENCB, NY). Nay (Jones 23478, UC p.p.). Oax (Mickel 6382, MEXU, NY, UC). Tab (Cowan 3157, CAS, ENCB, NY). Ver (Purpus 6450, UC, US; Wendt et al. 3472, CHAPA, NY). Unverified, Doubtful, or Mistaken Reports. SLP (reported by Mickel & Beitel, 1988, but not verified).

Asplenium abscissum is easily confused with A. delitescens, but the former is distinct in its erect rhizomes, entire rhizome scales, winged rachises, fewer (5–14 pairs) sori per pinna, and shorter (3–6 mm) indusia. In all likelihood the species as broadly construed here is a complex, as suggested by the different ploidy levels known. Hybrids with A. verecundum Chapman ex Underw. are known in Florida. Wendt et al. 3472, from Veracruz, and several specimens from Chiapas (e.g., Breedlove 22251, 22508, DS) have precociously fertile fronds less than ca. 8 ⫻ 2 cm and blades with 3 or 4 pinna pairs. Similar specimens are often annotated as A. pulchellum or A. virillae in herbaria, and the latter is construed by Adams (in Davidse et al., 1995) more broadly, to include very small plants of what we identify as A. abscissum (e.g., Breedlove 22475, 33105, DS). Adams distinguished A. virillae from A. abscissum by the

Roots fibrous, not proliferous; rhizomes suberect; rhizome scales brown or often black near tips, clathrate, long-attenuate, 1.5–5 ⫻ 0.8–2 mm, entire; fronds clumped, mostly 50–120 cm; stipes greenish gray, dull, (12–)20–40 cm ⫻ 2–5 mm, 1⁄3–1⁄2 of frond length, with scattered, hair-like, clathrate scales, 0.5–1 mm long, scales 2 cells wide at bases, lacking green wings; blades chartaceous to membranaceous, ovate-lanceolate, 15–80 ⫻ 8–26 cm, 2-pinnate with wings 0.5–0.8 mm wide on pinnae, proximal pinnae slightly reduced, apices pinnatifid, nonproliferous; rachises similar to stipes in color, bearing numerous linear or hairlike scales mostly 0.5–1.5 mm long, with wings 0.3 mm wide; pinnae oblong-lanceolate, 11–18 pairs, (4–)10–15 ⫻ 2–5 cm, sessile, pinnules to ca. 20 pairs per pinna, adnate at their bases and confluent with adjacent pinnules, not or only slightly narrowed, serrately incised; veins 1 per tooth, 3–6 pairs in each pinnule, tips evident adaxially; indument abaxially of scattered, appressed, tan, clavate hairs 0.1–0.2 mm; sori 2–8 pairs per pinna; indusia 1.5– 2.5 ⫻ 0.5–1 mm, margins subentire to erose or sometimes ciliate with hairs 0.1 mm long; spores reniform.

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81

including wings), 1⁄5–1⁄3 of frond length, glabrous or with rare scales, broadly winged 1–2.5 mm nearly to bases; blades thickherbaceous, linear-lanceolate, 1-pinnate, 14–25(–40) ⫻ 5–8(–10) cm, proximal 1 or 2 pinna pairs reduced, each blade terminating in a prolonged rachis with a bud at the tip; rachises brownish, glabrous, broadly winged ca. 1–2 mm from base to bud at the tip of the prolonged rachis; pinnae lanceolate, 10–22 pairs, 3.5– 6 ⫻ 1–1.8 cm, bases equilateral, truncate, exauriculate, apices obtuse or acute, margins biserrate; veins 1- or 2-forked, tips prominent adaxially; indument of scattered, tan, appressed clavate hairs 0.1 mm long; sori 7–14 pairs per pinna, on both sides of costae; indusia 3–8 ⫻ 0.3–0.5 mm, margins entire; spores reniform; 2n⫽72 (Jam). Distribution. Terrestrial or epipetric on moist slopes in wet woods; (250–)500–2300 m. Mexico; Guat, Hond, Salv, Nic, CR (Hammel 17802, US); Peru. Selected Specimens Examined. Chis (Breedlove 69110, CAS). Col (Espero´n Mun˜oz 18299, UC). Gro (Lorea 3020, FCME). Jal (Mexia 1252, UC). Oax (McVaugh 22363, IEB; Rzedowski 19555, NY). Pue (Lyonnet 511200001, MEXU; Riba s.n., MEXU). Ver (Purpus 4350, UC). Unverified, Doubtful, or Mistaken Reports. Mich (reported by Mickel & Beitel, 1988, but not verified).

This highly variable species is part of a complex that involves A. blepharophorum, A. eatonii, and A. solmsii. Material of A. achilleifolium from southern Mexico falls roughly into two morphological groups: 1) specimens with large rhizome scales 4–5 ⫻ 1.5–3 mm and erose indusia 0.8–1 ⫻ 2–2.5 mm; and 2) specimens with small rhizome scales 1.5–3 ⫻ 0.5–0.8 mm and subentire indusia 0.5–0.8 ⫻ 1.5–2 mm. Specimens in this second group also have somewhat more divided pinnules, with the ultimate segments acute. They vary in the direction of A. eatonii in western Mexico. Particularly problematic are specimens from Oaxaca and Chiapas, e.g., Breedlove 24715, 24917 (DS, NY) and 31323 (DS, MEXU, NY). The interrelationships of the species in this group need clarification. Asplenium grande, cited in synonymy, has indusia that are located along the costae where veins to the pinnules diverge, instead of on the pinnules. Asplenium achilleifolium was reported by Mickel and Beitel (1988) from Sinaloa, but specimens from this area appear to be A. eatonii, which see.

3. ASPLENIUM ALATUM Asplenium alatum Humb. & Bonpl. ex Willd., Sp. Pl., ed. 4, 5(1): 319. 1810. Type. Venezuela. “Nova Andalusia, regione temperata, prope Caripe et in declivitate montis Cocollar usque ad alt. 500 hexap.,” Humboldt & Bonpland 455 (B-Willd. 19889!, photo US; isotype P-Humb.!). Figs. 32A, B. Roots wiry, fibrous, not proliferous; rhizomes erect; rhizome scales brown, 1–3 ⫻ 0.8 mm, entire; fronds clumped, 20–60 cm long; stipes gray-brown to green, 3.5–8(–10) cm ⫻ 1–2.5 mm (not

Distribution. Terrestrial or on rocks in montane rain forests on the Atlantic slope; 1150–2700 m. Mexico; Guat, Salv, CR, Pan; Jam, Hisp; Col, Ven, Ec, Peru, s Braz, Bol. Reported from Nicaragua by Go´mez (1976: 52), but not verified. Selected Specimens Examined. Chis (Breedlove 34685, 35400, 38952, DS, MO, 49720, CAS, ENCB). Oax (Mickel 6577, NY, UC, 6714, ENCB, MEXU, NY, UC). Ver (Barrington 442, MEXU; Copeland herb. 62, MEXU, UC, US; Ventura A. 342, ARIZ).

The broadly alate, gray-brown rachises and stipes, proliferous blade apices, and small rhizome scales distinguish A. alatum from congeners in Mexico. The most similar species morphologically is A. pteropus. An even closer relative from Costa Rica to Ecuador is A. rosenstockianum Brade, which differs by having simple, rather than 1-forked veins in the pinnae.

4. ASPLENIUM ARCANUM Asplenium arcanum A. R. Sm., sp. nov. Type. Mexico. Nayarit: Vicinity of San Blas, trail to Ensenada de Matanche´n, Ferris 5360 (holotype DS! isotype GH!). Fig. 41B. Asplenium pumilum Sw. var. laciniatum Davenp., Bot. Gaz. 19: 390. 1894, non Asplenium laciniatum D. Don. Type. Mexico. [Jalisco:] Barranca de Tepic, 11 Oct 1893, Pringle 5534 (GH, not seen; isotypes GH!, K). Ab A. pumilo laminis dissectioribus, pinnis profunde laceratis vel dentatis apice, paginisque marginibus laminarum glabratis differt. (L., arcanus, mysterious, referring to somewhat uncertain affinities and origin.)

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Roots fibrous, not proliferous; rhizomes erect; rhizome scales blackish, clathrate, very narrow, 1–2 ⫻ 0.1–0.2 mm, entire; fronds clustered, mostly 7–20 cm long; stipes atropurpureous at least at bases, greenish distally, 1–12 cm ⫻ 0.3–0.6 mm, 1⁄2–2⁄3 of frond length, essentially lacking hairs or scales except at bases, not winged; blades membranaceous, bipinnate-pinnatifid, 2–10 ⫻ 3– 9 cm, deltate to pentagonal, apices pinnatifid, nonproliferous; rachises abaxially green or purple-streaked at the bases, lacking scales, adaxial wings 0.1–0.2 mm wide; pinnae 1–5 pairs, proximal pair largest, 2–5 ⫻ 1.5–3 cm, inequilateral, exaggerated basiscopically, sessile or stalked to 2 mm, long-acuminate, deeply lacerate, distal pinnae slightly adnate, segments or pinnules lacerate or toothed at tips; veins 1-forked, easily visible, tips evident adaxially; indument abaxially of scattered, appressed, light tan, hairs 0.1–0.2 mm long on veins and tissue between veins; sori 1–3 pairs per segment on both sides of segment midveins; indusia 2–5 ⫻ 0.5–0.8 mm, margins erose; spores reniform.

Asplenium pimpinellifolium Fe´e & Schaffner ex Fe´e, Me´m. Foug. 7: 52, t. 25, f. 5. 1857. Type. Mexico. [Veracruz:] Huatusco, 1854, Schaffner 50 (P!; isotype K!).

Roots fibrous, proliferous; rhizomes erect; rhizome scales orange-brown, clathrate, 3–4 ⫻ 1–2 mm, with golden hairs 0.8–2 mm long; fronds clumped 8–30(–40) cm long; stipes gray-green or brownish green, dull, (2–)8–12 cm ⫻ 0.8–1.2 mm, 1⁄5–1⁄2 of frond length, glabrous except for a few scales at bases, narrowly green-alate; blades thick-herbaceous, 3.5–9 cm wide, 1-pinnate with pinnatifid nonproliferous apices; rachises greenish, glabrous, narrowly alate with wings 0.1–0.3 mm wide; pinnae lanceolate, 8–14 pairs, 18–40 ⫻ 7–15 mm, stalked 1.5–2 mm, inequilateral, basal acroscopic auricles often overlapping the rachises, apices acuminate, tips acuminate to obtuse, coarsely toothed, crenate, or bicrenate, 10–20 teeth on acroscopic margins, incisions 1–2(– 3) mm deep; veins obscure, simple or 1-forked at the acroscopic base, tips evident adaxially; indument seemingly absent, or of sparse, appressed, tan hairs 0.1 mm long; sori 4–6 pairs per pinna, on both sides of midribs; indusia 5–6 ⫻ 0.8–1 mm, margins entire or with scarious fringe; spores reniform; 2n⫽144 (Oax).

Distribution. Terrestrial, on stream banks and wooded slopes; 200–700 m. Mexico. Specimens Examined. Nay (Wright 1360, DS, GH, MO). Sin (Breedlove 35626, CAS, MO, 35642, CAS; Mexia 257, MO, 257a, UC; Pray 1801, NY). Note: Mexia 257 (UC) is A. tenerrimum.

This species has previously been determined as A. cf. fournieri (⫽ A. tenerrimum), an extremely dissected form of A. pumilum (where it was treated as a variety by Davenport), or A. fragrans. It is more or less intermediate between the two first-named species and could have arisen through hybridization. However, specimens of A. arcanum seem to have well formed spores. Breedlove 35626 was previously determined as A. minimum vel aff. by Adams at CAS and as A. tenerrimum by Adams at MO.

5. ASPLENIUM AURICULATUM Asplenium auriculatum Sw., Kongl. Vetensk. Acad. Handl. 38: 68. 1817, non (Thunb.) Kuhn, 1868. Figs. 36H, K, L. Asplenium salicifolium L. var. auriculatum (Sw.) Proctor, Fl. Lesser Antill. 2: 321. 1977. Type. Brazil. Minas Gerais: Rio de Sao Francisco, Freyreis s.n. (S; isotypes BM, BR). Asplenium semicordatum Raddi, Pl. Bras. 1: 36, t. 52, f. 1. 1825. Type. Brazil. Rio de Janeiro, Raddi s.n. (isotypes FI, FI-WEBB).

Distribution. Epiphytic, epipetric or terrestrial in wet forests; 250–2800 m. Mexico; Guat, Hond, Salv, Nic, CR, Pan; Gr Ant, Guadeloupe; Col, Ven, Trin, Guy?, Ec, Peru, Braz, Bol. Adams (in Davidse et al., 1995) also reported Paraguay in the range. Selected Specimens Examined. Chis (Calzada et al. 8725, UC). Gro (Rzedowski & McVaugh 50, 107, NY). Hgo (Arreguin 492, MEXU). Jal (McVaugh 14246, MEXU). Me´x (Aguilar s.n., IZTA). Mich (Langlasse´ 784, US). Oax (Mickel 6458, ENCB, UC). Pue (Ventura A. 20238, ARIZ, ENCB, IEB). Qro (Rzedowski 42857, ENCB, IEB). Tam (Herrera 00288, ITCV). Ver (Purpus 1983, UC).

This species is closely related to A. salicifolium, from which it differs by the smaller fronds and pinnae, 1-forked veins, fewer acroscopic teeth (10–20), and fewer sori (4–6 pairs) per pinna. Both species have long golden hairs on the rhizome apices and sometimes on the stipe base scales, and they differ from other Mexican Aspleniums in this character. Proctor (1977, 1985) included A. auriculatum as a variety of A. salicifolium. Asplenium auriculatum is extremely variable: the pinnae are slender to broad, apices long-acuminate to obtuse, fronds long to short, blades thin to fleshy, small to large. It propagates veg-

ASPLENIUM BARBAENSE

etatively by root buds, so populations are nearly uniform. Gonza´lez-E. et al. 649 (XAL), from Chiapas, and Lorence 4099 (ENCB, NY), from Oaxaca, are large forms of this species, in size approaching A. salicifolium, but the high elevation (2500 m) and lack of pronounced basal acroscopic auricles suggest that they are both A. auriculatum. Asplenium hastatum Klotzsch ex Kunze (Linnaea 23: 235, 305. 1850), based on a cultivated specimen from Venezuela, differs from A. auriculatum in the larger, more equilateral pinnae and broader rachis wings (see Morton & Lellinger, 1966).

6. ASPLENIUM AURITUM Asplenium auritum Sw., J. Bot. (Schrader) 1800(2): 52. 1801. Type. Jamaica. Swartz s.n. (S; isotype B). Figs. 35C, D. Asplenium levyi E. Fourn., Bull. Soc. Bot. France 19: 257. 1872. Type. Nicaragua. Chontales, Levy 474 (P–photo US!; frag. US!).

Roots fibrous, not proliferous; rhizomes suberect; rhizome scales brown, clathrate, 3–4 ⫻ 1–1.3 mm, entire; fronds clumped, mostly 12–45 cm long; stipes green to brownish green, 4–18 cm ⫻ 1–2 mm, 1⁄3–1⁄2 of frond length, glabrous or with rare to sparse scales 0.8–1 mm long, non-alate or very narrowly alate; blades thick-chartaceous to subcoriaceous, narrowly deltate, 11–40 ⫻ 3– 12 cm, 1-pinnate, apices pinnatifid, not proliferous; rachises similar to stipes, glabrous, with wings 0.5 mm wide; pinnae narrowly lanceolate, 12–20 pairs, 2–6 ⫻ 0.5–1.2 cm, stalked 1–4 mm, bases narrowly to broadly cuneate, apices attenuate to obtuse, margins sharply bidentate; indument abaxially of rare to scattered hairlike, clathrate scales 0.5–1 mm long, especially in pinna axils, also with scattered clavate hairs on and between veins; veins obscure, 1-forked, tips evident adaxially; sori 4–8(–12) pairs per pinna; indusia 3–4 ⫻ 0.3–0.5 mm, margins entire; spores reniform, tan, 64 per sporangium; 2n⫽144 (Jam).

83

Selected Specimens Examined. Chis (Stevens & Martı´nez 25901, UC). Oax (Herna´ndez G. 1564, NY). Tab (Cowan 2970, CAS, ENCB, NY; Zamudio R. 1464, IEB). Ver (Wendt et al. 2690, CHAPA, MO; Wendt & Villalobos 3966, CHAPA, UC). Unverified, Doubtful, or Mistaken Reports. Qro (Takaki s.n., ENCB, cited by Dı´az-Barriga & Palacios-Rios, 1992, and Arreguı´n et al., 2001, but specimen not found, and the identification is suspect).

Asplenium macilentum Kunze ex Klotzsch is sometimes regarded as distinct (e.g., by Adams (in Davidse et al., 1995: 309), but we regard its placement as problematic (see comments under A. monodon). Asplenium levyi, treated as a synonym of A. macilentum by Adams, differs from A. auritum only its slightly more coriaceous segments and more nearly entire pinnae. Another synonym is A. plumbeum Christ, also treated as a synonym of A. macilentum by Adams. The A. auritum complex, which in Mexico includes A. auritum, A. cuspidatum, A. monodon, and A. sphaerosporum, needs modern biosystematic study. Morton and Lellinger (1966) treated this complex as two species, A. auritum (with 6 varieties) and A. cuspidatum (with 5 varieties). Until more detailed morphological, cytological, and isozymic studies are made, the division of this complex into taxa based mainly on blade and pinna dissection, blade texture, and spore type will have to stand. The blade dissection series seen in the auritum/cuspidatum complex varies from 1-pinnate in A. auritum, to deeply pinnatepinnatifid to 2-pinnate with dentate segments in A. cuspidatum, to 2-pinnate-pinnatifid to 3-pinnate in A. fragrans. This series is roughly paralleled in the dissection series seen in A. monodon (blades 1-pinnate with superior auricles to pinnate-pinnatifid proximally) to A. sphaerosporum (2-pinnate). Asplenium monodon and A. sphaerosporum differ from the first three species in having globose, blackish spores numbering only 32 per sporangium (indicating an apogamous life cycle) compared with the tan to brown, reniform spores, numbering 64 per sporangium in sexual diploids and tetraploids. All five of these species are basically epiphytic (occasionally epipetric) and occur at somewhat different, but overlapping, elevations in Mexico. Asplenium auritum is found at 100–800 m, A. cuspidatum at (350–)800–2500 m, A. fragrans at (300–)1500–3050 m, A. monodon at 100–1600(–2300) m, and A. sphaerosporum at 650–2700 m.

7. ASPLENIUM BARBAENSE Asplenium barbaense Hieron., Hedwigia 60: 214. 1918. Type. Costa Rica. Volca´n Barba, Hoffmann 26 (B, photo US). Figs. 43F, G. Asplenium williamsii Stolze, Amer. Fern J. 68: 57. 1978. Type. Guatemala. El Progreso: Montan˜a Canahui, Steyermark 43791 (US!; isotype F).

Distribution. Epiphytic or epipetric in wet to dry lowland to lower montane forests; 100–800 m. Mexico; Guat, Bel, Hond, Nic, CR, Pan; Jam; Col, Ven, Trin, Guy, Sur, Ec, Galapagos, Peru, Bol; Africa, Madagascar. Also reported from Brazil by Adams (in Davidse et al., 1995).

Roots wiry, fibrous, not proliferous; rhizomes erect; rhizome scales brown, clathrate, tips long-attenuate, 3–6 ⫻ 0.5–0.8 mm, entire; fronds clumped, 30–60 cm long; stipes gray-green to graybrown, 4–12 cm ⫻ 1–2 mm, 1⁄5–1⁄3 of frond length, glabrous, with green wings 0.5 mm wide, at least distally; blades 25–45 ⫻ 4–10 cm, 1-pinnate with pinnatifid, nonproliferous apices, 2–3

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pinna pairs slightly reduced proximally; rachises gray-green to gray-brown, glabrous, with greenish wings 0.5–0.8 mm wide; pinnae oblong to lanceolate, 14–25 pairs, 2–3 ⫻ 0.8–1.5 cm, bases of basiscopic margins excavate, bases of acroscopic margins usually curved, often overlapping the rachises, apices obtuse to subacute, margins serrate to biserrate with obtuse to subacute teeth, sinuses mostly 1–3(–5) mm deep, deepest sinuses separating acroscopic basal lobe; veins evident, 1-forked on acroscopic sides of pinnae, distal and basiscopic veins simple, tips evident adaxially; indument absent, blades essentially glabrous; sori 2–6(–14) on basiscopic sides, 4–10(–17) on acroscopic sides, (0–)1 or 2 diplazioid sori on basal vein; indusia 3–5(–8) ⫻ 0.5–0.8 mm, margins entire to subentire; spores reniform.

Distribution. Usually epiphytic in wet forests; 1600–2700 m. Mexico; Guat, CR, Pan. Also reported from Honduras, Venezuela, and Ecuador by Adams (in Davidse et al., 1995), but the range uncertain because of confusion with several related species.

Asplenium diplosceum Hieron. (Hedwigia 60: 232. 1918; Cuba, Wright 849, B, isotype BM!) of Cuba, Hispaniola, and Jamaica resembles A. barbaense closely. If they prove to be conspecific, A. diplosceum has priority; Adams (in Davidse et al., 1995: 399) regarded them as distinct, but did not elaborate as to his reasons.

8. ASPLENIUM BLEPHARODES Asplenium blepharodes D. C. Eaton, Zoe 1: 197. 1890. Type. Mexico. Baja California Sur: Sierra de Laguna, 22 Jan 1889, T. S. Brandegee s.n. (YU!). Fig. 39J. Roots thin, fibrous, not proliferous; rhizomes short-creeping to suberect; rhizome scales black, lumina occluded at maturity, 2–3 ⫻ 0.2–0.3 mm, entire; fronds clumped, 12–35 cm long; stipes dark reddish brown, lustrous, mostly 2–6 cm ⫻ 0.6–1.5 mm, 1⁄5– 1⁄4 of frond length, often curved at bases, glabrous, not winged; blades chartaceous, 1-pinnate, buds absent, blades not rooting at tips; rachises dark red-brown, lustrous, abaxially with scattered minute appressed hairs 0.1–0.2 mm long, adaxially with tan wings to 0.2 mm wide; pinnae (10–)15–25 pairs, mostly opposite or subopposite, mostly 1–1.6 ⫻ 0.4–0.7 cm, sessile, proximal ones deflexed, mostly somewhat auriculate at the acroscopic bases, apices oblong, obtuse, finely serrate or biserrate on both sides and tips with rounded teeth, articulate with age; veins faintly visible in young fronds, 1-forked, tips evident adaxially; indument with scattered appressed tan hairs 0.1–0.2 mm abaxially; sori 6–9 per pinna, on both sides of midveins; indusia whitish, 2–3 ⫻ 0.5–0.8 mm, margins fringed with slender, whitish jointed hairs 0.2–1 mm long; spores reniform.

Selected Specimens Examined. Chis (Breedlove 15410, DS, US, 33685, DS, MEXU, NY, 34696, DS). Oax (Lorence 4371, MO; Mickel 4301, 5371, NY, UC, 6581, ENCB, NY, UC, 6583, 6718, ENCB, NY, US, 6753, ENCB, NY, UC, US).

Both specimens of A. williamsii cited by Palacios-Rios (1992) from Veracruz are other species in this complex: Finck 50 (NY) is A. miradorense, and Bourgeau 2369 (NY) is A. potosinum. Several of the collections cited as A. williamsii by Mickel and Beitel (1988) are also A. miradorense: Mickel 1535 (NY, US); Mickel 7306 (NY); Hallberg 1551 (NY). The stated elevation for A. williamsii in both references is consequently too low. Mexican specimens of A. barbaense generally have less attenuate, shorter pinnae than most specimens from Costa Rica and Panama, and it is slightly uncertain whether plants from the two areas are conspecific. Adams (in Davidse et al., 1995), was the first person to unite them. Asplenium barbaense is distinguishable from the closely related A. miradorense by the dull gray-green stipes that are alate to the bases, broadly alate rachises (wings 0.5–0.8 mm wide), longer rhizome scales (4–6 mm), longer sori (3–5 mm), and acroscopic bases of pinnae strongly overlapping the rachises. The two species may grow together (compare, e.g., Mickel 6582, 6719 (NY), both A. miradorense in Oaxaca). See discussion under A. potosinum for comparison with that species.

Distribution. Bases of rocks and crevices of cliffs, also moist pine-oak slopes; 1000–1750 m. Mexico. Specimens Examined. BCS (Alexander et al. 2307, ARIZ, DS, MEXU, UC; Brandegee 660, NY, UC; Breedlove 43272, CAS; Gentry 4282, ARIZ, MO, UC; Gentry & Fox 11762, ARIZ, UC, US; Jones 24162, MO, NY; Thomas 7830, DS, ENCB).

This differs from congeners in Mexico, and especially from its nearest ally, A. fibrillosum, in having rachises scantily pubes-

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85

cent (vs. distinctly fibrillose), forked veins (vs. simple), and indusia delicately ciliate (vs. deeply laciniate).

9. ASPLENIUM BLEPHAROPHORUM Asplenium blepharophorum Bertol., Novi Comment. Acad. Sci. Inst. Bononiensis 4: 443. 1840. Type. Guatemala “Vulcano d’acqua” [Volca´n de Agua], Vellasquez s.n. (BOLO). Typification discussed by R. E. G. Pichi Sermolli & M. P. Bizzarri. 1990–91. Mem. Accad. Lunigianese Sci. “Giovanni Capellini” 40–41: 209–210, t. 43. Figs. 37B, C, E, F. Athyrium macrocarpon Fe´e, Me´m. Foug. 5: 188. 1852. Asplenium macrocarpon (Fe´e) T. Moore, Index Fil. 143. 1859, hom. illeg., non Desv., 1827. Type. Mexico. Oaxaca: Llano Verde, Galeotti 6555 (P; isotypes K!, US!). Asplenium myapteron Fe´e, Me´m. Foug. 8: 82. 1857. Syntypes. Mexico. [Veracruz:] Orizaba, Schaffner 70 (RB); Popocate´petl, Schaffner 294 (RB). Asplenium commutatum Mett. ex Kuhn, Linnaea 36: 99. 1869. Syntypes. Mexico. Schaffner s.n. (B!–photo US!; isosyntype K, photo BM); Oaxaca, Galeotti 6555 (isosyntypes K!, US!); Veracruz: Orizaba, Mu¨ller s.n. (B!; isosyntypes BM, NY!) [“Mu¨ller 1487”]; Guatemala. Salvin & Godman 67, 93; Skinner s.n. (all presumably B). Asplenium onustum Christ in Pittier, Prim. Fl. Costaric. 3(1): 24. 1901. Type. Costa Rica. Copey, Tonduz s.n. [Herb. no. 11931] (P; isotype BR, US!). Asplenium subvestitum Copel., Univ. Calif. Publ. Bot. 19: 289, t. 39. 1941. Type. Mexico. Guerrero: Dto. Mina, W of Petlacala, Mexia 9065 (UC!; isotypes CAS!, F, K!, NY!, US!).

Roots coarse, fibrous, not proliferous; rhizomes erect; rhizome scales brown to tan, clathrate, 5–15 ⫻ 1–2.5 mm, with a few marginal cilia; fronds clumped, (30–)45–100 cm long; stipes brownish green to gray-brown, dull, 8–35 cm ⫻ 2–5 mm, 1⁄3 of frond length, densely clothed with clathrate scales, lacking wings; blades chartaceous, 15–65 ⫻ 6–27 cm, usually 2-pinnatebipinnatifid, ovate-lanceolate, proximal pinnae the largest or 1–2 pairs sometimes slightly reduced, apices pinnatifid or abortive, ending in terminal buds, or buds in axils of distal pinnae, rarely buds absent; rachises yellow to gray-brown, with scattered clathrate scales 0.8–1 mm long, somewhat flattened, but not alate; pinnae 15–25 pairs, 3–14 ⫻ (1–)2–4 cm, sessile, bases truncate, sometimes narrowed on proximal pinnae, costae alate, pinnules deeply pinnatifid or pinnate (at least the larger ones), with 3–4 pairs of lobes, these obtuse to acute, sometimes bifid at tips; veins evident on both sides, tips not significantly enlarged or more visible; indument abaxially of numerous clathrate scales on costae and costules, also with tan to whitish scattered, appressed, clavate hairs 0.2–0.3 mm on veins and tissue between veins; sori 2–4 pairs per pinnule; indusia whitish, thick, 2–3 ⫻ 0.8–1 mm, margins entire or suberose; spores globose.

Distribution. Terrestrial or epipetric, moist ravines, montane rain forests, cloud forests, fir forests; 1150–3100 m. Mexico; Guat, Hond, CR, Pan; Hisp. Cited from El Salvador by Adams (in Davidse et al., 1995: 298). Also reported from Nicaragua by Go´mez (1976: 52). Selected Specimens Examined. Chis (Little & Sharp 9880, UC, US). Col (Sanders 10633, UC). DF (Lyonnet 3250, MEXU, US). Gro (Breedlove 65180, CAS). Hgo (Sa´nchez Mejorada 128, US). Jal (McVaugh 10041, DS, MEXU, NY, UC, US). Me´x (Lyonnet 2090, CHAPA, MO, US). Mich (Rzedowski 41965, ENCB, IEB). Mor (Lyonnet 1862, MEXU). Oax (Pringle 6072, GH, LL, MEXU, MO, NY, UC, US). Pue (Pringle 10300, MEXU, MO, NY, UC, US). Qro (Rzedowski 48346, IEB). SLP (Virlet d’Aoust 52, P–photos MO, US). Ver (Ventura 293, DS, NY). Unverified, Doubtful, or Mistaken Reports. Nay (reported by Mickel, 1992, but not verified; possibly the specimen seen was A. eatonii).

The A. blepharophorum complex, which also includes A. eatonii and A. achilleifolium, is variable with regard to the presence or absence of subterminal buds. Specimens with buds (Mickel 3810b, 4129, 4692, 4916, NY, UC), here referred to A. blepharophorum, generally have blades broadest at the bases, or the proximal pair of pinnae only slightly reduced. Of the specimens without buds, some have the proximal pinnae greatly reduced; these correspond to what is here called A. eatonii, in western Mexico. Other budless specimens have blades more or less identical to A. blepharophorum, and are here referred to that species. Asplenium achilleifolium differs in having much less scaly stipes, rachises, costae, and costules. More work is necessary to determine the relationship and status of these variations. There is an indication (large globose spores) that apogamy may be operative in A. blepharophorum.

10. ASPLENIUM BREEDLOVEI Asplenium breedlovei A. R. Sm., Proc. Calif. Acad. Sci., ser. 4, 40: 209, f. 1, a–b. 1975. Type. Mexico. Chiapas: Mpio. Villa Corzo, E base of Cerro Tres Picos, Breedlove 30041 (DS!; isotypes CAS!, F, MO!, NY!). Fig. 40B. Roots fibrous, not proliferous; rhizomes short-creeping; rhizome scales dark brown to blackish, clathrate, 1–3 ⫻ 0.2–0.3 mm;

86

ASPLENIUM

fronds few, 1–3 per plant, to ca. 50 cm long; stipes atropurpureous or brown, dull, mostly 10–25 cm ⫻ ca. 2 mm, 1⁄3–1⁄2 the frond length, glabrous, without adaxial wings; blades chartaceous, deltate, to 35 ⫻ 15 cm, nearly 4-pinnate proximally, gradually tapering toward apices, buds absent, blades not rooting at tips; rachises darkened proximally, greenish distally, glabrous, not winged; pinnae to 8–12 pairs, ascending, to 10 ⫻ 3 cm, not articulate, stalked to 3 mm, pinnules pinnate-pinnatifid, cuneiform, mostly 2–5-lobed, ultimate segments to 1.5 mm wide, acute or subacute at tips; veins 2–3-forked in the ultimate segments, visible on both sides, terminating in linear hydathodes near the margins; indument abaxially of scattered, tan appressed clavate hairs ca. 0.1 mm long; sori 1–2 per ultimate segment; indusia whitish to tan, 1.5–3 ⫻ 0.3–0.6 mm, margins entire or slightly erose; spores globose (Adams in Davidse et al., 1995).

Distribution. Terrestrial in montane rain forests; 1500–1800 m. Mexico; Guat., Hond (Hawkins 979, MO). In Mexico known only from the type collection. The most similar Mexican species are A. solmsii and A. insolitum, from which A. breedlovei differs in having creeping rhizomes. The nearest affinities of A. breedlovei are obscure; it seems to be a rather isolated species, and is unusual in the genus because of the creeping rhizomes. It does not appear closely related to sect. Hymenasplenium, species of which regularly have creeping rhizomes.

11. ASPLENIUM CASTANEUM Asplenium castaneum Schltdl. & Cham., Linnaea 5: 611. 1830. Asplenium monanthes L. var. castaneum (Schltdl. & Cham.) Stolze, Fl. Ecuador 23: 45. 1986. Type. Mexico. Veracruz: Mt. Orizaba, Schiede s.n. [768] (HAL; isotype B, frag. NY!). Fig. 33K. Asplenium rubinum Davenp., Bot. Gaz. 19: 391. 1894. Type. Mexico. Me´xico: Sierra de las Cruces, Pringle 5191 (GH!; isotypes B!, MEXU!, US!, YU!). The holotype was indicated incorrectly as at B by Mickel & Beitel (1988).

Roots fibrous, not proliferous; rhizomes suberect; rhizome scales black, clathrate, not occluded, 4–6 ⫻ 0.3–0.6 mm, entire, tips long-attenuate; fronds clumped, numerous, mostly 12–35 cm

long; stipes castaneous, lustrous, mostly 5–11(–17) cm ⫻ 0.7–2 mm, 1⁄4–1⁄2 of frond length, glabrous or with scattered hair-like scales (1–2 mm long) proximally, not alate; blades thickherbaceous to subcoriaceous, 1-pinnate, linear, 15–30 ⫻ 1–2 cm, apices pinnatifid, not proliferous; rachises reddish brown, with scattered filiform scales to ca. 1 mm and a few spreading septate hairs 0.2–0.4 mm, adaxial wings entire or rarely minutely papillate, 0.3 mm wide; pinnae nearly round or rhomboid, (10–)20– 30 pairs, 6–10 ⫻ 5–6 mm, sessile, articulate, rachis color running out onto pinna stalks, the largest ones slightly auriculate acroscopically, margins entire to faintly crenulate; veins simple or 1forked, tips evident adaxially; indument absent, blades essentially glabrous, or with sparse to scattered clavate hairs; sori 2–6 pairs per pinna, 2–4 on basiscopic sides, 2–5(–6) on acroscopic sides; indusia whitish to light tan, 1–3 ⫻ 0.8–1.2 mm, margins erose; spores globose to ovoid.

Distribution. Terrestrial or epipetric, often in rock crevices, usually above timberline, or in fir forests; (2450–)3300–4350 m. Mexico; Guat, CR, Pan; Col, Ven, Ec, Peru, Bol. Also reported from El Salvador and Chile by Adams (in Davidse et al., 1995), but Marticorena and Rodrı´guez (1995) did not include it in the range for Chile. Selected Specimens Examined. Chis (Matuda 2353, K, MEXU, US). DF (Balls 4099, UC, US). Gro (Lorea 3980, 4205, 4360, FCME). Gto (Rzedowski 47913, IEB; 47913a, UC). Jal (Beaman 2372, UC, US). Me´x (Beaman 2840, UC). Mich (Dı´az B. 1009, ENCB, IEB, MEXU). Mor (Sa´nchez 310, MEXU). Oax (Mickel 4582, NY). Pue (Diggs 2060, ENCB, NY). Tlax (Acosta P. & Sa´nchez 2968, CIB). Ver (Beaman 2288, UC, US). Unverified, Doubtful, or Mistaken Reports. Dgo (reported by Mickel & Beitel, 1988, but not verified; most likely, specimens so determined are A. sanchezii). NL (Hinton 17189, TEX, cited by Hinton & Hinton, 1995).

Asplenium castaneum is distinguished from A. monanthes in having castaneous rachises and clathrate, non-occluded rhizome scales. Another related species is A. polyphyllum, but that has a subterminal proliferous bud along the rachis of each blade, the pinnae are more oblong, and the indusia subentire. Stolze (1986) and Tryon and Stolze (1993) treated A. castaneum as a variety of A. monanthes, but the two seem readily distinct and, in Mexico at least, seldom if ever grow together.

ASPLENIUM CIRRHATUM

Occasional specimens of A. castaneum are very small, with as few as 8 pinna pairs (e.g., Ortega O. 538, XAL, from Veracruz). These specimens approach A. fibrillosum in size, but the indusia are subentire or erose and the rachises less scaly. Small specimens of A. castaneum also resemble A. sanchezii, which see for differences.

12. ASPLENIUM CHIHUAHUENSE Asplenium chihuahuense Baker, Ann. Bot. (Oxford) 5: 305. 1891. Type. Mexico. Chihuahua: Arroyo Ancho, Pringle 1444 (K; isotypes CAS!, MEXU!, NY!, US!). Fig. 41A. Asplenium dubiosum Davenp., Garden & Forest 4: 483. 1891, nom. provis. Davenport cited Pringle 1444, the type of A. chihuahuense, as representative.

Roots fibrous, not proliferous; rhizomes short-creeping or suberect; rhizome scales dark brown to blackish, clathrate, 2–3(–4) ⫻ 0.2–0.4 mm, entire; fronds clumped, 4–8(–15) cm long; stipes castaneous to atropurpureous, lustrous, 1–5 cm ⫻ 0.5–1 mm, 1⁄4– 2⁄3 of frond length, glabrous except for scattered filiform scales and minute hairs, non-alate; blades firm-herbaceous to subcoriaceous, ovate-deltate, acute and tapering at apices, rather irregularly pinnate-pinnatifid to 2-pinnate, 2.5–5(–10) ⫻ 2–3(–6) cm; rachises reddish brown proximally, green at least in the distal half, dull, glabrescent or with filiform scales to 1 mm long, not winged adaxially; pinnae 4–10 pairs, proximal pinnae deltate to broadly ovate-deltate, 1–4 ⫻ 1–2.5 cm, more distal pinnae deltatelanceolate, blades tapering to pinnatifid apices, ultimate segments ovate to spatulate, cuneate proximally, sharply toothed or cleft with ascending teeth, margins thickened and pale; veins not evident except at tips adaxially; indument absent, blades essentially glabrous; sori 1–6 pairs per segment, on both basiscopic and acroscopic sides of midribs of pinnae and pinnules; indusia whitish, 1–3 ⫻ 0.2–0.3 mm, margins entire to erose; spores reniform; 2n⫽216 (Chih).

Distribution. In deep pockets and crevices of rhyolitic rocks and in limestone canyons; 1800–2600 m. USA (Ariz, Colo, Utah); Mexico. Specimens Examined. Chih (Correll & Gentry 23042, UC, US; Knobloch 597, 977, US; McVaugh 11508, US; Pringle 1711, NY, UC; Reeves 4877, 4946, ASU; Toolin 1351, ARIZ, US). Dgo (Reeves et al. 5006, ASU).

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From artificial hybridization studies, Rasbach et al. (1994) demonstrated that this species is an allohexaploid and possibly formed from hybridization between A. adiantum-nigrum L. var. adiantum-nigrum, a tetraploid, and some unknown diploid. Asplenium adiantum-nigrum L. is principally a Eurasian species but is also found in North America (Ariz, Colo, Utah), where it is very rare, and in Hawaii. It is an allotetraploid derived from hybridization of two European diploids, A. cuneifolium Viv. and A. onopteris L. New World specimens of A. adiantumnigrum (and A. chihuahuense) are generally much smaller (fronds one-third to one-half the length of Old World specimens) than those of A. adiantum-nigrum from elsewhere in the range, but otherwise not very different. Windham (1983) summarized evidence, both cytological and chemical, for treating Eurasian and American plants of A. adiantum-nigrum as conspecific.

13. ASPLENIUM CIRRHATUM Asplenium cirrhatum Rich. ex Willd., Sp. Pl., ed. 4, 5(1): 321. 1810. Figs. 38B, C. Asplenium radicans L. var. cirrhatum (Rich.) Rosenst., Hedwigia 46: 102. 1906. Type. Guadeloupe. Richard s.n. (B-Willd. 19894!, microfiche UC!; frag. and tracing NY!).

Roots wiry, fibrous, not proliferous; rhizomes suberect or erect; rhizome scales dark brown, clathrate, ca. 4–6 ⫻ 0.4–0.6 mm, entire; fronds clumped, 20–35 cm long (including flagelliform tips); stipes dark purplish to castaneous, lustrous, 4–10 cm ⫻ 0.8– 1 mm, 1⁄4–1⁄3 of frond length, glabrous or with a few hair-like scales, not alate; blades thick-herbaceous, lanceolate, 1-pinnate, 10–25 ⫻ 3.5–6(–8.5) cm, broadest at bases or nearly so, apices with gradually smaller pinnae, ending in flagelliform tips 3–8 cm long terminated by a small bud; rachises atropurpureous to castaneous (at least proximally), lustrous, with sparse hair-like scales, lacking wings or adaxially with narrow green wings to ca. 0.2 mm; pinnae oblong, 10–15 pairs, 1.5–3(–4) ⫻ 0.6–1.2 cm, sessile, not articulate, excavate basiscopically, truncate and nonauriculate to slightly auriculate acroscopically, margins subentire to crenate, apices obtuse to acutish, sinuses to ca. 1 mm deep; veins obscure on both sides, 0–1-forked, tips prominent adaxially; indument abaxially of scattered, tan, appressed, clavate hairs 0.1–0.2 mm long; sori 3–7(–9) pairs per pinna, on both sides of midveins; indusia 2–4(–6) ⫻ 0.4–0.7 mm, margins entire to suberose; spores reniform.

88

ASPLENIUM

pinna, ultimate segments elliptic or oblong, 4–6 mm long, cuneate proximally, acute distally, slightly falcate; veins single in the ultimate segments, readily visible on both sides, tips evident adaxially; indument abaxially of whitish to tan, appressed, clavate hairs 0.1–0.3 mm long; sori one per ultimate lobe; indusia 1–3 ⫻ 0.3–0.5 mm, margins entire; spores reniform; 2n⫽72 (Fla), 144 (Trin, Galapagos).

Distribution. Terrestrial, in wet montane forest; 450–2200 m. Mexico; Bel, Guat, Nic, CR, Pan; Jam, PR, L Ant; Col, Ven, Ec, Peru, s Braz, Bol. Also reported from Honduras, and French Guiana by Adams (in Davidse et al., 1995). References to this species from Trinidad (Adams in Davidse et al., 1995) may pertain to A. acutiserratum (Hieron.) Mickel. Specimens Examined. Oax (Mickel 927, UC, 5570, 5673, 6610, ENCB, MEXU, NY, UC, 6459, NY, UC, 7085, ENCB, NY, UC; Palacios-Rios 2331, UC; Vera Santos 3422, US).

This species, with 1-pinnate blades and non-auriculate to slightly auriculate pinnae, is part of a related group of species with castaneous to atropurpureous, lustrous stipes and long, flagelliform, proliferous blade apices. This complex includes taxa that are progressively more divided, ending with the fully 3-pinnate A. uniseriale. For discussion of this complex see under A. radicans. In Mexico, Asplenium cirrhatum is most closely related to the 1-pinnate A. yelagagense (which see). Mickel and Beitel (1988) reported this species from Chiapas, but we are now unable to verify this record.

14. ASPLENIUM CRISTATUM Asplenium cristatum Lam., Encycl. 2: 310. 1786. Type. Puerto Rico. LeDru s.n. (P-JU 1276!, photo US). Fig. 38G. Asplenium cicutarium Sw., Prodr. 130. 1788. Type. Jamaica. Swartz s.n. (S; isotype B-Willd., photo NY!).

Roots relatively coarse, fibrous, not proliferous; rhizomes erect; rhizome scales black, clathrate, 3–5 ⫻ 0.3–0.7 mm, entire; fronds clumped, mostly 30–55(–70) cm long; stipes 6–10(–15) cm long, ca. (1⁄8–)1⁄4–1⁄2 of frond length, dull gray-brown, glabrous, with narrow wings 0.2–0.8 mm wide at least distally; blades membranaceous, 2-pinnate-pinnatifid to 3-pinnate, (15–)30–50 ⫻ 7– 15 cm, lanceolate to deltate, apices long-acuminate, pinnatifid, nonproliferous; rachises brownish, dull, glabrous, with narrow wings 1 mm wide; pinnae 15–30 pairs, spreading, mostly 2.5–8 ⫻ 1–2.5 cm, sessile, with proximal pinnules overlapping the rachises, usually with 1 or 2 pairs (infrequently to 9 pairs) greatly reduced and slightly deflexed proximal pinnae, proximal pinnules on larger deeply pinnatifid to 1-pinnate, 9–14 pairs per

Distribution. Terrestrial or epipetric in dense woods; 200– 1600(–2100) m. USA (Fla); Mexico; Guat, Bel, Hond, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Ec, Peru, Braz, Bol. Cited also from the Guianas, Galapagos, and Paraguay by Adams (in Davidse et al., 1995). Selected Specimens Examined. Chis (Croat 40661, MO, UC). Oax (Herna´ndez G. 1377, 1943, CHAPA, MO, NY). Pue (Sa´nchez Mejorada R. 631, MEXU, US). Qro (Rzedowski 46351, IEB). SLP (Pringle 3365, CAS, DS, MEXU, MO, NY, US, atypical). Tab (Cowan 2965, CAS, MEXU). Tam (Palmer 299, MO, NY, UC, US; Webster 213, TEX). Ver (Purpus 1971, MO, UC, US). Unverified, Doubtful, or Mistaken Reports. Me´x (reported by Mickel & Beitel, 1988, but not verified). Mich (reported by Leavenworth, 1946, not verified, likely misidentified).

Asplenium cristatum differs from A. rutaceum by having nonproliferous blade apices and dull, non-lustrous stipes and rachises. The opposite pinnae, longer stipes, proximal pinnules that strongly overlap the rachises, more broadly cuneate segments that often have 2- or 3-lobed apices, narrower indusia 0.3–0.5 mm wide, and occurrence at lower elevations distinguish A. cristatum from A. myriophyllum (which see), but the taxa may hybridize, producing hybrids with intermediate morphology and malformed spores, as reported by Smith (1981) from Chiapas. A specimen from Oaxaca (Mickel 7301b, NY) growing with typical A. cristatum resembles this putative hybrid and also shows irregular spore size. Pringle 4072 (MEXU), from San Luis Potosı´, Tamasopo Canyon, appears to be a hybrid involving A. cristatum and perhaps A. semipinnatum, known from the same locality. Spores of this specimen appear to be strongly malformed and most of the sporangia are collapsed. The same number at UC is more typical A. cristatum. Breedlove 22294 (DS, NY), from Chiapas, also seems to be a hybrid, possibly involving A. rutaceum or possibly A. myriophyllum; the sporangia appear abortive. In Florida, the

ASPLENIUM CUSPIDATUM

abortive-spored hybrid, A. plenum E. P. St. John ex Small, may have A. cristatum and A. abscissum as parents. Asplenium ocoe¨nse C. Chr. (Kongl. Svenska Vetenskapsakad. Handl., ser. 3, 16(2): 53, t. 13, figs. 6–8. 1937; type from Dominican Republic, Ekman H11785, isotype US!), from Hispaniola and Puerto Rico, seems indistinguishable from specimens of A. cristatum with reduced proximal pinnae. Proctor (1989) commented that similar specimens occurred in Mesoamerica and South America, and we agree. Several specimens determined and cited by Mickel and Beitel (1988) as this species are A. uniseriale (e.g., Herna´ndez G. 1182, 1472, 1672, 1905, CHAPA, NY), which differs in having darker, lustrous stipes and rachises and flagelliform apices.

15. ASPLENIUM CRUEGERI Asplenium cruegeri Hieron., Hedwigia 60: 254. 1918. Type. Trinidad. Am Camerun-Wasserfall, Crueger 44 (B p.p.). Fig. 43E. Roots thin, fibrous, not proliferous; rhizomes suberect; rhizome scales brown, clathrate, 3–4 ⫻ 0.2–0.4 mm, entire; fronds tightly clustered, 10–15(–25) cm long; stipes brownish to brown-purple, dull, 2–5(–10) cm long, 1⁄4–1⁄3 of frond length, glabrous, adaxially with green wings 0.1–0.2 mm wide; blades thin-chartaceous, 1pinnate, oblong-lanceolate, 8–20 ⫻ (2–)2.5–3.5 cm, not or only slightly reduced proximally, apices pinnatifid, not proliferous; rachises tan to greenish 2⁄3 of length, glabrous, adaxially with very narrow green wings 0.1–0.2 mm wide; pinnae 15–20(–25) pairs, 1.4–1.8 ⫻ 0.5–0.8 cm, sessile to short-stalked to 1 mm, pinna bases inequilateral, dimidiate, slightly auriculate acroscopically, apices obtuse, margins subentire basiscopically, crenulate to bicrenulate acroscopically and at tips; veins 1(–2)-forked acroscopically, simple basiscopically, tips evident adaxially; indument abaxially of scattered, appressed, whitish to light tan, clavate hairs 0.1–0.2 mm long on tissue between veins; sori 6–10 per pinna, 2–4 on basiscopic sides, 4–6 on acroscopic sides; indusia 2–4 ⫻ 0.5–0.8 mm, margins subentire; spores reniform.

89

Specimens Examined. Chis (Breedlove 39059, DS, ENCB, NY).

This is closely related to A. claussenii Hieron., widespread in South America from Colombia and Venezuela to Brazil and Paraguay, and A. hostmannii Hieron. (Hedwigia 60: 256. 1918), from northern South America (Venezuela, Guianas, and Brazil). It belongs to the A. miradorense group and is distinguished from other Mexican species by the smaller number of pinna pairs, generally 15–20 per blade, and pinnae rounded at tips, more or less entire along the basiscopic margins, shallowly crenate or toothed along acroscopic margins. The veins are 1-forked in the proximal half of the pinnae and simple toward the tips. From A. laetum, it differs by the erect rhizomes, blunt-tipped and shorter pinnae, and greater number of pinna pairs per blade. Adams (in Davidse et al., 1995) referred the Chiapas specimen cited here to A. hostmannii, which differs by the more coarsely dentate, or often bidentate pinna margins and more tapering pinnae (isolectotype Hostmann 168, K, photo UC!). He also cited specimens of A. hostmannii from Guatemala and Panama, as well as from northern South America, but it is not clear whether he regarded A. cruegeri as a synonym. Both species were published in the same paper by Hieronymus.

16. ASPLENIUM CUSPIDATUM Asplenium cuspidatum Lam., Encycl. 2: 310. 1786. Type. Peru. J. de Jussieu s.n. (P-JU 1253!, photos BM, NY!, US!; isotype PLA, photo US). Figs. 36D–F. Asplenium lacerum Schltdl. & Cham., Linnaea 5: 612. 1830. Type. Mexico. [Veracruz:] prope Jalapa, Schiede & Deppe s.n. [775] (B!, frag. NY!, photo US!; isotypes HBG–photos NY!, US!; LE!–2 sheets, MO!, P. Asplenium mexicanum M. Martens & Galeotti, Me´m. Foug. Mexique 62, pl. 15, f. 4. 1842. Asplenium auritum Sw. var. mexicanum (M. Martens & Galeotti) Hieron., Bot. Jahrb. Syst. 34: 468. 1904. Asplenium fragrans Sw. var. mexicanum (M. Martens & Galeotti) Christ, Bull. Herb. Boissier, se´r. 2, 5: 729. 1905. Type. Mexico. Veracruz: Xalapa and Mirador, Galeotti 6391 (BR!– photos UC!, US!; isotypes BR! P–frag. NY!). Asplenium auritum Sw. var. bipinnatifidum Kunze, Linnaea 18: 332. 1844 [1845]. Type. Mexico. Leibold 14 (LZ destroyed). Asplenium pyramidatum Liebm., Mexic. Bregn. 249 (reprint 97). 1849, non Desv., 1827. Lectotype (chosen by Smith, 1981: 51). Mexico. Oaxaca: [Ixtla´n] Between Tanetze and Cacalotepec, Liebmann s.n. [Fl. Mex. 333] (C!; isolectotype US!). Asplenium auritum Sw. var. bipinnatisectum Mett., Abh. Senckenberg. Naturf. Ges. 3: 147. 1859. Lectotype (chosen by Morton & Lellinger, 1966: 20). Mexico. Schiede & Deppe s.n. [773] (B). See comments under Asplenium sphaerosporum.

Similar to A. auritum except: blades deeply pinnate-pinnatifid to 2-pinnate with dentate segments, proximal pinnae with at least an acroscopic basal lobe cut nearly to costa.

Distribution. Terrestrial in oak forest; 1000 m. Mexico; Hond, CR; Ven, Trin.

90

ASPLENIUM ⫻

Distribution. Epiphytic or epipetric, rarely terrestrial; (350–) 800–2500 m. Mexico; Guat, Nic, Salv, CR, Pan; Cuba, L Ant; Col, Ven, Ec, Peru, Bol. Also reported from Honduras, Brazil, and Paraguay by Adams (in Davidse et al., 1995), but the species is somewhat differently circumscribed by him.

1 mm, 1⁄10–1⁄15 length of fronds, or less, scaly throughout, not winged; blades subcoriaceous, elliptic, pinnatifid, lobed to within 1–2 mm of blade midribs, ca. 4–15(–20) ⫻ 1.5–6 cm, gradually tapered proximally, apices obtuse, not rooting; rachises light brown to tan, dull, not winged, with numerous, brownish, strongly clathrate scales 0.5–2 mm, these linear, uniseriate at tips, to ca. 6 cells wide and lanceolate; pinnae confluent with adjacent pinnae, deltate, 8–20 pairs, to 3 ⫻ 1.5 cm, widest at the bases, rounded at tips, margins entire to crenulate; veins obscure, 2–3forked, sometimes casually anastomosing near margins, tips not evident adaxially; indument abaxially of whitish to tan, appressed clavate hairs 0.1–0.2 mm long; sori 3–8 pairs per pinna, on both basiscopic and acroscopic sides of pinnae; indusia whitish, mostly 3–10 ⫻ 0.5–0.6 mm, margins entire; spores reniform, 64 per sporangium; 2n⫽72 (India).

Selected Specimens Examined. Chis (Stevens & Martı´nez 25860, MO, UC). Hgo (Moore 5305, UC, US). Jal (McVaugh 16188, MEXU, NY, TEX). Mich (Cota 17096, UC). Mor (Lyonnet 775, NY). Oax (Mickel 6364, UC). Pue (Sa´nchez M. 598, US). Qro (Rzedowski 42721, IEB, XAL). SLP (Pringle 3364, CAS, DS, NY, UC, US). Tam (Stanford et al. 983, ARIZ, DS, MO, NY, US). Ver (Purpus 5945, UC). Unverified, Doubtful, or Mistaken Reports. Gro (reported by Mickel & Beitel, 1988, and Lorea-Herna´ndez & Vela´zquez M., 1998, but these specimens are most likely A. fragrans).

See discussion under A. auritum. Adams (in Davidse et al., 1995) recognized A. rigidum Sw. (Kongl. Vetensk. Acad. Handl. 1817: 68. 1817), type from southern Brazil (Freyreis s.n., S, photo UC!), as distinct from A. cuspidatum and ranging into southern Mexico and Central America. We are doubtful that the two species, both of which are 2-pinnate or sometimes slightly more divided, are specifically different, and even if they are distinct, we have seen nothing in Mexico that quite matches the Brazilian type. We therefore choose not to recognize this species from Mexico. Asplenium cuspidatum has been variously circumscribed, in its tortuous history, often more broadly, to include what we have here called A. fragrans (which see for further discussion).

17. ASPLENIUM DALHOUSIAE Asplenium dalhousiae Hook., Icon. Pl. 2: t. 105. 1837. Figs. 41J, K. Ceterach dalhousiae (Hook.) C. Chr., Index Filic. 170. 1905. Ceterachopsis dalhousiae (Hook.) Ching, Bull. Fan Mem. Inst. Biol. 10: 9. 1940. Type. India. Simla, Dalhousie s.n. (K). Asplenium rupium Goodd., Muhlenbergia 8: 92. 1912. Syntypes: U.S.A. Arizona: Cochise Co., Mule Mts., Asplenium Canyon, Goodding 67, 969 (RM; isosyntype of Goodding 67 at UC!).

Roots thin, wiry, fibrous, not proliferous; rhizomes erect; rhizome scales black with brown margins, clathrate, 2–5 ⫻ 0.6–1 mm, sparingly denticulate; fronds mostly 10–20 cm long; stipes dark to light brown throughout, dull, nearly absent to 1(–3) cm

Distribution. Terrestrial or at bases of rocks in moist pine-oakarbutus slopes, rocky ravines; 1050–2000 m. USA (Ariz); Mexico; Asia in the Himalayas. Specimens Examined. Chih (Bye 1772, COLO not seen, UCR). Sin (Breedlove 16464, CAS, NY, 16904, CAS, 18370, CAS, MO; Gentry 6503, ARIZ, MICH, MO, US). Son (Fishbein 1042, ARIZ; Phillips 547, ARIZ; Steinmann 1317, NY; Toolin 306, ARIZ).

This species is highly distinctive and not closely related to any other in Mexico. It is the only Mexican species of Asplenium with deeply pinnatifid blades. Its affinities appear to be with Old World species. It is a member of the segregate genus Ceterach, and differs from congeners in Mexico by having shallowly pinnatifid blades. Most of the collections seen are from the Sierra Surutato, Edo. Sinaloa.

18. ASPLENIUM DELICATULUM Asplenium delicatulum C. Presl, Reliq. Haenk. 1(1): 47, t. 7, f. 5. 1825. Type. Ecuador. “in Cordilleris Quitensibus,” Haenke s.n. (PRC; isotype HBG, photos F, GH, US). Figs. 40G, H. Roots filiform, proliferous and giving rise to new plantlets at long intervals (thus forming colonies); rhizomes suberect; rhizome scales brown, clathrate, obtuse at tips, concave on inner (adaxial) surfaces, 0.5–1 ⫻ 0.1–0.3 mm, entire; fronds tightly clustered, 5–15 cm long; stipes brownish green, dull, 1–4 cm ⫻

ASPLENIUM DELITESCENS

0.3–0.5 mm, 1⁄4–1⁄3 of frond length, glabrous, with wings 0.1 mm wide; blades membranaceous, 3–6 ⫻ 1.5–2.5 cm, 2–3-pinnate, narrowly deltate, widest at bases, apices pinnatifid, not proliferous; rachises greenish, glabrous, with narrow greenish wings 0.1 mm wide; pinnae 4–6 pairs, 1–1.5 cm long, stalked to 2 mm, ultimate segments linear or narrowly elliptic, 0.5–0.8 mm wide, often notched or bifurcate at tips; veins evident, simple, tips evident adaxially; indument abaxially of scattered, appressed, whitish to tan, clavate hairs 0.1 mm long; sori 1 per ultimate segment and usually located at bases of segments near bifurcation of veins; indusia 1–1.5 ⫻ 0.5–0.6 mm, margins erose; spores reniform.

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Diplazium delitescens Maxon, Contr. U.S. Natl. Herb. 10: 497, t. 56, f. 1. 1908. Type. Cuba. Oriente: Vicinity of San Luı´s, Pollard & Palmer 348 (US!; isotypes F, MO, NY!).

Roots coarse, fibrous, not proliferous; rhizomes short- to longcreeping; rhizome scales clathrate, uniseriate at tips, 0.5–2 ⫻ 0.1– 0.3 mm, entire; fronds 1–3 mm apart, 32–56 cm long; stipes stramineous to brown, dull, 16–26 cm ⫻ 2–3 mm, 2⁄5–1⁄2 of frond length, bases swollen to ca. 3–4 mm, glabrous or with occasional scales 0.5–1.5 ⫻ 0.1–0.2 mm long, adaxially with green wings 0.1–0.3 mm wide; blades chartaceous, broadly deltate, 1-pinnate, 20–32 ⫻ 13–26 cm, apices pinnatifid, not proliferous; rachises green, with wings ca. 0.1–0.2 mm wide, glabrous or with occasional filiform scales; pinnae lanceolate to linear-lanceolate, 7–9 pairs per blade, (3.5–)7–14 ⫻ 0.8–2 cm, stalked to ca. 1.5 mm, subequilateral to inequilateral, excavate basiscopically, acroscopic bases parallel to rachises and slightly auriculate, apices acuminate, margins serrate; veins visible on both sides of blades, 2–3forked, vein tips visible adaxially; indument abaxially of redbrown appressed clavate hairs 0.1 mm long on and between veins; sori 9–22 pairs per pinna, occasionally diplazioid especially on terminal segments; indusia mostly 5–15 ⫻ 0.3–0.4 mm, margins suberose; spores reniform, 64 per sporangium (Murakami & Moran, 1993).

Distribution. On rocks along streams; 300–950 m. Mexico; Guat, Hond, Nic, CR, Pan; Ec, Peru. Also reported from Cuba, Hispaniola, Colombia, Venezuela, Chile, and Argentina by Adams (in Davidse et al., 1995), but the species is circumscribed differently by him; Asplenium delicatulum is not included for Chile by Marticorena and Rodrı´guez (1995). Specimens Examined. Chis (Breedlove 48315, 50838, CAS; Dressler 1627, US). Oax (Herna´ndez G. 1483, 1669, CHAPA, MEXU, MO, NY).

Asplenium delicatulum is most similar to very small forms of A. cuspidatum and differs by its much smaller size, thinner blade texture, proliferous roots that produce new plantlets, and very small, ovate, more or less appressed and inrolled rhizome scales. Another close relative is A. tenerrimum, which differs in the more broadly deltate blades, darker, more polished stipe bases, and linear-lanceolate rhizome scales. Throughout its range, A. delicatulum occurs only at relatively low elevations, below 1000 m. Asplenium cuspidatum Lam. var. triculum Lellinger (Proc. Biol. Soc. Wash. 98: 367. 1985. Type. Costa Rica. San Jose´: Basin of El General, Skutch 4776, US; isotypes F, G) is also very similar to this species and differs primarily by having fronds not borne at intervals along thread-like roots, larger and less dissected blades, and patent pinnae.

19. ASPLENIUM DELITESCENS Asplenium delitescens (Maxon) L. D. Go´mez, Brenesia 8: 52. 1976. Figs. 43A, B.

Distribution. Terrestrial or epipetric in limestone regions of Atlantic lowlands; 50–800 m. Mexico; Guat, Bel, Hond, Nic, CR, Pan; Cuba; Col, Ven, Ec, Peru, n Braz. Also reported from French Guiana (Boggan et al., 1997). Selected Specimens Examined. Chis (Breedlove 57997, CAS; Chater 128, MEXU, MO; Martı´nez S. 7848, ARIZ, MO, 8136, ARIZ, MO, NY). Oax (Herna´ndez G. 1920, CAS, CHAPA, MEXU, MO, NY; Mickel 6843, ENCB, NY, UC, US). Tab (Croat 47908, CR, MO). Ver (Finck 159, UC; Leeds 166, US; Reeder & Reeder 1974, US).

Smith (1977) discussed the placement of this and other species in Asplenium rather than in Diplazium, where traditionally they have been treated. Murakami and Moran (1993) provided a revision of this and related species (A. hoffmannii, A. laetum, and A. riparium in Mexico; Asplenium sect. Hymenasplenium) for tropical America. Available evidence suggests that Hymenasplenium forms the sister group for all other species in Asplenium and probably also all the segregate genera of Aspleniaceae, in the sense that this family is circumscribed by nearly all recent workers (Schneider et al., ms.).

92

ASPLENIUM

Asplenium delitescens is distinct from A. abscissum in having creeping dorsiventral rhizomes, non-alate rachises, very broad, truncate marginal pinna serrations, more sori per pinna (10–15 pairs), and longer (5–15 mm) indusia. It is suspected of hybridization with A. laetum in Belize (Schipp 49, MO) and Costa Rica (Go´mez 18620, MO) (Murakami & Moran, 1993).

20. ASPLENIUM DENTATUM Asplenium dentatum L., Sp. Pl. 2: 1080. 1753 (as A. trichomanes-dentatum, to be cited as A. dentatum according to the Tokyo Code, Ex. 14 under Art. 23.8); Syst. Nat., ed. 10, 2: 1323. 1759. Lectotype (chosen by Morton & Lellinger, 1966: 25). Plumier, Traite´ Foug. Ame´r., pl. 101C. 1705. Adams (in Davidse et al., 1995) regarded Plumier’s Desc. Pl. Ame´r., pl. 50, 1693, as lectotype. Figs. 42F, G. Roots thin, fibrous, proliferous; rhizomes erect; rhizome scales blackish throughout, clathrate, 1–2 ⫻ 0.2–0.3 mm, entire; fronds clumped, subdimorphic, the sterile forming a rosette, spreading or prostrate, 3–12 cm long, short-stiped, the fertile central, erect, 10–25 cm long, long-stiped, stipes nearly as long as blades; stipes darkened proximally, greenish distally, mostly 2–15 cm ⫻ 0.5–1 mm, 1⁄5–2⁄3 of frond length, glabrous, adaxially with whitish, thickened, lateral ridges bordering the grooves; blades herbaceous, 1-pinnate, 3–10(–15) ⫻ 1–2.5 cm, usually with subflabellate terminal segments, not proliferous; rachises green throughout, glabrous, not or obscurely winged, but adaxially with two ridges; pinnae oblong to trapeziform (fertile subflabellate), 6–13 pairs, more widely spaced on fertile fronds, 3–11 ⫻ (2–)4–6 mm, the proximal ones short-stalked 1–2 mm, cuneate and unequal proximally, rounded or dentate at apices, margins subentire to serrate acroscopically, entire basiscopically; veins evident, mostly simple or first acroscopic one 1-forked, tips evident adaxially; indument abaxially of scattered appressed, whitish to brown clavate hairs 0.1–0.2 mm long; sori 2–4(–5) pairs, on both sides of midveins; indusia whitish, 2–6 ⫻ 0.5–0.6 mm, margins entire; spores reniform; 2n⫽108 (Jam).

Distribution. Terrestrial or epipetric on calcareous rocks, cave entrances; 100–800 m. USA (Fla); Mexico; Guat, Bel; Berm,

Bah, Gr & L Ant; Col, Ven, Trin. Also cited from Honduras by Adams (in Davidse et al., 1995). Specimens Examined. Camp (Butterwick 282, TEX, UC). Chis (Breedlove 29932, DS, MEXU, 29939, DS, 33892, DS, MEXU, MO, NY, 38185, DS, 42850, DS, MO). Ver (Va´zquez 2548, CHAPA, CIB, NY). Yuc (Butterwick 262, UC; Schott 747, MO, NY, US).

This differs from congeners in Mexico by the subdimorphic fronds, the fertile more erect and twice the length (or more) of the sterile fronds in fully developed plants. The sterile fronds are spreading or arching fronds and form a rosette. Asplenium dentatum superficially resembles species of the A. monanthes group, but it is not closely related, differing by the greenish or yellowgreen stipes and rachises and the lack of true wings adaxially on these same axes.

21. ASPLENIUM DIANAE Asplenium dianae A. R. Sm, sp. nov. Type. Mexico. Tamaulipas: Gomez Farias region, Sep 1965, F. & M. Webster 221 (holotype TEX!). Figs. 44E, F. Ab A. myriophyllo laminis angustioribus, tantum 2-pinnatis, pinnulis (praeter pinnulam acroscopicam basalem in quoque pinna) valde adnatis, non vel vix constrictis basi, laminis chartaceis differt. (Honoring Diana (Dee) Ferguson, who has assisted with details of descriptions of Asplenium for this work, as well as descriptions of species in a few other genera.)

Roots fine, wiry, not proliferous; rhizomes erect; rhizome scales dark brown to blackish, strongly clathrate, to ca. 3 ⫻ 0.8 mm, entire; fronds clumped, 9–13 cm long; stipes brownish, dull, 1–2 cm ⫻ 0.5–0.8 mm, ⬍1⁄5 of frond length, glabrescent or with a few hair-like or linear-lanceolate scales to ca. 4 cells wide, adaxially with narrow green wings for much of their length; blades chartaceous, lanceolate or narrowly elliptic, 2-pinnate, 8–12 ⫻ ca. 2 cm, narrowed proximally, to 6 proximal pairs gradually reduced with the lowermost pair trifid and less than 4 mm long, blade apices confluent, gradually tapered, pinnatifid, not proliferous; rachises brownish or greenish distally, dull, with sparse hair-like scales and green lateral wings 0.2–0.3 mm wide; pinnae deltate-lanceolate, to ca. 20 pairs, to 1.2 ⫻ 0.5 cm, sessile, not articulate, inequilateral, excavate at basiscopic bases, auriculate acroscopically, pinnules to 4 pairs per pinna, strongly ascending ca. 30–40⬚ from costae, simple and undivided except for the largest basal acroscopic one, that cuneate-flabellate and 2–4toothed at apices, to 5 mm ⫻ 2.5 mm, but mostly ca. 1–2 ⫻ 1 mm; veins Ⳳ evident on both sides of blades, simple or occasionally 1-forked, tips prominent adaxially; indument abaxially of scattered, appressed, clavate hairs 0.1 mm long; sori one per ultimate segment; indusia 1–2 ⫻ 0.5–0.6 mm, margins entire; spores reniform, probably 64 per sporangium.

ASPLENIUM EATONII

93

1

Distribution. Habitat unknown; elevation unknown, but Go´mez Farı´as is ca. 500 m. Known only from the type. Of Mexican species, this is most similar to A. myriophyllum. It differs from that in the narrower, less divided blades (only 2-pinnate); strongly adnate pinnules (except the basal acroscopic one of each pinna), not or only slightly constricted at the bases; and chartaceous (vs. thin-herbaceous) blades. The spores are reniform and normal in appearance; although in the type specimen most sporangia have shed their spores, some were found with at least 45 spores, indicating that this species is probably sexual, with 64 spores per sporangium. The closest relative of A. dianae may be A. sciadophilum Proctor, endemic to Jamaica. That species, although basically 2-pinnate like A. dianae, has larger fronds (20–40 ⫻ 5.5 cm) and much longer stipes (4–11 cm), and is, like A. myriophyllum, thintextured; it also has lighter green blades than A. dianae. Additional relatives, all apparently less closely related, include several species mentioned by Adams (in Davidse et al., 1995) under A. cladolepton Fe´e, i.e., A. barclayanum C. D. Adams from Isla del Coco, A. macraei Hook. & Grev., from Hawaii, and A. haenkeanum (C. Presl) Hieron., from Colombia to Peru (Tryon & Stolze, 1993). The type was regrettably mounted abaxial side down with nearly indestructable glue, causing many of the important characters to be obscured. Surgery on the specimen revealed that all of the half dozen developed fronds are fully fertile.

22. ASPLENIUM DISSECTUM Asplenium dissectum Sw., Prodr. 130. 1788. Type. Jamaica. Swartz s.n. (S, photo US!; isotype B-Willd. 19916!, microfiche UC!). Fig. 31G. Asplenium bissectum Sw., J. Bot. (Schrader) 1800(2): 55. 1801, nom. superfl. (based on A. dissectum Sw.).

Roots fibrous, wiry, not proliferous; rhizomes creeping; rhizome scales blackish or dark brown, sparsely ciliate, subclathrate (lumina somewhat occluded), tips long-attenuate, mostly 3–6 ⫻ 0.2–0.5 mm, entire; fronds clumped, mostly 30–60 cm long; stipes gray-brown to atropurpureous, dull, 6–18 cm ⫻ 1–1.5 mm,

⁄4–1⁄3 of frond length, with scattered to dense short (0.1–0.5 mm) glandular hairs and also attenuate or hair-like clathrate scales, not alate; blades thin-chartaceous, 25–50 ⫻ 5–12(–18) cm, 1-pinnate, lanceolate, bases not or only slightly reduced, apices gradually reduced and pinnatifid, not radicant; rachises atropurpureous, dull to sublustrous, with numerous attenuate to hairlike scales, not winged; pinnae narrowly lanceolate, 25–60 pairs per frond, 3–9 cm ⫻ 6–10 mm, less than 5 mm wide in distal 1⁄2, stalked to 1 mm, bases inequilateral, excised and narrowly cuneate basiscopically, truncate and sometimes slightly auriculate acroscopically, apices long-attenuate, margins lacerate or often bilacerate (longer teeth shallowly notched) with sharp angled teeth; veins evident on both sides, mostly 1-forked, 2-forked at bases, simple toward pinna tips, diverging at ca. 10⬚ angle from the costae; indument abaxially of scattered brownish stellate scales mostly 0.1–0.5 mm long on and between veins, the larger ones with attenuate tips and fringed bases; sori 4–10 pairs per pinna, on both sides of midveins, almost parallel to and appearing borne on the pinna midribs; indusia red-brown, 1.5–3 ⫻ 0.3–5 mm, margins entire; spores reniform.

Distribution. Epiphytic in wet montane forests; 1200–1450 m. Mexico; Guat, Nic, CR, Pan; Cuba, Jam, Hisp; Col, Ven, Ec, s Braz. Specimens Examined. Oax (Mickel 1057, ENCB, ISC, US, 5728, 5741, 7095, NY).

Among the 1-pinnate members of sect. Sphenopteris (scaly rachises and sori at narrow angles with respect to the costae and costules), the thin blade texture, deeply lacerate or bilacerate pinnae, pinnatifid and very gradually tapering blade apices, more numerous pinna pairs, and glandular hairs on the stipes readily distinguish A. dissectum from A. serra, the closest Mexican relative.

23. ASPLENIUM EATONII Asplenium eatonii Davenp., Bot. Gaz. 21: 258. 1896. Figs. 40E, F. Asplenium cicutarium Sw. var. paleaceum Davenp., Bot. Gaz. 19: 390. 1894. Type. Mexico. [Jalisco:] Colima Mts., Pringle 5531 (GH!).

94

ASPLENIUM

Differs from A. achilleifolium in having up to ca. 5 (or more) pairs of reduced proximal pinnae, the smallest sometimes only 1–2 cm long. Differs from A. blepharophorum in lacking buds in the axils of distal pinnae, narrower scales mostly two cells wide along the rachises and costae abaxially, and in the less densely scaly stipes and rachises.

liferous bud, the buds usually dormant and developing further only in very moist situations, pinnules oblong, subentire to toothed, each one sometimes with a bud at the tip; veins obscure, tips not evident adaxially; indument abaxially of scattered whitish to tan clavate hairs 0.1 mm long; sori to ca. 15 per pinna, on both sides of costae, 1–2(–4) per pinnule; indusia whitish, 1–1.5 ⫻ 0.3–0.5 mm, margins suberose to erose; spores reniform; 2n⫽144 (India).

Distribution. In wet montane forests; 300–1500 m. Mexico. Selected Specimens Examined. Col (Lott 2925, MO, UC; Sanders 10889, ENCB, MO, UC). Jal (McVaugh 16205, MEXU, US; Mexia 1252, CAS, DS, MO, NY, UC, US, 1797, MO, NY, US). Mich (Langman 3283, US). Nay (McVaugh 18991, IEB, NY; Palmer 1950, ARIZ, NY, UC, US). Sin (Gentry 7277, ARIZ, DS, UC, US; Ortega 5097, MEXU, US).

This species is most closely related to A. achilleifolium and differs by the characters given in the diagnosis above. These two, with the addition of A. blepharophorum and A. solmsii, form a natural group in Mexico and Mesoamerica.

24. ASPLENIUM EXIGUUM Asplenium exiguum Bedd., Ferns S. India 49, t. 146. 1863. Type. India. Nilgris, Beddome s.n. (K!). Fig. 37H. Athyrium gracile E. Fourn., Mexic. Pl. 1: 102. 1872. Asplenium gracile (E. Fourn.) Hemsl., Biol. Cent.-Amer., Bot. 3: 634. 1885, hom. illeg., non Don, 1825, nec Fe´e, 1852, nec Pappe & Raws, 1858. Type. Mexico. [Distrito Federal:] “Vallis Mexicensis pr. Pedregal,” Bourgeau 252 (P!–3 sheets; isotypes MO!, NY!). Asplenium glenniei Hook. & Baker, Syn. Fil., ed. 2, 488. 1874. Syntypes. Mexico. Glennie s.n. (K!), Bourgeau 252 (K!; isosyntypes P, US!; isosyntype and frag. NY!).

Roots fibrous, not proliferous; rhizomes short-creeping or erect; rhizome scales black, clathrate with occluded lumina, 2.5–3 ⫻ 0.5–0.8 mm, entire or nearly so; fronds clumped, 6–18 cm long; stipes castaneous, or sometimes darkened only at the base, 1–5 cm ⫻ 0.4–1 mm, 1⁄10–1⁄8 of frond length, sparsely scaly, the scales hair-like or lanceolate, not winged; blades 5–14 ⫻ 1.5–2.5(– 3.5) cm, narrowly elliptic, 1–2-pinnate-pinnatifid, proximal pinnae gradually reduced, the apices sometimes proliferous, flagelliform; rachises brown proximally, greenish distally, with scattered hair-like scales 1–3 mm long, not winged; pinnae 10–18 pairs, 6–16 ⫻ 5–9 mm, apices emarginate, each with a pro-

Distribution. Terrestrial or epipetric on non-calcareous rocks, damp cliffs, and crevices of ledges; 1000–2600(–3500) m. USA (Ariz); Mexico; Guat; n India, s China, Philippines. Selected Specimens Examined. Chih (Pringle 833, DS, UC, US). Chis (Breedlove 26979, DS). Coah (Wendt 2046, MEXU). DF (Lyonnet 233, MEXU, NY, US). Dgo (Pringle 10150, MEXU, MO, NY, UC, US). Gro (Sharp 441421, MEXU, US). Gto (Rzedowski 41582, ENCB, IEB). Jal (Bautista 17149, GUAD, UC). Me´x (Lyonnet 521100001, MEXU). NL (Cedillo Montoya s.n., 6 Sep 1986, UNL). Oax (Mickel 6488, ENCB, UC). Qro (Rzedowski 46690, IEB, XAL). SLP (Parry & Palmer 1003.5, MO, NY). Son (Lloyd 503, US). Tam (Heredia 00260, ITCV). Unverified, Doubtful, or Mistaken Reports. Ags (Gonza´lezAdame 70, HUAA, cited by Siqueiros-Delgado & Gonza´lez-Adame, 2004, but not verified). Hgo (reported by Mickel & Beitel, 1988, but not verified).

Asplenium exiguum has small proliferous apical buds on the pinnae and sometimes pinnules, and also has subterminal rachis buds, but these are tiny and easily overlooked. The rachises may also become flagelliform and naked when humidity is sufficiently high (Mickel, 1976). No other Mexican spleenwort is so diversified in its means of vegetative reproduction. The report of this species from Nuevo Leo´n by Hinton and Hinton (1994), based on Hinton 20142, applies to A. ultimum. Asplenium exiguum was correctly reported from Nuevo Leo´n by Rodrı´guez-Lo´pez and Arreguı´n-Sa´nchez (1992), citing Sa´nchez s.n., herb. no. 18919, and Cedillo M. s.n., herb. no. 18931 (both UNL!).

25. ASPLENIUM FEEI Asplenium feei Kunze ex Fe´e, Me´m. Foug. 5: 194. 1852. Type. Mexico. Galeotti s.n. (P!, photo US). Fig. 35L.

ASPLENIUM FIBRILLOSUM Asplenium sanguinolentum Kunze ex Mett., Abh. Senckenberg. Naturf. Ges. 3: 142, t. 4, f. 10. 1859. Lectotype (chosen by Morton & Lellinger, 1966: 10). Brazil. Beyrich s.n. (L). Asplenium sarcodes Maxon, Contr. U.S. Natl. Herb. 10: 494, t. 56, f. 3. 1908. Type. Cuba. Oriente: La Perla, Maxon 4390 (US!).

Roots coarse, fibrous, not proliferous; rhizomes suberect, densely scaly at apices; rhizome scales red-brown, clathrate, 5–10 ⫻ 1.5–3(–4) mm, entire or with occasional marginal teeth; fronds clumped, 20–90 cm long; stipes brown, dull, (8–)15–30 cm ⫻ 2–4 mm, 1⁄4–1⁄2 of frond length, at bases with dense narrow, twisted or curled clathrate scales 3–4 mm long, not winged; blades thickherbaceous, 1-pinnate, not reduced proximally, 25–35(–60) ⫻ 10– 25 cm, apices subconform, not proliferous; rachises green to brown, dull, with scattered scales 1.5–2 mm long, narrowly winged; pinnae lanceolate, 6–14 pairs, 4–12 ⫻ (0.8–)1.5–2.5 cm, stalked 1–2(–4) mm, bases equilateral to inequilateral, excavate basiscopically, sometimes roundish-auriculate acroscopically, apices attenuate, margins crenate, serrate, or biserrate; veins 1-or 2-forked, tips evident adaxially; indument abaxially of brown, clathrate, attenuate to stellate-attenuate scales to 1 mm on and between veins, scales often pectinate at bases; sori elliptical, 10–12 pairs per pinna, on both sides of midveins; indusia 3–7 ⫻ 1–2 mm, vaulted, margins entire; spores reniform.

95

Asplenium anisophyllum Kunze (Linnaea 10: 511. 1836.), type from tropical Africa, resembles this species closely, and is possibly conspecific (Moran & Smith, 2001). If this is true, the name A. anisophyllum has priority. However, most African specimens seen (except from Sa˜o Tome´) have more incised pinnae than do American specimens of A. feei. Also, unpublished molecular data by Schneider et al. indicate that A. feei and A. anisophyllum, although closely related, may not be sister species.

26. ASPLENIUM FIBRILLOSUM Asplenium fibrillosum Pringle & Davenp. ex Davenp., Bot. Gaz. 21: 257, pl. 18, f. 1–4. 1896. Type. Mexico. Morelos: above Cuernavaca, Pringle 6191 (GH!; isotypes CAS!, MEXU!–2 sheets, MO!, NY!, UC!, US!). Figs. 39G, H. Roots filamentous, fibrous, not proliferous; rhizomes suberect; rhizome scales black, clathrate, 2–4 ⫻ 0.3–0.5 mm, entire; fronds clumped, 7–15 cm long; stipes castaneous, lustrous, 1.5–3 cm ⫻ 0.4–0.5 mm, ca. 1⁄8 of frond length, with brown, flexuous, hairlike (uniseriate at tips) scales, not winged; blades 1-pinnate, linear, 6–13 ⫻ 1.1–1.5 cm, the bases slightly to decidedly reduced, the apices tapering and with a terminal hastate or basally pinnatifid segment, not proliferous; rachises castaneous, lustrous, with scattered hair-like (uniseriate at tips) scales, adaxially with wings 0.1 mm wide or less; pinnae oblong, dimidiate, 15–25 pairs, the proximal 1–3 pairs often reduced and sometimes deflexed, 5–7 ⫻ 3–4 mm, sessile, bases slightly auriculate, apices obtuse, margins crenulate to dentate; veins obscure, but tips evident adaxially; indument abaxially of scattered appressed, whitish to tan clavate hairs 0.1–0.2 mm long; sori 1–4 pairs per pinna, on both sides of midveins; indusia 1–2 ⫻ 0.5–0.8 mm, margins fimbriate with processes 0.4–0.5 mm long; spores reniform, 64 per sporangium (Pringle 6191, UC).

Distribution. Epiphytic and terrestrial in montane rain forests; 800–1250 m. Mexico; Guat (no specimens cited by Stolze, 1981, but Bernoulli hb. guatemalens. 407, NY, is this species), Hond, Salv, Nic, CR, Pan; Cuba, Hisp, PR; Col, Ven, Sur, Galapagos, Braz, Bol. Also reported from French Guiana by Bogan et al. (1997). Specimens Examined. Chis (Breedlove 33116, DS, MEXU, 68692, CAS; Purpus 6765, MO, UC, US, 7110, UC, 7242, UC, US). Ver (Lira 16, ENCB, MEXU, XAL; Ramı´rez 850, XAL).

The larger frond size, numerous scales on stipes and rachises, long rhizome scales, and short and wide indusia (3–7 ⫻ 1–2 mm) distinguish A. feei from A. tuerckheimii and A. riparium, both of which have subconform terminal pinnae. Asplenium riparium has creeping rhizomes, veins running to the pinna margins, and fewer (6–8) pairs of sori per pinna, while A. tuerckheimii has fewer pinnae (3–6), each with an equilateral base. Fronds of A. feei often dry a distinctive dark green.

Distribution. Terrestrial and epipetric in ravines and barrancas in pine-oak forests, on mossy banks, and on sheltered ledges and grottos in lava fields; 1700–3900 m. Mexico. Selected Specimens Examined. Ags (McVaugh 18396, IEB, MEXU, NY). DF (Pringle 8791, CAS, MEXU, MO, NY, UC, US). Dgo (Keil 8879, NY). Gro (Lorea 4254, NY, UC). Gto (Correll & Correll 28820, MEXU, US). Jal (McVaugh 13097, MICH, US). Me´x (Arreguin 622,

96

ASPLENIUM

ENCB). Mich (Arse`ne 6794, MEXU). Mor (Pringle 11257, MO, NY). Qro (Medina-Cota & Barrios 3060, IEB).

1080, US, 5704, NY, UC). Pue (Riba s.n., Jardı´n Botanico 894, MEXU). Ver (Purpus 5789, UC, US, Purpus 8254, MO, NY).

The small plant size, hair-like clathrate stipe and rachis scales, and fimbriate indusia distinguish this species. The closest relative is very likely A. blepharodes, from Baja California, which see. Diminutive forms of A. castaneum are also similar, but that species has less scaly rachises and entire to slightly erose indusia. Rzedowski 47913 (IEB), from Guanajuato, has less strikingly fimbriate indusia than typical forms but is certainly A. fibrillosum, rather than A. castaneum, as cited by Dı´az-Barriga and PalaciosRios (1992).

Asplenium flabellulatum is part of the A. radicans complex, a group having castaneous, lustrous stipes and long-flagelliform, proliferous blade tips. We no longer maintain var. partitum as distinct from var. flabellulatum, as was done by Smith (1981) and Adams (in Davidse et al., 1995). Some of the collections cited by Mickel and Beitel (1988) as A. uniseriale (e.g., Mickel 1080) are A. flabellulatum. For additional discussion, see under A. radicans.

28. ASPLENIUM FORMOSUM 27. ASPLENIUM FLABELLULATUM Asplenium flabellulatum Kunze, Linnaea 9: 71. 1834, non Klotzsch, 1847, nec Mett., 1859. Fig. 44H. Asplenium flabellulatum Kunze var. dentatum Klotzsch, nom. superfl., Linnaea 20: 357. 1847. A. radicans L. var. dentatum (Klotzsch) Bonap., Notes Pterid. 7: 349. 1918. Lectotype (chosen by Morton & Lellinger, 1966: 37). Peru. Pampayaco, Poeppig 1145 (B!, frag. NY!; isolectotype W, photo BM). Asplenium flabellulatum Kunze var. partitum Klotzsch, Linnaea 20: 357. 1847. Asplenium partitum (Klotzsch) C. Chr., Index Filic. 125. 1905. Asplenium radicans L. var. partitum (Klotzsch) Hieron., Bot. Jahrb. Syst. 34: 464. 1904. Lectotype (chosen by Morton & Lellinger, 1966: 37). Venezuela. Caracas, Otto 651 (B!, frags. NY!, US).

Differs from Asplenium radicans by: rhizome scales 3–5 ⫻ 0.5– 0.8 mm; pinnules lobed or deeply dentate (especially proximal pinnules) to basal acroscopic pinnules lobed, each with a free basal segment (thus the blades 3-pinnate); ultimate segments cuneate or petiolulate proximally; sori 1–4 per segment; indusia 1–2 ⫻ 0.3 mm, margins suberose.

Distribution. Terrestrial, in wet montane forests; 600–2450 m. Mexico; Guat, Hond, CR, Pan; Cuba, Jam, Hisp; Col, Ven, Ec, Peru. Reported from El Salvador by Seiler (1980) and from Nicaragua by Go´mez (1976), both as var. A. radicans var. partitum. Also reported from Paraguay by Adams (in Davidse et al., 1995). Selected Specimens Examined. Chis (Breedlove 15409, DS, US, 21809, 31313, DS, MEXU, NY). Oax (Maya J. 1097, CHAPA, NY; Mickel

Asplenium formosum Willd., Sp. Pl., ed. 4, 5(1): 329. 1810. Type. Venezuela. Caracas, Bredemeyer s.n. (B-Willd. 19908!, photo US!). Fig. 33J. Asplenium subalatum Hook. & Arn., Bot. Beechey Voy. 312, t. 71. 1838. Type. Mexico. Nayarit: Tepic, Dec 1827–Feb 1828, Lay s.n. [Beechey] (K!).

Roots filiform and wiry, fibrous, not proliferous; rhizomes suberect to erect; rhizome scales bicolorous, central portion black, occluded, margins light brown, clathrate, 2–3 ⫻ 0.3–0.5 mm, entire; fronds clumped, mostly 10–45 cm long; stipes castaneous to atropurpureous, lustrous, mostly 1–5 cm ⫻ 0.5–1 mm, 1⁄10–1⁄5 of frond length, with rare hair-like scales 1–2 mm long and scattered appressed tan hairs 0.1–0.2 mm long, adaxially with tan wings 0.1–0.3 mm wide; blades chartaceous, 1-pinnate, linear, (6–)15–30 ⫻ 2–3.5 cm, bases with 5–10 pairs of gradually reduced pinnae, apices pinnatifid, not proliferous; rachises castaneous to atropurpureous, lustrous, with scattered tan appressed hairs 0.1 mm long, adaxially with tan wings 0.1–0.3 mm wide; pinnae oblong, dimidiate, (15–)25–50 pairs, (6–)10–18 ⫻ 3–6 mm, sessile, bases inequilateral, apices acute or lacerate, acroscopic margins bidentate or bilacerate, sinuses 1–3 mm deep, distal part of basiscopic margins also often toothed or lacerate; veins 1-forked, tips evident adaxially; indument abaxially of tan, appressed, clavate hairs 0.1 mm long; sori mostly 1–3 per pinna, on basiscopic side of midveins and usually parallel to basiscopic margins, or on both sides and then often 2–3 pairs; indusia 2–5 ⫻ 0.5–0.8 mm, margins erose to subentire; spores reniform; 2n⫽72 (Jam).

ASPLENIUM GENTRYI

97

mm, long-attenuate, pinnules (3–)4–5(–9) pairs per pinna, acute or obtuse, acroscopic pinnule of proximal pinnae with at least a basal lobe cut almost to costule; segments cuneate proximally, lacerate to toothed distally, 1–3 veins per segment; 2n⫽144 (Oax, Jam).

Distribution. Terrestrial or epipetric, especially on rocks near streams; 50–1250(–2000) m. Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Gr & L Ant; Col, Ven, Guy, Ec, Galapagos, Peru, s Braz, Bol, Parag, n Arg; Africa, s India, Sri Lanka. Also reported from French Guiana by Adams (in Davidse et al., 1995). Selected Specimens Examined. Chis (Breedlove 70707, CAS). Col (Orcutt 4638, DS, NY, US). DF (without collector and number, FCME). Dgo (Gentry 5264, ARIZ, DS, MEXU, MO, NY). Gro (Mexia 8858, ARIZ, CAS, MO, NY, UC, US). Jal (Mexia 1632a, DS, UC, US). Me´x (Santiago S. s.n., FCME). Mich (King & Soderstrom 4870, MEXU, NY, UC, US). Mor (Pringle 11255, MEXU, UC). Nay (McVaugh 18915, CAS, IEB, NY). Oax (Mickel 5159, NY, UC, US). Pue (Maldonado s.n., FCME). Rev (Moran 5776, MEXU). Tab (Cowan 3489, UC). Ver (Purpus 2178, MO, UC, US).

Asplenium formosum is distinguished from congeners in Mexico by the deeply incised pinna margins, especially acroscopically, and its frequent occurrence on moist rocks by streams at low elevations. It has a remarkably wide distribution, extending to Africa and south India (Moran & Smith, 2001). It is a member of the Asplenium trichomanes group, but is rather isolated within this group and not easily confused with any other species.

29. ASPLENIUM FRAGRANS Asplenium fragrans Sw., Prodr. 130. 1788.

Fig. 36G.

Asplenium auritum Sw. var. tripinnatum E. Fourn., Mexic. Pl. 1: 107. 1872. Asplenium cuspidatum Lam. var. tripinnatum (E. Fourn.) C. V. Morton & Lellinger, Mem. New York Bot. Gard 15: 29. 1966. Type. Jamaica. Swartz s.n. (S, photo US; isotype UPS-Herb. Thunb. 24802, microfiche UC!). Asplenium foeniculaceum Kunth, Nov. Gen. Sp. 1: 15. 1815. Asplenium cuspidatum Lam. var. foeniculaceum (Kunth) C. V. Morton & Lellinger, Mem. New York Bot. Gard. 15: 29. 1966. Type. “Novae Andalusiae” [Venezuela]. Humboldt & Bonpland s.n. (P-Humb.!; isotype B-Willd. 19860!). Asplenium coriifolium Liebm., Mexic. Bregn. 249 (reprint 97). 1849. Lectotype (chosen by Smith, 1981: 43). Mexico. Oaxaca: Tepitongo and Totontepec, Liebmann s.n. [Pl. Mex. 2270, Fl. Mex. 336] (C!; possible isotypes K!, US!).

Differs from A. cuspidatum by: blades 2-pinnate-pinnatifid to 3-pinnate, narrowly to broadly deltate; pinnae petiolulate 2–5

Distribution. Epiphytic, infrequently epipetric, rarely terrestrial; (300–)1500–3050 m. Mexico; Guat, Hond, Salv, Nic, CR (Smith 2030, UC), Pan; Gr Ant; Col, Ven, Ec, Peru, Bol. Selected Specimens Examined. Chis (Davidse et al. 29871, MEXU). Dgo (Sa´nchez 751, MEXU). Gro (Hinton 10732, CAS, MO, NY). Jal (Iltis et al. 29562, IEB, UC). Me´x (Hinton 2390, DS, NY). Mich (McVaugh 22799, CAS, IEB, MEXU, NY). Mor (Pringle 8955, ARIZ, ASU, CAS, ENCB, MEXU, MO, NY, UC). Nay (Bartholomew 2645, CAS, MO, NY). Oax (Davidse et al. 30242, MO, UC). Pue (Tenorio L. 15420, MEXU). Sin (Breedlove 17109, CAS, NY). Ver (Mu¨ller 688, NY). Unverified, Doubtful, or Mistaken Reports. Qro (Ferna´ndez N. 2121a, ENCB, uncertain determination). Tam (reported in the range by Sharp, 1954: 74, but the specimen is most likely A. cuspidatum).

The name A. cuspidatum has often been applied to the species here called A. fragrans, e.g., by Smith (1981), Mickel and Beitel (1988), and Mickel (1992), but Adams (in Davidse et al., 1995) regarded it as distinct, and we now agree. Most authors have treated it as a synonym of A. cuspidatum var. tripinnatum (e.g., Morton & Lellinger, 1966, for Venezuela; Tryon & Stolze, 1993, for Peru; Proctor, 1985, for Jamaica). Asplenium fragrans is a more dissected species, having blades that are 3–4-pinnate, rather than 2-pinnate to 2-pinnate-pinnatifid at the bases. See additional discussion under A. auritum. Juvenile or young sterile specimens have nearly filiform ultimate segments, e.g., Mickel 4140, 5275 (UC), from Oaxaca.

30. ASPLENIUM GENTRYI Asplenium gentryi A. R. Sm., sp. nov. Type. Mexico. Chihuahua: Sierra Charuco, Arroyo Hondo, 4500–5500 ft, Gentry 8064 (holotype UC!; isotypes ARIZ!, US!). Fig. 42E. Ab A. monanthes frondibus plerumque parvioribus, soris usque 7-jugatis utrinque costarum, rhachidibus atratis vel atropurpureis differt; ab A. resilienti hydathodis conspicuis adaxialiter, alis brunneolis rhachidum differt. (Honoring

98

ASPLENIUM

Howard Scott Gentry, 1903–1993, student of Agave, agricultural explorer for the U.S. Department of Agriculture, especially in the Sonoran Desert region, also affiliated with the Desert Botanical Garden in Phoenix.)

Roots thin, wiry, not proliferous; rhizomes erect; rhizome scales bicolorous, central portion occluded, blackish, margins brown, clathrate or subclathrate, apices attenuate, 2–4 ⫻ 0.1–0.4 mm, entire; fronds clumped, mostly 7–25(–40) cm long; stipes black to atropurpureous, 1–10 cm ⫻ 1.5 mm, 1⁄10–1⁄3 of frond length, glabrous, with narrow brownish ridges 0.1 mm wide flanking the adaxial side; blades 1-pinnate, linear, 6–30 ⫻ 1.5–3 cm, apices pinnatifid, not proliferous; rachises black to atropurpureous, lustrous, glabrescent or with sparse, twisted, hair-like scales 1–2 mm long, adaxially with brownish wings 0.1–0.2 mm wide; pinnae quadrangular to oblong, 20–40 pairs, 7–15 ⫻ 2–6 mm, reduced proximally (lowest deltate, ca. 1⁄3–1⁄2 the length of the longest ones), sessile, dimidiate, basiscopic edges straight or curved upward, acroscopic edge slightly auriculate at the base, crenulate or subentire, sinuses to 0.8 mm deep; veins mostly simple, 1-forked at the acroscopic base of segments, tips evident adaxially; indument abaxially of scattered, appressed, whitish to tan, clavate hairs 0.1–0.2 mm long; sori up to 7 pairs per pinna, on both sides of costae; indusia mostly 2–3 ⫻ 0.5–0.6 mm, margins entire; spores reniform.

31. ASPLENIUM HALLBERGII Asplenium hallbergii Mickel & Beitel, Mem. New York Bot. Gard. 46: 58. 1988. Type. Mexico. Oaxaca: Distrito Pochutla, Oaxaca–Pochutla rd., Km 153 at Campamento Rı´o Molino, Mickel 1280 (NY!). Figs. 33L, M. Roots wiry, fibrous, not proliferous; rhizomes short-creeping; rhizome scales bicolorous, central portion black, heavily sclerotic, lumina occluded, with narrow brown clathrate wings, 1–2 ⫻ 0.5 mm, entire; fronds clumped, (8–)12–30(–40) cm long; stipes castaneous to atropurpureous, lustrous, 2–4 cm ⫻ 0.5–1.2 mm, 1⁄10– 1 ⁄3 of frond length, glabrous, adaxially with wings entire proximally, 0.1–0.2 mm wide, distally the wings fringed with minute castaneous papillae; blades linear, 1-pinnate, (10–)18–25(–35) cm ⫻ 6–18(–22) mm, apices pinnatifid, not proliferous; rachises castaneous, lustrous, glabrous, with wings 0.1–0.3 mm wide, fringed with minute papillae less than 0.05 mm long; pinnae oblong or rhomboid, dimidiate, 20–40 pairs, 4–8(–12) ⫻ 2–5 mm, sessile, articulate, rachis color running onto pinna stalks, bases slightly auriculate, apices rounded, margins entire to slightly crenatedentate at tips; veins obscure, but tips evident adaxially; indument abaxially of scattered, tan, appressed, clavate hairs 0.1–0.2 mm long; sori 1–4 pairs per pinna on both sides of midveins; indusia 2–3(–4) ⫻ 0.5–0.8(–1) mm, margins erose to suberose; spores reniform.

Distribution. Terrestrial in rocky canyons in pine-oak forests; 1300–2600 m. Mexico. Specimens Examined. Ags (de la Cerda L. 863, IEB). Chih (Correll & Gentry 22963, LL, 23032, MO, NY, UC, US; Fishbein 1807, 1817, ARIZ; Nesom & Vorobik 5603, 5726, TEX; Spellenberg et al. 8748, UC). Dgo (Ortega 5314, US; Reeder & Reeder 2477, US, 2494, MEXU, US; Sa´nchez 748a, US). Sin (Breedlove 18467, CAS, DS, NY; Correll & Correll 28851, LL, 28873, LL, MEXU; Gentry 7233, ARIZ, US; Gentry & Arguelles 18189, ARIZ; Gonza´lez O. 6028, MEXU; Ownbey & Ownbey 1942, 1943, US; Sanders 21121, UCR). Son (Fishbein 1724, ARIZ; Martin et al. s.n., ARIZ; Turner 74-375, ARIZ).

Specimens of A. gentryi have been identified in herbaria as either A. monanthes or A. resiliens, from which it seems amply distinct by the characters given in the diagnosis. Apparently, this species grows with A. sanchezii. The two were collected at the same general locality and date by Sa´nchez, but A. gentryi is readily distinguished by having larger fronds and pinnae, more numerous pinna pairs, and blackish stipes and rachises.

Distribution. Terrestrial in pine-oak forests; 1500–3200 m. Mexico. Also reported from Guatemala by Mickel and Beitel (1988), but not verified; Adams (in Davidse et al., 1995) did not treat the species for Mesoamerica. Selected Specimens Examined. Ags (Cerda L. 863, ENCB, IEB). Chih (Fraser-Jenkins 13506, NY). DF (Pringle 8754, UC). Dgo (Breedlove 18819, CAS, NY). Hgo (Mun˜oz B. 3, ENCB). Gro (Lorea 2284, FCME). Gto (Dı´az B. 3971, ENCB, IEB). Jal (Corres Zepeda 16112, UC). Me´x (Rose & Painter 7204, US). Mich (King & Soderstrom 5096, UC, US). Mor (Copeland herb. 67 p.p., UC, US). Oax (Mickel 4440, ENCB, NY, UC). Pue (Tenorio L. 15120, MEXU). Qro (Dı´az B. & Carranza 7360, IEB). SLP (Parry & Palmer 978 1⁄2B, NY). Tlax (Sa´nchez 368, CIB). Ver (Ventura A. 185, ENCB).

Asplenium hallbergii is very closely related to A. monanthes, and often identified as that species or as A. castaneum. In general,

ASPLENIUM HARRISII

it has smaller (shorter and narrower) pinnae than A. monanthes, and can be readily distinguished by the papillate rachis wings and the occluded lumina of the smaller (1–2 mm) rhizome scales (which are clathrate in A. castaneum). The indusia are not as wide as those of A. castaneum and the pinnae are more oblong. Asplenium polyphyllum has larger, clathrate rhizome scales, nonpapillate rachis wings, and subterminal proliferous buds in axils of distal pinnae. Like A. castaneum and A. polyphyllum, A. hallbergii occurs at relatively high elevations. Larger frond size, occluded rhizome scales, and papillate rachis wings distinguish A. hallbergii from A. oligosorum and A. olivaceum. Prominent adaxial hydathodes separate A. hallbergii from A. resiliens and A. palmeri. Tejero-Dı´ez 2865 (IZTA), from Edo. Me´xico, agrees with A. hallbergii in nearly all characters except the lack of papillae along the adaxial rachis wings; the spores seem well formed. Gonza´lez T. 418 (LL), from Jalisco, also lacks papillae on the rachis wings, but has many collapsed sporangia. We have seen a number of mixed collections of A. hallbergii and A. monanthes, e.g., Purpus s.n., Feb 1903 (UC), from Ixtaccihuatl; these mixed gatherings give us increased confidence of the distinctness of A. hallbergii.

32. ASPLENIUM HARPEODES Asplenium harpeodes Kunze, Linnaea 18: 329. 1844. Fig. 34J. Asplenium erectum Bory var. harpeodes (Kunze) Mett., Abh. Senckenberg. Naturf. Ges. 3: 166. 1859. Lectotype (chosen by Morton & Lellinger, 1966: 15). Mexico. Leibold 26 (B!–2 sheets, photo US!; isolectotype P!). Asplenium pendulum Fe´e, Me´m. Foug. 5: 196. 1852. Type. Mexico. Veracruz: Galeotti 6407 (P?; isotype BR!). Asplenium donnell-smithii Christ, Bot. Gaz. 20: 544. 1895. Type. Guatemala. Dept. Quiche: Heyde & Lux s.n. [Donn. Sm. 4678] (P?; isotype US!, frag. NY!).

Roots wiry and thin, fibrous, not proliferous; rhizomes suberect; rhizome scales brown, clathrate, tips long-attenuate, 6–10 mm long, margins with occasional teeth; fronds clumped, arching or pendent, (25–)35–70(–100) cm long; stipes castaneous to black, dull or sublustrous, 7–16 cm ⫻ 1–2 mm, 1⁄10 of frond length, glabrous, without wings; blades thin-chartaceous, 1-pinnate with a non-radicant, pinnatifid apices, (25–)40–70(– 90) ⫻ 6–12 cm, proximal pinnae deflexed, linear; rachises castaneous to black, lustrous, glabrous, with narrow wings 0.2–0.3 mm wide; pinnae lanceolate, falcate, over 30 pairs, (3–)4.5–7 ⫻ 0.8–1.5 cm, sessile, basal basiscopic margins cuneate and excavate, apices long-attenuate, distal 1⁄3–1⁄2 of pinnae less than 3 mm wide, margins serrate, sinuses 1–3 mm deep; veins mostly simple except for proximal portion of pinnae, vein tips evident adaxially; indument apparently absent, blades essentially glabrous; sori (8–)10–15 pairs per pinna (usually none on basal, forked veins), on both sides of midveins; indusia 2–4 ⫻ 0.2–0.5 mm, margins entire; spores reniform.

99

Distribution. Epiphytic in wet forests; (1000–)1600–2800 m. Mexico; Guat, Hond, Salv, Nic, CR, Pan; Jam, Hisp; Col, Ven, Guy, Ec, Peru, s Braz, Bol. Reported from Nicaragua by Go´mez (1976); also reported from French Guiana by Adams (in Davidse et al., 1995). Selected Specimens Examined. Chis (Breedlove 23234, DS, MEXU, NY; Somers et al. 225, NY). Gro (Lorea 2003, 2270, FCME). Oax (Mickel 4255, 4303, MEXU, NY). Pue (Ventura A. 13642, ENCB, MEXU, XAL). Ver (Copeland herb. 60, MEXU, UC, US; Ventura A. 7417, ARIZ, 7962, CHAPA).

This is most similar to A. miradorense and A. potosinum, differing in its epiphytic habit, darker and more lustrous stipes and rachises, and falcate, less deeply serrate, more attenuate pinnae. From A. pteropus and A. pseuderectum, A. harpeodes differs in its non-alate rachises, or rachises adaxially narrowly winged (vs. laterally and broadly winged in A. pteropus and A. pseuderectum).

33. ASPLENIUM HARRISII Asplenium harrisii Jenman, Gard. Chron., ser. 3, 17: 68. 1896. Type. Jamaica. Blue Mt. Peak, Harris s.n. [7474] (isotype K!). Figs. 42H, J. Roots filamentous, wiry, not proliferous; rhizomes erect; rhizome scales bicolorous, center dark, strongly clathrate, margins narrow, lighter, lumina not occluded, 1–2 ⫻ 0.5 mm, entire; fronds clumped, to ca. 15 cm long; stipes greenish to tan, dull, usually less than 1.5 cm ⫻ 0.5 mm, ca. 1⁄10 of frond length, with occasional red-brown hairs ca. 0.2 mm long and rare red-brown hair-like scales 1.2 ⫻ 0.2 mm, adaxially with green wings ca. 0.1– 0.2 mm wide; blades membranaceous, 1-pinnate, 7–14 ⫻ 1.5 cm, linear, usually with flagelliform apices eventually bearing small buds; rachises greenish, dull, with rare red-brown hairs ca. 0.2 mm and reduced scales 1 or 2 cells wide and 0.2–1 mm long, both hairs and scales with enlarged, darkened apical cells, adaxially with green wings ca. 0.1 mm wide; pinnae dimidiate, medial pinnae oblong to almost round, mostly 15–30 pairs, 3–7 ⫻ 2.5– 4 mm, short-stalked to 0.5 mm, proximal 3–6 pairs gradually reduced nearly to rhizomes, pinnae similarly shaped but smaller distally, basiscopic sides cuneate and straight, acroscopic bases truncate, rounded at apices, margins of largest pinnae subentire

100

ASPLENIUM

to erose; veins 1- or 2-forked, ca. 3 pairs per pinna, evident on both sides of blades, tips evident adaxially; indument abaxially of scattered tan appressed clavate hairs 0.1–0.2 mm long on blades; sori 1 or 2 pairs per pinna, on both sides of midveins, sori on basiscopic sides Ⳳ parallel to lower margins; indusia ca. 1–1.5 ⫻ 0.3–0.5 mm, margins erose to entire; spores reniform; 2n⫽144 (Jam).

Distribution. Epiphytic on tree bases (Chis) or in rock crevices in cloud forests; 2800 m. Mexico; Costa Rica (Hammel 19261, MO, UC); Jam. Specimens Examined. Chis (Breedlove 31750, DS). ?Hgo (Sa´nchez Mejorada R. 755, MEXU).

The sole specimen from Hidalgo, from Plan del Ayacahuite, El Chico, “creciendo en las grietas de las pen˜as hu´medas,” is scrappy and without blade apices, so the identity is somewhat uncertain. If it is not this species, then it is possibly undescribed. Asplenium harrisii differs from other very small species in the A. trichomanes group, like A. soleirolioides, in having greenish or brownish, dull stipes and rachises. It is one of only a few other species of Asplenium in Mexico that have flagelliform apices, the others being A. soleirolioides, A. palmeri, and species in the A. radicans group. None of these appears to be closely related.

34. ASPLENIUM HETEROCHROUM Asplenium heterochroum Kunze, Linnaea 9: 67. 1834. Type. Cuba. Poeppig s.n. (LZ destroyed; isotypes BM!, NY!). Fig. 39E. Roots thin to filamentous, wiry, not proliferous; rhizomes erect; rhizome scales bicolorous, center strongly clathrate, margins lighter, lumina not occluded, ca. 1.5 ⫻ 0.3–0.5 mm, entire; fronds numerous, clumped, (5–)8–15(–35) cm long; stipes black, lustrous, 1–4(–7) cm ⫻ 0.3–1 mm, 1⁄10–1⁄5 of frond length, with hair-like scales ca. 2 ⫻ 0.1–0.2 mm long, adaxially with tan wings 0.1–0.2 mm wide; blades thin-chartaceous, linear, 1-pinnate, 7– 12 cm ⫻ 7–18 mm, slightly narrowed proximally (sometimes nearly to the rhizomes), tapering at tips to confluent, sometimes ligulate terminal segments; rachises black, lustrous, glabrous, with tan wings ca. 0.1 mm wide on adaxial shoulders; pinnae

oblong, 15–25(–45) pairs, 3–6 ⫻ 1.5–3(–4) mm, stalked to 0.2 mm, bases inequilateral, cuneate basiscopically, apices rounded to serrulate, acroscopic margins serrulate, sinuate-crenate, or subentire; veins simple to 1-forked, vein tips obscure adaxially; indument abaxially of whitish appressed clavate hairs 0.1 mm long; sori mostly 2–5 pairs per pinna, on both sides of midveins; indusia 0.5–1 ⫻ 0.3–0.5 mm, margins erose; spores reniform; 2n⫽144 (Fla).

Distribution. Epipetric on limestone rocks, or terrestrial in calcareous soils; 200–1400 m. USA (Fla); Mexico; Bel; Berm, Cuba, Hisp, PR; Col, Ven, Ec. Also reported from Guatemala by Stolze (1981), but the species is more broadly circumscribed in that work, with the inclusion of A. palmeri, regarded here as distinct; Adams (in Davidse et al., 1995), using the same circumscription adopted here, also cited a collection from Guatemala. Specimens Examined. Chis (Breedlove 22379B, DS, 32911, DS, MEXU, 33216, DS; Sohns 1713, MEXU, US). Ver (Copeland herb. 67, BM).

Asplenium heterochroum is distinguishable from all the lustrous- and dark-stiped, 1-pinnate Aspleniums (except A. nesioticum, A. resiliens, and A. palmeri) by the lack of conspicuous hydathodes adaxially on the blades. The closest relative in Mexico, and perhaps elsewhere, is A. nesioticum, from which A. heterochroum differs by the thinner blade texture (veins readily visible by transmitted light) and more sharply crenate-dentate pinna margins (Maxon, 1913). The smaller, thinner, more dentate pinnae, more delicate, arching fronds without strongly deflexed pinnae, smaller rhizome scales (2 mm), and occurrence at lower elevations, distinguish A. heterochroum from A. resiliens (which see). The occurrence at lower elevations and lack of terminal buds on the blades distinguish A. heterochroum from A. palmeri (which see).

35. ASPLENIUM HOFFMANNII Asplenium hoffmannii Hieron., Hedwigia 60: 258. 1918. Type. Costa Rica. [Alajuela:] Aguacate, Hoffmann 836 (B, photo BM!; frag. NY! as “826”). Fig. 34B.

ASPLENIUM JUGLANDIFOLIUM Asplenium membranifolium Maxon, Amer. Fern J. 24: 72. 1934. Type. Panama. Ca. 10 mi. E of Panama City, Killip 2567 (US!, frag. NY!).

Roots thin, fibrous, not proliferous; rhizomes short-creeping to erect; rhizome scales red-brown, clathrate, lumina not occluded, 1.5–2 ⫻ 0.3–0.5 mm, entire; fronds few, 2–5 per plant, 1–2 mm distant, 4–23 cm long; stipes green, dull, 3–9 cm ⫻ 0.5–1 mm, 1⁄4–1⁄3 of frond length, with rare red-brown, clathrate scales ca. 1.5 ⫻ 0.3 mm long, adaxially with green to gray-green wings ca. 0.3 mm wide; blades membranaceous, oblong-deltate, 1pinnate, 8–15 ⫻ 3–8 cm, apices pinnatifid, not proliferous; rachises greenish, dull, glabrous, with lateral gray-green wings ca. 0.3–0.5 mm wide; pinnae oblong, 4–10 pairs, 2–4 ⫻ 1–1.5 cm, sessile, proximal pair often slightly reduced and deflexed, basiscopically excavate ca. 1⁄2 the length, not auriculate, apices acute to usually obtuse, margins deeply crenate-serrate, sinuses 1–2 mm deep; veins simple to 1-forked, readily visible on both sides of blades, tips obscure; indument absent, blades essentially glabrous; sori 3–10 pairs per pinna, occasionally back-to-back on the same vein, on both sides of midveins; indusia 1–4 ⫻ 0.5–0.8 mm, margins erose; spores reniform, 64 per sporangium (Murakami & Moran, 1993).

101

Roots coarse, fibrous, not proliferous; rhizomes short-creeping; rhizome scales blackish, clathrate, walls thick, lumina very small, 1–2 ⫻ 0.5–0.8 mm, entire; fronds few, 1–2 mm apart, 35–45 cm long; stipes brownish, dull, ca. 16 cm ⫻ 1.5 mm, ca. 1⁄2 of frond length, glabrous or bases with rare scales, adaxially with green wings ca. 0.2–0.3 mm wide; blades membranaceous, deltate, not or only slightly reduced (1 or 2 pairs) at bases, apices attenuate, 2-pinnate, 18–25 ⫻ 4–8 cm, not proliferous; rachises brown, dull, glabrous, with green wings ca. 0.2–0.3 mm wide; pinnae deltate, 15–20 pairs, 1.5–6 ⫻ 1.5–5 cm, stalked to ca. 2 mm, pinnate, pinnules flabellulate, 4–8 pairs per pinna, 5–9 ⫻ 3–5 mm, basally cuneate, apically dentate; veins 1- or 2-forked, or to 4 pairs in the larger pinnules, visible on both sides of blades, tips barely evident adaxially; indument abaxially of brownish clavate hairs ca. 0.1 mm long; sori 1–5 per ultimate segment, occasionally diplazioid on basal segments, on both sides of midveins; indusia whitish, 1–2.5 ⫻ 0.3 mm, margins suberose to entire; spores reniform.

Distribution. Terrestrial in subtropical forests by streams and in pine-oak and mixed hardwood forests; 1850–2200 m. Mexico. Distribution. Epipetric, on muddy banks and rocks by streams in wet woods; 150–500 m. Mexico; Guat, Hond (as reported by Murakami & Moran, 1993), Salv, Nic, CR, Pan; Col, Ven, Trin. Selected Specimens Examined. Chis (Breedlove 29938, DS, 33893, DS, MEXU, MO, NY, 42851, DS, MO). Gro (Lorea 4917, FCME). Jal (Dı´az L. 21291, NY). Nay (McVaugh 19198, CAS, IEB, MEXU, NY). Oax (Herna´ndez G. 2021, CAS, CHAPA, MEXU, MO, NY; Mickel 7411, NY, UC). QR (Torres et al. 75, MEXU). Ver (Dorantes 2722, XAL).

Asplenium hoffmannii differs from A. laetum in having fewer leaves per plant, membranaceous blades, fewer pinna pairs, and more incised pinnae. It is said to hybridize with that species to produce A. incisoserratum (see A. laetum for discussion).

36. ASPLENIUM INSOLITUM Asplenium insolitum A. R. Sm., Amer. Fern J. 70: 15, f. 1, 2. 1980. Type. Mexico. Chiapas: Mpio. Motozintla de Mendoza, Cerro Mendoza, Breedlove 41653 (DS!). Fig. 37A.

Specimens Examined. Chis (Breedlove 25760, DS). Gro (Vela´zquez 138, 853, FCME). Oax (Mickel 3810a, NY).

This is similar to Asplenium breedlovei and differs in having broader, less divided segments (blades only 2-pinnate to 2pinnate-pinnatifid vs. 3–4-pinnate). There is also similarity to A. venturae, which differs in having erect rhizomes. The tapered, non-flagelliform, nonproliferous blade apices and dull brown stipes distinguish it from A. flabellulatum.

37. ASPLENIUM JUGLANDIFOLIUM Asplenium juglandifolium Lam., Encycl. 2: 307. 1786. Syntypes. “Antilles,” collector not stated (P-JU 1283A! and P-JU 1283B; photos UC!, US; isotype P-LA, photo US). Fig. 44C. Asplenium falcinellum Maxon, Contr. U.S. Natl. Herb. 13: 14. 1909. Type. Guatemala. Alta Verapaz: Cubilquitz, Tu¨rckheim II.1910 (US!).

Roots coarse, fibrous, not proliferous; rhizomes short-creeping to suberect; rhizome scales conspicuous, clathrate, iridescent, lu-

102

ASPLENIUM

mina not occluded, uniseriate at tips and contorted distally, 5– 12 ⫻ 0.5–0.7 mm, entire; fronds few (often ca. 2–4), clustered, 35–90 cm; stipes brown or purplish brown, dull, 12–25 cm ⫻ 2–4 mm, 1⁄3–2⁄3 the frond length, with rare whitish to light tan clavate hairs, not winged; blades thick-herbaceous, yellowish green, deltate-ovate, 1-pinnate, 30–55 ⫻ 18–30 cm, each with a Ⳳ conform terminal pinnae; rachises tan to stramineous or greenish, dull to sublustrous, glabrous, flattened or angled, but not winged; pinnae lanceolate to falcate, 2–8(–12) pairs, 10–17 ⫻ 1.5–3 cm, stalked to 4 mm, bases inequilateral, broadly to sometimes narrowly cuneate at bases, sometimes inequilaterally so (acroscopic side broader and more broadly cuneate, at least on distal pinnae), attenuate at apices, margins slightly sinuate to entire; veins mostly 2(–3)-forked, evident on both sides of blades, tips conspicuous adaxially; indument abaxially of scattered whitish to tan, appressed, clavate hairs 0.1–0.3 mm long; sori 7–14 pairs per pinna, on both sides of midveins; indusia 8–25 ⫻ 0.8–1 mm, margins entire; spores reniform; 2n⫽288 (Trin).

1985: 371). Schkuhr, 24. Kl. Linn. Pfl.-Syst. 1: 65, t. 70. 1805. The holotype (Swartz s.n., West Indies) has not been found. Fig. 34A. Asplenium lugubre Liebm., Mexic. Bregn. 243 (reprint 91)]. 1849. Lectotype (chosen by Smith, 1981: 47). Mexico. Veracruz: Barranca de Mirador, Liebmann s.n. [Pl. Mex. 2272, Fl. Mex. 322] (C!; isolectotypes K!, US!).

Roots fibrous, wiry, not proliferous; rhizomes short-creeping; rhizome scales clathrate, lumina not occluded, 1–2.5 ⫻ 0.1–0.3 mm, entire; fronds few, 1–2 mm apart, 15–40 cm long; stipes dark brown, lustrous, 3–16 cm ⫻ 1–2 mm, 1⁄5–2⁄5 of frond length, with filiform scales ca. 3 ⫻ 0.1–0.2 mm at bases, adaxial wings, if any, ⭐ 0.1 mm; blades chartaceous, oblong-lanceolate, 1-pinnate, 10– 25 ⫻ 3–9 cm, proximal 1–3 pinna pairs slightly reduced, apices pinnatifid, not proliferous; rachises brown, somewhat lustrous, glabrous, adaxially with greenish wings ⭐ 0.1 mm wide; pinnae oblong, 12–22 pairs, 2–6 cm ⫻ 4–12 mm, stalked 0.5–3 mm, bases inequilateral, strongly excavate basiscopically, occasionally slightly auriculate acroscopically, apices obtuse to usually acute, margins serrate to biserrate; veins 1- or 2-forked, visible on both sides of blades, tips evident adaxially; indument abaxially of whitish appressed hairs 0.1 mm long; sori (2–)4–7(–9) pairs per pinna, occasionally diplazioid, on both sides of costae; indusia 4–10 ⫻ 0.3–0.8 mm, margins erose; spores reniform, 64 per sporangium (Murakami & Moran, 1993); 2n⫽72 (Jam).

Distribution. Epiphytic in lower montane and montane rain forests; 500–950 m. Mexico; Guat, Bel, Nic, CR, Pan; Gr Ant; Col, Ven, Trin, Ec, Peru, Braz, Bol. Also cited from Honduras by Adams (in Davidse et al., 1995). Specimens Examined. Chis (Breedlove 32947, DS, MEXU, 48327, CAS, ENCB, 48360, CAS; Martı´nez S. 17617, XAL; Purpus 7249, BM, MO, NY, UC, US; Rovirosa 1044, PH).

Tryon and Stolze (1993) synonymized A. falcinellum and gave good reasons for doing so; Adams (in Davidse et al., 1995) followed suit. The reputed differences between the two species do not hold upon examination of specimens from throughout the range. Breedlove 33375 (DS), from Chiapas, at 350 m, is close to A. juglandifolium, but the fronds are very small (ca. 20 cm long) with only 2–4 pairs of lateral pinnae, and the pinnae are shallowly crenate. It could be a hybrid or merely a small variant of A. juglandifolium.

38. ASPLENIUM LAETUM Asplenium laetum Sw., Syn. Fil. 79, 271. 1806, non Schkuhr, 1809, nec Wall., 1828, nom. nud. Neotype (designated by Proctor,

Distribution. Terrestrial or on wet rocks near streams in wet forests; 100–1000(–1600) m. Mexico; Guat, Bel, Hond, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Fr Gui, Ec, Peru, Braz, Bol, nw Arg. Also cited from El Salvador and Paraguay by Adams (in Davidse et al., 1995). Selected Specimens Examined. Chis (Breedlove 22506, DS, 55423, CAS; Purpus 7241, UC, US). Oax (Herna´ndez G. 1008, CHAPA, MO, 1787, NY; Mickel 7188, UC). Tab (Cowan & Magan˜a 3139, CAS, ENCB, MEXU, MO, NY; Gilly & Herna´ndez X. 278, US). Ver (Dorantes 2555, UC; Purpus 2846, UC). Unverified, Doubtful, or Mistaken Reports. SLP (reported by Mickel & Beitel, 1988, but not verified).

Asplenium laetum differs from A. harpeodes in its creeping rhizomes, much smaller rhizome scales (1.5–2.5 mm long), fewer pinnae (10–19 pairs), pinna apices acute to obtuse, terrestrial or epipetric habit, and occurrence at lower elevations. Herna´ndez

ASPLENIUM MINIMUM

103

G. 1008 is very atypical in the prominently auriculate pinnae (auricles to 6 mm long), biserrate pinna margins, and broad pinnae 2 cm wide, including auricles (pinnae to 1.4 cm wide, excluding auricles). Hallberg 1467 (NY), from Oaxaca, was cited as the hybrid between A. laetum and A. hoffmannii by Murakami and Moran (1993), but this specimen seems reasonably typical A. laetum to us, especially in the atropurpureous, lustrous stipes and rachises, pinna shape and cutting, and blade texture. The spores, said by Murakami and Moran to be abortive, appear mostly well formed to us. The putative hybrid between A. hoffmannii and A. laetum, called A. incisoserratum (Rosenst.) Murakami & R. C. Moran, was said to occur in southern Mexico (only the above-mentioned specimen cited), Costa Rica, Trinidad, and Colombia.

39. ASPLENIUM LAMPROCAULON Asplenium lamprocaulon Fe´e, Me´m. Foug. 5: 197. 1852. Type. Mexico. Oaxaca: Llano Verde, Galeotti 6340 (P!, photos BM, BR!, MO!, UC!, US!; isotype RB). Fig. 37G. Asplenium nigricans D. C. Eaton, Proc. Amer. Acad. Arts 8: 619. 1873, hom. illeg., non Kunze, 1834. Asplenium melanorachis C. Chr., Index Filic. 121. 1905. Type. Mexico. Chiapas: Ghiesbreght 377 (YU!, photos BM!, US!, frag. US!; isotypes K!).

Roots thin, wiry, fibrous, not proliferous; rhizomes shortcreeping to suberect; rhizome scales bicolorous, central portion black, margins lighter, clathrate, 2–3 ⫻ 0.8–1 mm, entire; fronds clumped, relatively few (3–5 per plant), 30–50 cm long; stipes castaneous, lustrous, 10–12 cm ⫻ 1–1.5 mm, 1⁄5–2⁄5 of frond length, glabrous, adaxially with narrow light brown wings 0.1– 0.2 mm; blades thick-herbaceous, 1-pinnate, pinnatifid at apices, not radicant or proliferous, 17–26 ⫻ 5–7 cm, the lowest pinnae slightly reduced and deflexed; rachises atropurpureous to black, lustrous, glabrous, adaxially with light brown wings 0.3 mm wide; pinnae trapezoidal, 12–25 pairs, 2–3(–4) ⫻ 0.7–1.5 cm, sessile, inequilateral, bases slightly auriculate, apices rounded to obtuse, margins crenulate; veins 1-forked, obscure, tips evident adaxially; indument abaxially of appressed clavate hairs ca. 0.1 mm long; sori to 4 on basiscopic sides, 0–3 on acroscopic sides; indusia 3–7 ⫻ 1 mm, margins subentire; spores not seen.

Distribution. Terrestrial or epiphytic in montane rain forests; 1500–2150 m. Mexico; CR. Specimens Examined. Chis (Breedlove 21797, DS, ENCB, 31441, DS, MEXU, NY, 32757, 34451, DS, 66277, CAS; Ton 5393, CICY, ENCB, MEXU). Oax (Mickel 6051, NY; Wendt et al. 4909, CHAPA, NY). Unverified, Doubtful, or Mistaken Reports. Me´x (reported by Mickel & Beitel, 1988, but not verified; see comments below).

The dark stipes and relatively small number of large pinnae characterize this uncommon taxon in the A. monanthes group. Asplenium lamprocaulon is intermediate, in pinna number and size, between A. monanthes (large number of small pinnae) with A. obesum (small number of large pinnae). Fisher 51 (US) and Fisher s.n. (MO), from Amecameca was identified as this species (by Maxon and by Adams), and is probably the source of the attribution of range to Edo. Me´xico, but the specimen is too incomplete for satisfactory identification; perhaps it is A. polyphyllum.

40. ASPLENIUM MINIMUM Asplenium minimum M. Martens & Galeotti, Me´m. Foug. Mexique 55, pl. 15, f. 1. 1842. Lectotype (chosen by Mickel & Beitel, 1988: 62). Mexico. Oaxaca: Chinantla, Galeotti 6424 (BR!–photo NY!), figured in the original publication. Figs. 31C, D. Differs from A. pumilum by: rhizome scales longer (2–3 mm) and wider (0.5–0.8 mm), not as hair-like, occasionally denticulate along margins; fronds larger, with stipes often 15–20 cm, blades often 10–15 cm; stipes black abaxially to proximal pair of pinnae and color continuing along proximal part of rachises; pinnae thick, veins obscure; indument essentially absent on stipes, rachises, and veins of both blade surfaces and margins, abaxially glabrous or with scattered, brown, appressed hairs 0.1 mm long on abaxial leaf surfaces; indusia 4–8(–11) ⫻ 0.5–0.8 mm, with erose margins, lacking hairs.

104

ASPLENIUM

Distribution. Terrestrial and epipetric on open banks, shaded limestone ledges; 40–950 m. Mexico. Also cited from Guatemala, Panama, and Colombia by Adams (in Davidse et al., 1995), but no specimens have been seen from these countries.

Roots relatively coarse, fibrous, not proliferous; rhizomes suberect to erect; rhizome scales black, weakly clathrate with occluded lumina, 1.5–3 ⫻ 0.5–0.8 mm, entire or with sparse marginal cilia; fronds clumped, mostly (10–)20–50 cm long; stipes castaneous, dull, 1–10 cm ⫻ 1–2 mm, 1⁄10–1⁄5 of frond length, glabrous except for scales at bases, not winged; blades chartaceous, 1-pinnate, 25–45(–55) ⫻ 3–5(–7) cm, linear-oblong, apices pinnatifid, not proliferous; rachises castaneous, dull, glabrous, with greenish wings 0.3–0.5 mm wide; pinnae oblong-lanceolate, 16–55 pairs, reduced proximally, 1.5–3 ⫻ 0.5–1 cm, sessile or stalked to 1 mm, inequilateral, basiscopically excavate, acroscopic margins straight to curved, bases often overlapping rachises, apices acute or obtuse, margins serrate to biserrate, sinuses mostly 1–2(–4) mm deep; veins simple or 1–2-forked near the pinna bases, tips evident adaxially; indument abaxially of appressed tan clavate hairs 0.1 mm long; sori mostly 5–10 pairs per pinna, more numerous acroscopically than basiscopically; indusia 1.5–3 ⫻ 0.3– 0.6 mm, margins entire; spores reniform.

Selected Specimens Examined. Chis (Rovirosa 862, NY, PH). Mor (Rose & Painter 6945, US). Oax (Mickel 7328 p.p., NY, UC). Qro (Ferna´ndez N. 3465, ENCB; Rzedowski 48084, IEB, MEXU, XAL). SLP (Pringle 3389, DS, UC). Tab (Rovirosa 862, MEXU). Tam (Palmer 305, NY, US). Ver (Calzada 7943, MEXU, XAL; Purpus 6055, UC).

The wider rhizome scales (more than 3 cells at bases), thicker blades, lack of catenate hairs and ciliate indusial margins, and longer sori appear to correlate and distinguish Asplenium minimum from the closely related A. pumilum. More collections have been seen from Veracruz (23) than from all other states combined, where A. minimum is uncommon or rare. Several mixed collections of A. minimum and A. pumilum have been found in herbaria, from Oaxaca (Mickel 7328, NY, UC) and Veracruz (Bourgeau 2783, NY; Ventura A. 19852, XAL), and the differences between the two species are readily apparent in these mixed gatherings. Asplenium minimum is based on specimens with juvenile leaves that are precociously fertile, a condition seen in both A. minimum and A. pumilum, as well as many other species of Asplenium. The type is glabrous and appears to be conspecific with much larger specimens cited above, but further study is needed.

41. ASPLENIUM MIRADORENSE Asplenium miradorense Liebm., Mexic. Bregn. 243 (reprint 91). 1849. Lectotype (chosen by Smith, 1981: 47). Mexico. Veracruz: Near Mirador, Liebmann s.n. [Pl. Mex. 2275, Fl. Mex. 323] (C!; isotypes BM!, K). Figs. 32C, D. Asplenium bradeorum Hieron., Hedwigia 60: 217. 1918. Lectotype (chosen by Lellinger, 1977, Proc. Biol. Soc. Wash. 89: 706). Costa Rica. San Jose´: Tablazo, A. & A. C. Brade [33a], Rosenst. Fil. Costar. Exs. 11 (B; isolectotypes NY!, UC!, frag. US!). Asplenium schlechtendahlianum Hieron., Hedwigia 60: 218. 1918. Type. Mexico. Veracruz: bei Jalapa, Schiede 772 (B). Asplenium standleyi Maxon, Proc. Biol. Soc. Wash. 43: 85. 1930. Type. El Salvador. Finca Colima, Standley 20136 (US!).

Distribution. Terrestrial or epipetric; (500–)1000–2100(–2500) m. Mexico; Guat, Hond, Salv, Nic, CR, Pan. Also reported from Colombia and Brazil by Adams (in Davidse et al., 1995), but these reports may involve confusion with other related species in the group. Selected Specimens Examined. Chis (Breedlove 22504, DS, MO, 69099, CAS). Gro (Vela´zquez 22, FCME). Oax (Hallberg 1493, NY, UC; Wendt et al. 5165, NY). Pue (Sa´nchez 73, US; Sa´nchez Mejorada R. 613, MEXU, US; ). Ver (Purpus 6198, UC, US; Ventura A. 11272, ASU, 14631, ARIZ, MO).

See discussions under A. pteropus, A. barbaense, and A. potosinum for comparison with those species. Asplenium seileri, A. cruegeri, and A. pulchellum are additional relatives, albeit all smaller species. Asplenium miradorense is also sometimes confused with A. harpeodes, but the latter is epiphytic, at least in Mexico, while A. miradorense is strictly terrestrial. Asplenium miradorense lacks the long-attenuate pinna apices usually found in A. harpeodes. Elevationally, the two species appear nonoverlapping, or co-occurring only infrequently. Other closely related species are A. argentinum Hieron. (S. America), A. bangii Hieron. (Peru, Bolivia), A. boliviense Rosenst., A. brasiliense Raddi (S. America), A. claussenii Hieron. (Antilles, S. America), A. diplosceum Hieron. (Greater Antilles), A. otites Link, A. pul-

ASPLENIUM MONANTHES

chellum Raddi, and A. raddianum Gaudich. Adams (in Davidse et al., 1995) included A. standleyi as a synonym of A. potosinum, but we believe it is a form of A. miradorense. This whole group, including A. miradorense, is badly in need of monographic revision, a fact about which all students of the group are in agreement. Breedlove 31319 (DS, MEXU, NY), cited by Murakami and Moran (1993: 14) as A. laetum at NY (p.p.) and determined by Adams as A. cruegeri vel prox. aff., is neither species, but may be a hybrid between A. miradorense and A. abscissum. Most young sporangia are collapsed, and mature sporangia are seemingly without spores; rhizomes are erect (which precludes this specimen being A. laetum). Both putative parents are known from the area (Breedlove 31317, 31327, DS, A. miradorense; 31307, DS, A. abscissum), and the supposed hybrid is intermediate morphologically.

42. ASPLENIUM MODESTUM Asplenium modestum Maxon, Bull. Torrey Bot. Club 31: 657. 1904. Type. Mexico. Chihuahua: Hacienda San Miguel, Batopilas, 1885, Palmer 162 (US!). Figs. 41E, F. Roots thin, fibrous, not proliferous; rhizomes suberect; rhizome scales brown, clathrate, 2 ⫻ 0.3 mm, entire; fronds clumped, ca. 8 cm long; stipes green-stramineous, to 2.3 cm ⫻ 0.5 mm, 1⁄4–1⁄3 of frond length, bases with a few scattered scales similar to those of rhizomes, not winged; blades thick-herbaceous, oblonglanceolate, 4.5–6 ⫻ 1.5 cm, 2-pinnate proximally; rachises greenish to yellowish, glabrous, not winged; pinnae ovate-deltate to irregularly rectangular, 6–8 subopposite to alternate pairs, ca. 10 ⫻ 10 mm, short-stalked to 1 mm, proximal pinnae pinnately divided into 3 cuneate lobes, the acroscopic and basiscopic lobes often nearly or quite free, the obtriangular lobes in turn deeply and sharply cleft, more distal pinnae less deeply divided with commonly only the acroscopic lobe free; veins obscure, subflabellate-pinnate, tips evident adaxially; indument absent, blades essentially glabrous; sori 1–3 per lobe; indusia 1.5–2 ⫻ 0.5 mm, margins suberose; spores reniform.

Distribution. Epipetric on shaded, mossy rocks; ca. 500 m. Mexico. Known only from the type.

105

This taxon is remarkably unlike anything else known from Mexico, and has apparently not been recollected since 1885. This causes one to wonder if it could be a hybrid between two strongly divergent taxa, such as A. trichomanes and A. chihuahuense, both known from Chihuahua. However, SEM photographs (at US) of the spores of the type, by Viane, show that the spores are well formed. In fact, A. modestum closely resembles a European hybrid, A. dolosum Milde (Verh. Zool. Bot. Ges. 14: 165, t. 4. 1864), which is A. adiantum-nigrum L. ⫻ trichomanes L. Further study is warranted.

43. ASPLENIUM MONANTHES Asplenium monanthes L., Mant. Pl. 130. 1767. Figs. 33A, B, D, 44G. Asplenium monanthemum L., Syst. Veg., ed. 14, 933. 1784, hom. illeg. Type. South Africa. Cape of Good Hope (LINN 1250.17, photo US). Asplenium arcuatum Liebm., Mexic. Bregn. 241 (reprint 89). 1849. Type. Mexico. Veracruz: Mirador and Zacuapan, Liebmann s.n. [Pl. Mex. 2257, Fl. Mex. 296] (C!; isotype US!, probable isotype K!, frag. NY!). Asplenium obtusissimum Fe´e, Me´m. Foug. 5: 197. 1852. Type. Mexico. Galeotti 6446 p.p. (P?). Asplenium galeottii Fe´e, Me´m. Foug. 7: 50, t. 16, f. 2. 1857. Type. Mexico. Oaxaca: Galeotti 6369 (P; isotypes BR–photo BM! RB!). Asplenium leptophyllum Fe´e, Me´m. Foug. 7: 50, t. 14, f. 2. 1857. Syntypes. Colombia. Ocan˜a, Schlim 328, 479 (479 at RB; isosyntype of 479 at B; also at B is Moritz 328, which may be an isosyntype, possibly attributed mistakenly to Schlim); Mexico. Galeotti 6446 (RB; isosyntype BR!). Asplenium curvatum Fe´e, Me´m. Foug. 9: 16. 1857. Type. Mexico. Oaxaca: south of Sola, Cordillera Occidental, Galeotti 6370 p.p. (BR!).

Roots thin, wiry, not proliferous; rhizomes erect; rhizome scales bicolorous, central portion occluded, margins brown, clathrate, apices long-attenuate and uniseriate at tips, 4–6 ⫻ 0.5–0.8 mm, entire; fronds clumped, mostly (15–)25–70 cm long; stipes castaneous to atropurpureous, 2–15(–30) cm ⫻ 1–2 mm, 1⁄5–1⁄3 of frond length, glabrous, adaxially sometimes with narrow brownish wings 0.1–0.2 mm; blades 1-pinnate, linear, 15–40(–60) ⫻ 1– 3.5(–4.5) cm, apices pinnatifid, not proliferous or sometimes with buds in axils of proximal pinnae on older fronds; rachises castaneous to atropurpureous, lustrous, with sparse to scattered, twisted, hair-like scales 1–2 mm long, adaxially with brownish wings 0.1–0.3 mm wide; pinnae quadrangular to oblong, to 60 pairs, 6–22 ⫻ 2–7 mm, reduced proximally (lowest deltate, ca. 1 ⁄3 the length of the longest ones), sessile, dimidiate, basiscopic edges straight, acroscopic edge slightly auriculate at the base, crenulate with mucronate teeth or subentire, sinuses to 0.8 mm deep; veins simple or 1-forked (2-forked at acroscopic base), tips evident adaxially; indument abaxially of scattered, appressed, tan, clavate hairs 0.1–0.3 mm long; sori often 1–3 per pinna, often confined to basiscopic side and parallel to the margin, or 2–4 pairs per pinna on both sides of midveins; indusia 2–3(–5) ⫻ 1 mm, margins entire; spores reniform; 2n⫽108, apogamous (Ver, Oax).

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ASPLENIUM

Distribution. Terrestrial on rocky slopes in light pine-oak woods; (900–)1450–3600 m. S USA; Mexico; Guat, Hond, Salv, CR, Pan; Jam, Hisp; Col, Ven, Ec, Peru, Braz, Bol, Chile, n Arg; Africa; Hawaii. Also reported from Nicaragua and Uruguay by Adams (in Davidse et al., 1995). Selected Specimens Examined. BCS (Carter 2360, ARIZ, DS, MEXU, UC, US). Chih (Correll & Gentry 23242, UC, US). Chis (King 2798, NY, UC, US). Coah (Villarreal 5666, CHAPA, MEXU). Col (Goldsmith 47, UC). DF (Copeland herb. 66, UC). Dgo (Goodding 2148, UC). Gro (Mexia 9025, ARIZ, CAS, MO, UC, US). Gto (Dı´az B. 3053, ENCB, IEB). Hgo (Barnett et al. 78, MO, UC). Jal (Dı´az L. 5317, UC). Me´x (Vera Santos 2214, UC). Mich (King & Soderstrom 5220, NY, UC, US). Mor (Lyonnet 1381, MEXU, MO). Nay (Mexia 707a, UC). NL (Dorr 2598, MO, UC). Oax (Breckon & Christman 627, UC). Pue (Purpus 6433, UC). Qro (Zamudio & Dı´az B. 5287, ENCB, IEB). Sin (Breedlove 16862, CAS). SLP (Parry & Palmer 978, MO, NY). Son (Martin et al. s.n., ARIZ). Tam (Stanford et al. 2072, DS, MO, NY, UC, US). Tlax (Sanchez & Acosta P. 631, CIB). Ver (Nee 32916, UC). Zac (Rose 2726, US). Unverified, Doubtful, or Mistaken Reports. Ags (Gonza´lezAdame 605, HUAA, cited by Siqueiros-Delgado & Gonza´lez-Adame, 2004, but not verified).

Asplenium monanthes is a common and variable species nearly throughout Mexico. The prominent hydathodes adaxially on the blades distinguish it from Asplenium resiliens, A. heterochroum, and A. palmeri. Asplenium castaneum and A. polyphyllum differ in having non-occluded clathrate rhizome scales, lighter redbrown rachises, and more pairs of sori per pinna. Asplenium monanthes is most often confused with A. hallbergii (which see), which differs in having papillate rachis wings and short (1–2 mm long) occluded rhizome scales. In its typical form, perhaps representing 80% of the specimens seen, A. monanthes bears 1–3 sori on only the basiscopic sides of pinnae. In this feature, it can be easily distinguished from many of the other related taxa already mentioned. However, in southern Mexico, especially Oaxaca and Chiapas, many plants (but a small minority of the specimens) have been collected that have 2–3 sori (to 5 in Breedlove 22670, DS, MEXU, from Chiapas) on the acroscopic sides of the pinnae; Ventura A. 2546 (IEB), from Distrito Federal, Koch 7559 (MEXU), from Edo. Me´xico, and Lo´pez V. 113 (XAL), from Veracruz, are also unusual in having sori along both sides of the costae of each pinna. Some specimens from western Mexico, especially from

Sinaloa (e.g., Correll & Correll 28863, LL), also show this tendency. Pringle 11256 (MO), from Distrito Federal, has the pinna size, shape, and soriation of A. hallbergii, but lacks the papillate adaxial rachis wings of that species and the indusia are entire; thus, it will key to A. monanthes. Further study may show that these variants represent a separate species, or are apogamous variants of this or related species. One of the most extreme forms of A. monanthes is Breedlove 21707 (DS, NY), which has 3–4 pairs of sori per pinna, nearly blackish stipes and rachises, and broad pinnae 7–8 mm wide with rounded tips. This specimen appears to represent an undescribed species in the A. monanthes group, but more material is needed to be more certain of its discreteness. A few specimens of A. monanthes, e.g., Dı´az B. & Carranza 6601 (IEB), from Quere´tero, bear buds or plantlets in the axils of proximal pinnae.

44. ASPLENIUM MONODON Asplenium monodon Liebm., Mexic. Bregn. 247 (reprint 95). 1849. Figs. 36A, B. Asplenium auritum Sw. var. monodon (Liebm.) E. Fourn., Mexic. Pl. 1: 106. 1872. Lectotype (chosen by Smith, 1981: 48). Mexico. Veracruz: Hacienda de Jovo, Liebmann s.n. [Pl. Mex. 2295, Fl. Mex. 302] (C!; isotypes K!, US!). ?Asplenium auritum Sw. var. subsimplex Kunze, Linnaea 18: 332. 1844. Type. Mexico. Leibold 13 (Herb. Roemer, LZ-destroyed; isotype B–photo US!).

Similar to A. auritum except: blades 1-pinnate, each pinna with a superior auricle often cut almost to costa but with acroscopic auricle adnate at the base, margins deeply biserrate, cut to ca. 1⁄3 distance to midvein; sori (3–)6–8 pairs per pinna, on both sides of midveins; indusia 3–5 ⫻ 0.5–0.8 mm; spores globose, large, dark brown to blackish, 32 per sporangium.

Distribution. Epiphytic or on moist rocks, rarely terrestrial; 100–1600(–2300) m. USA (Fla); Mexico; Guat (Steyermark 44317, UC), Hond (Croat & Hannon 64388, UC), CR (Stork 1421, UC), Pan (Cornman 815, UC); Cuba, Jam, Hisp, Martinique; Ven, Trin, Ec. Also reported from Colombia by Adams (in Davidse et al., 1995). Selected Specimens Examined. Chis (Breedlove 22149, DS, 35387, DS, MO). Oax (Mexia 9162, CAS, GH, MO, NY, UC; Mickel 5840,

ASPLENIUM MYRIOPHYLLUM

107

NY, UC). Pue (Vera Santos 2579, UC). Ver (Bourgeau 1829, US; Copeland herb. 63, US; Orcutt 2827, DS, US; Purpus 2415, UC, 2847, MO, UC, US). Unverified, Doubtful, or Mistaken Reports. Tab (reported by Magan˜a, 1992, but not verified). See discussion under A. auritum.

Asplenium macilentum Klotzsch ex Kunze (1857) was regarded as a distinct species, also occurring in Mexico, by Adams (in Davidse et al., 1995). He excluded A. monodon from Flora Mesoamericana (which we would not) and characterized A. macilentum as having reniform spores, 64 per sporangium. The lectotype of this is of unknown provenance (selected by Morton & Lellinger, 1966: 18, B-Willd. 19890, microfiche UC!), is similar in form, and may also belong in synonymy under this species (Smith, 1981: 48); however, until spores of the lectotype can be examined, it would be imprudent to place the name. Some specimens from northern South America (especially Venezuela) with the morphology of A. macilentum have large, globose spores, like those of A. monodon, while others have small, reniform spores, like those of A. auritum sensu stricto. A plant identified as A. macilentum from Trinidad was reported as 2n⫽288, apogamous, by Walker (1985), but Adams (in Davidse et al., 1995) stated that all material in the A. auritum complex from Trinidad was referable to A. monodon. Nearly all plants identified by Adams as A. macilentum from Central America we regard as A. auritum, including those he cited (1995: 309) from Chiapas, Belize, Honduras, and Panama.

45. ASPLENIUM MUENCHII Asplenium muenchii A. R. Sm., Proc. Calif. Acad. Sci., ser. 4, 40: 211. 1975 [“munchii”]. Type. Mexico. Chiapas: San Pablo, Mu¨nch 114 (DS!). Figs. 35J, K. Asplenium rhizophyllum Kunze var. proliferum Davenp., Bot. Gaz. 19: 390. 1894. Type. Mexico. Can˜ons of Colima Mts., Pringle 5533 (GH!; isotypes GH!– 2 sheets).

Roots relatively coarse, fibrous, not proliferous; rhizomes suberect; rhizome scales black, clathrate, 3–4 ⫻ 1–1.3 mm, with occasional small teeth; fronds clumped, 25–50 cm long; stipes castaneous to reddish brown, dull, 3–9 cm ⫻ 1–2 mm, 1⁄10–1⁄4 of frond length, glabrous, with adaxial wings 0.1 mm wide; blades thin-chartaceous, (17–)25–40 ⫻ (3.5–)5–9 cm, elliptic to narrowly elliptic, apices attenuate, each blade with a proliferous bud near apex, 2-pinnate-pinnatifid to 3-pinnate; rachises graybrown, dull, glabrous, with adaxial wings 0.1–0.3 mm wide; pinnae broadly lanceolate, 20–25 pairs, to 5 ⫻ 2 cm, lowest somewhat deflexed, sessile, inequilateral, pinnules more developed acroscopically, costae winged, pinnules spreading to often somewhat ascending, ultimate segments ca. 1 mm wide, falcate, cuneate; veins forking, tips evident adaxially; indument abaxially of scattered, appressed, light tan hairs 0.1 mm long on veins and tissue between veins, blades essentially glabrous; sori one per ultimate segment; indusia whitish to tan, mostly 1–2(–3) ⫻ 0.4– 0.6 mm, margins entire; spores reniform.

Distribution. Terrestrial or epipetric; 1300–2800 m. Mexico. Selected Specimens Examined. Chis (Breedlove 6222, DS, ENCB, US, 33648, DS). Gro (Hinton 14302, ARIZ, LL, MO, NY, TEX, US; Martı´nez S. 7526, CAS, MO, XAL). Jal (Dı´az L. 6549, UC, US). Me´x (Hinton 2389, BM, K, NY, 8941, ARIZ, MO, NY, US). Mich (Rzedowski 46275, ENCB, MEXU, TEX, XAL). Mor (Lyonnet 2876, U). Oax (Mickel 3818a, NY).

Citation of Guatemala in the range of A. muenchii by Mickel and Beitel (1988: 64) is based on a misidentification of A. sessilifolium (Heyde & Lux s.n. [Donn. Sm. 4670], NY, an isosyntype of A. sessilifolium var. guatemalense). This species has often been misidentified as A. myriophyllum, which has more fusiform terminal segments and nonproliferous blades. Asplenium muenchii is more closely allied to A. sessilifolium, which is only pinnate-pinnatifid rather than 2-pinnatepinnatifid to 3-pinnate. Adams (in Davidse et al., 1995) also recognized A. cladolepton Fe´e (Me´m. Foug. 7: 55, t. 22, f. 4. 1857; type from Colombia, Schlim 324) as occurring in Chiapas and southern Mexico. Specimens of A. cladolepton from Colombia and Venezuela differ from A. muenchii in having narrower blades and lacking buds or plantlets in the axils of distal pinnae, and the resemblance seems to us the result of convergence. However, two of the three Chiapas specimens cited here (including Breedlove 6222, DS, the specimen cited as A. cladolepton by Adams) lack buds (all other specimens cited above bear rachis buds). For now, we maintain the name A. muenchii for Mexican specimens, and exclude A. cladolepton from the Mexican flora.

46. ASPLENIUM MYRIOPHYLLUM Asplenium myriophyllum (Sw.) C. Presl, Rel. Haenk. 1: 48. 1825. Figs. 38D, 40J, K. Caenopteris myriophylla Sw., J. Bot. (Schrader) 1800(2): 60. 1801. Type. Jamaica. Swartz s.n. (S–photo US!; isotype B-Willd. 19859, photo NY, microfiche UC!). Asplenium monteverdense Hook., Sec. Cent. Ferns, pl. 41. 1860. Type. Cuba. Oriente: Monte Verde, Wright 1029 (K!; isotypes B!, GH, NY!, US!). Asplenium verecundum Chapm. ex Underw., Bull. Torrey Bot. Club 33: 193. 1906. Type. U.S.A. Florida: Jackson Co., Chapman s.n. (NY!).

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ASPLENIUM

Roots thin, fibrous, proliferous; rhizomes erect; rhizome scales blackish, clathrate, but lumina small, occluded, 0.8–1.5 ⫻ 0.5– 0.7 mm, entire; fronds clumped, 6–30 cm long; stipes reddish brown to gray-green, dull, 1–12 cm ⫻ 0.5–1 mm, ca. 1⁄10(–1⁄3) of frond length, glabrous, wings seemingly absent; blades membranaceous, elliptical, 5–25 ⫻ 2.5–8 cm, 3-pinnate, tapering proximally and distally, apices elongate, non-conform, not proliferous; rachises reddish brown to gray-green, dull, glabrous, adaxially with narrow green wings 0.1–0.2 mm wide; pinnae broadly lanceolate, 10–20 pairs, 1.5–6 ⫻ 1–2 cm, sessile to stalked to 2 mm, 2-pinnate, ultimate segments obovate to elliptical, ca. 1 mm wide, narrowed proximally, apices acute, non-falcate; veins 1 per ultimate segment, tips evident adaxially; indument abaxially of scattered, tan, appressed clavate hairs 0.1–0.2 mm long; sori 1 per ultimate segment; indusia 1–1.5(–2) ⫻ 0.5–0.8 mm, margins entire; spores reniform; 2n⫽144 (USA).

Distribution. Epipetric (on limestone) and terrestrial; 700– 2700 m. USA (Fla); Mexico; Guat; Berm, Gr Ant; Ven. Also cited from Honduras, Nicaragua, Costa Rica, Panama, Colombia, Ecuador, Peru, and Bolivia by Adams (in Davidse et al., 1995), but no doubt this wider range is a reflection of the broader circumscription he adopted for the species. For example, Cornman 935, cited as this species from Panama by Adams, is A. cristatum, in our opinion. We doubt that the species is this widely distributed, as we circumscribe it.

tatum: Mickel 7301B, NY, from Oaxaca. In Florida, Asplenium myriophyllum hybridizes with A. abscissum to form A. ⫻curtissii Underw. See discussion under A. cristatum and A. rutaceum for differences with those taxa.

47. ASPLENIUM NESIOTICUM Asplenium nesioticum Maxon, Contr. U.S. Natl. Herb. 17: 142, f. 4. 1913. Type. Jamaica. Ledges above Green River, trail from Cinchona to Blue Mt. Peak, Maxon 1487 (US! isotype US!). Figs. 39A, B. Roots filamentous, fibrous, not proliferous; rhizomes suberect; rhizome scales black, 1.5–2 ⫻ 0.2–0.3 mm, lumina occluded; fronds clumped, 6–15(–22) cm long; stipes blackish, 0.5–1(–4) cm ⫻ 0.5–1 mm, less than 1⁄10 of frond length, glabrous, adaxially with whitish wings 0.1 mm wide; blades thick-chartaceous, 5– 15(–20) ⫻ 1.5–2 cm, 1-pinnate, narrowly elliptic-linear, proximal 4–5 pairs of pinnae reduced nearly to rhizomes, dwindling slightly at tips to confluent, sometimes hastate terminal segments; rachises atropurpureous, lustrous, glabrous, adaxially with narrow whitish wings 0.1–0.2 mm wide; pinnae oblong, 15–20(– 30) pairs, 7–10 ⫻ 2–4 mm, dimidiate, sometimes with small basal acroscopic auricles, apices obtuse, serrulate to crenulate-dentate along acroscopic margins and tips; veins mostly simple, obscure on both surfaces of pinnae, tips not evident adaxially; indument abaxially of scattered, appressed to spreading, whitish hairs 0.1 mm long on veins and tissue between veins; sori (1–)2–5 pairs per pinna, on both sides of midveins; indusia whitish, 1–2 ⫻ 0.3– 0.4 mm, margins erose; spores ovoid-globose, 32 per sporangium (Purpus 4880, UC), or reniform, 64 per sporangium (see below).

Selected Specimens Examined. Chis (Breedlove 22302, 31539, 34697, DS). DF? (Valle de Me´xico, Schaffner 105, NY). Oax (Mickel 3818, MEXU, NY, UC, 5350, ENCB, NY, UC, US, 5369, ENCB, MEXU, NY, UC). Ver (Bourgeau 2002, MO, 3059, MO, US; Pringle 6094, LL, NY, UC, US).

This species was treated as Asplenium divaricatum Kunze (Linnaea 9: 71. 1834, type from Peru, Cerro de San Cristo´bal, prope Cassapi, Poeppig [1151], isotypes B!, NY!, P!, W) by Mickel and Beitel (1988), but Smith (1981) and Tryon and Stolze (1993), restricted the application of A. divaricatum to plants from Peru. Adams (in Davidse et al., 1995) circumscribed A. divaricatum more broadly, to include plants from Guatemala, Trinidad, Brazil, and Chile, but he did not admit Mexico in the range. Putative hybrids, intermediate in morphology between the suspected parents, occur between A. myriophyllum and A. cris-

Distribution. Presumably on rocks (pedregal) and ledges; 1350– 2100 m. Mexico; Jam, Hisp. Specimens Examined. DF (Schaffner 70, 109, NY). Gto (Correll & Correll 28819, LL p.p.). NL (Webster & Preston 2885, TEX). SLP ( Purpus 4880, UC, US, 5487, UC).

Specimens of this species from Mexico have previously been determined as A. heterochroum, A. palmeri, and A. aff. resiliens,

ASPLENIUM OLIVACEUM

all of which are closely related. Asplenium nesioticum differs from A. heterochroum in having thicker blade texture (veins not visible), shorter stipes, and less sharply crenate-dentate pinna margins. From A. resiliens, A. nesioticum differs in having inframedial to medial sori, thinner blade texture, and acuminate (vs. hairlike) scale tips. From A. palmeri, A. nesioticum differs in lacking proliferous blade tips. Webster & Preston 2885, from Nuevo Leon, was annotated by Wendt as Asplenium aff. resiliens, “but sexual, 64 spores/sporangium.” We confirm this number, and suggest that this species may be one of the sexual progenitors of the apogamous A. resiliens, 2n⫽108 with 32 large spores per sporangium. Further work is needed on this difficult complex of anhydathodous spleenworts of the A. trichomanes group.

48. ASPLENIUM OBESUM Asplenium obesum Baker in Hooker & Baker, Syn. Fil. 209. 1867. Type. Guatemala. Volca´n de Fuego, Salvin & Godman 119 (K!, drawing NY!). Fig. 44D. Asplenium kellermanii Maxon, Contr. U.S. Natl. Herb. 17: 152. 1913. Type. Guatemala. Solola´: Volca´n Atitla´n, Kellerman 5792 (US!). Asplenium pinnatum Copel., Univ. Calif. Publ. Bot. 19: 289, t. 40. 1941. Type. Mexico. Guerrero: Dist. Mina, Sierra Madre del Sur, Petlacala W of Reyes, Mexia 9048 (UC!; isotype US!).

109

49. ASPLENIUM OLIGOSORUM Asplenium oligosorum Mickel & Beitel, Mem. New York Bot. Gard. 46: 64. 1988. Type. Mexico. Oaxaca: Distrito Teotitla´n, Mpio. Tenango, first half of the trail from San Martı´n Caballero to Tenango, Hallberg 1444 (NY!; isotype UC!). Fig. 33H. Roots filiform, fibrous, not proliferous; rhizomes shortcreeping to suberect; rhizome scales black, clathrate, 1–1.5 ⫻ 0.1– 0.3 mm, entire; fronds clumped, 5–16 cm long; stipes atropurpureous, lustrous, (1–)2.5–3.5 cm ⫻ 0.5–0.6 mm, 1⁄3–2⁄5 of frond length, glabrous, adaxially with very narrow brown wings; blades 1-pinnate, 4–13 ⫻ 1.3–1.5 cm, apices pinnatifid, not proliferous; rachises atropurpureous, lustrous, glabrous, adaxially with brown, wings 0.1–0.3 mm wide; pinnae oblong, arcuate, 12–18 pairs, (5–)7–8 ⫻ 3–5 mm, opposite proximally, alternate distally, sessile, dimidiate, articulate, rachis color not extending onto pinna stalks, acroscopic bases of pinnae usually overlapping rachises, apices rounded to obtuse, entire to undulate-serrate; veins simple or 1-forked near their bases, tips evident adaxially; indument absent, except for scattered, appressed hairs 0.1 mm long on laminae abaxially; sori 1 or 2 per pinna, on one or both sides of midveins, if only one then this parallel to and adjoining the basiscopic margin; indusia whitish, 2–4 ⫻ 0.8–1 mm, margins undulate; spores reniform.

Differs from Asplenium lamprocaulon by: blades each with a larger, subconform, terminal pinna; pinnae larger, 4–5 ⫻ 2–2.5 cm, fewer per blade (4–9 pairs); sori more numerous, 4–5 pairs per pinna on both sides of midveins.

Distribution. Epipetric on limestone in primary forests; 1300– 1750 m. Known only from the type collection and one additional gathering from the same area. Specimen examined. Oax (Geith 95-M023, NY).

Distribution. Terrestrial in wet forests; 1800 m. Mexico; Guat. Known only from the three type collections, and in Mexico only from Guerrero; reported from Costa Rica by Lellinger (ms.) Asplenium obesum agrees with A. lamprocaulon in having dark, lustrous stipes and rachises, but differs by the characters mentioned above. Its distinctiveness and rarity are reinforced by the fact that it has been described three times, the only collections known for the species.

Asplenium oligosorum is closely related to A. olivaceum and differs from that in the less rounded (more arcuate) pinnae that usually overlap the rachises, more coarsely undulate pinna margins, prominent brownish wings on the stipes and rachises adaxially, and larger indusia.

50. ASPLENIUM OLIVACEUM Asplenium olivaceum A. R. Sm., Proc. Calif. Acad. Sci., ser. 4, 40: 212. 1975. Type. Guatemala. Alta Verapaz: Senahu, summit of Cerro Sillab, Hatch & Wilson 149 (US!). Figs. 42A, B.

110

ASPLENIUM

Roots thin, fibrous, not proliferous; rhizomes erect; rhizome scales black, clathrate, 2–3 ⫻ 0.5–0.8 mm, entire; fronds clumped, to ca. 15 cm long; stipes atropurpureous, lustrous, 1.5–2 cm ⫻ 0.6 mm, 1⁄5–1⁄3 of frond length, with scattered, hair-like scales 0.8– 1 mm long, not winged; blades 1-pinnate, 5–15 ⫻ 0.8–1.5 cm, apices pinnatifid, not proliferous; rachises reddish brown to atropurpureous, lustrous, with rare to sparse clavate hairs, adaxially with minute wings; pinnae oblong, 6–15 pairs, opposite, 4–8 ⫻ 3.5–5 mm, dimidiate, pinnae articulate, rachis color not extending onto pinna stalks, apices obtuse, margins entire to slightly undulate acroscopically; veins simple or 1-forked at the acroscopic base, tips evident adaxially; indument abaxially with sparse, appressed, clavate hairs; sori 1 or 2 per pinna, largest sorus of a given pinna parallel to the basiscopic pinna margins, sori on acroscopic side of pinnae oblique with respect to midveins, or 2–3 pairs per pinna on both sides of midveins; indusia whitish to gray, (1–)2–3 ⫻ 0.8–1 mm, margins minutely Ⳳ entire to slightly erose; spores reniform.

51. ASPLENIUM PALMERI Asplenium palmeri Maxon, Contr. U.S. Natl. Herb. 13: 39. 1909. Type. Mexico. Jalisco: Shaded mountain near Etzatla´n, Rose & Painter 7582 (US!). Fig. 33G. Asplenium trichomanes L. var. repens Davenp., Bull. Torrey Bot. Club 13: 130. 1886. Type. Mexico. Near Chihuahua, Pringle 444 (GH!; isotypes MO!–2 sheets, NY!–3 sheets, UC!, US!).

Roots thin, fibrous, not proliferous; rhizomes erect; rhizome scales bicolorous, center black, occluded, margins narrow, brown, clathrate, 2–5 ⫻ 0.2–0.5 mm, entire; fronds clumped, 5–30 cm long; stipes black to castaneous, lustrous, 0.5–2 cm ⫻ 0.2–0.7 mm, 1⁄10 or less of frond length, glabrous, adaxially with whitish to tan wings 0.1 mm wide; blades chartaceous to subcoriaceous, 5–15(–20) cm long, 1-pinnate, linear-lanceolate, reduced proximally nearly to the rhizomes, apices pinnatifid or with rachises often flagelliform and producing a bud (ultimately plantlet) at the tips; rachises atropurpureous, lustrous, with scattered hairlike clathrate scales 1–1.5 mm long, adaxially with tan wings 0.1– 0.2 mm wide; pinnae oblong, (15–)20–40 pairs, not deflexed, 3–8 ⫻ 1.5–3 mm, articulate, rachis color not extending onto pinna stalks, scarcely occasionally with a tiny auricle acroscopically, apices obtuse, crenate-serrate to serrate with sinuses 0.3–0.5 mm deep; veins simple, tips not evident adaxially; indument abaxially of scattered, appressed, whitish to tan clavate hairs 0.1 mm long; sori 2–7 pairs per pinna, on both sides of midveins; indusia 0.8–1 ⫻ 0.3–0.5 mm, margins entire; spores globose to ovoid.

Distribution. Limestone cliffs; 1300–2900 m. Mexico; Guat. Specimens Examined. Chis (Breedlove 22379A, DS, ENCB; Rzedowski 669, ENCB). Oax (Mickel 7059c, NY). Unverified, Doubtful, or Mistaken Reports. Me´x (Flores X. 45, IZTA, cited by Tejero-Dı´ez & Arreguı´n-Sa´nchez, 2004, but the specimen is A. castaneum).

Among the small-bladed taxa with lustrous, castaneous stipes and rachises, A. olivaceum has prominent hydathodes (unlike A. resiliens and A. palmeri), is nonproliferous (unlike A. palmeri and A. stolonipes), has minute wings (unlike A. resiliens, A. palmeri, and A. oligosorum, which have wings 0.1–0.3 mm wide), nonoccluded clathrate rhizome scales (unlike A. resiliens, A. palmeri, and A. stolonipes), and small (1–3 mm long) indusia (unlike A. oligosorum). Breedlove 31818B (DS), from Chiapas, is close to A. olivaceum but differs in the more oblong, narrower segments. Possibly it is undescribed.

Distribution. Epipetric on steep hills; 800–2000(–2750) m. USA (Ariz, N Mex, Tex); Mexico; Guat, Bel, Hond; Bol (Kessler 9569, UC). Also reported from Colombia, Venezuela, and Ecuador by Adams (in Davidse et al., 1995), but collections from this area are probably A. heterochroum s.str., with which A. palmeri is sometimes synonymized (e.g., by Stolze, 1986). Selected Specimens Examined. Ags (McVaugh & Koelz 127, IEB, MEXU, NY). Chih (Correll & Gentry 22800, MO, NY, UC, US). Coah (Johnston 8272, MEXU, MO). DF (Lyonnet 1385 p.p., MEXU, US). Dgo (Palmer 555, MEXU, MO, NY, UC, US). Gro (Carbajal A. 7, FCME). Gto (Rzedowski 40578, ENCB, IEB). Hgo (Chase 7436, MO, US). Jal (Pringle 1838, DS, MO, UC). Me´x (Magallo´n B. 91, ENCB). Mich (Rzedowski 39267, IEB). Nay (Breedlove 45373, CAS). NL (Correll & Johnston 19923, NY). Oax (Mickel 4945, ENCB, NY, UC, US). Pue

ASPLENIUM POTOSINUM

111

(Purpus 3152, UC). Qro (Carranza 3653, IEB, UC). Sin (Breedlove 17907, CAS). Son (Sanders 13047, NY). Tam (Stanford et al. 2620, NY). Ver (Purpus 6619, MO, UC, US). Zac (McVaugh 17761, MICH).

Asplenium palmeri is distinguishable from related dark-stiped aspleniums (except A. heterochroum, A. nesioticum, and A. resiliens) by the lack of conspicuous vein tips adaxially. It differs from all three in many features, the most conspicuous being the proliferous tips present on many blades and its different habit. From A. resiliens, A. palmeri differs by having smaller, thinner-textured pinnae with distinctly serrulate margins and delicate, arching blades with opposite spreading pinnae. From the more northerly A. trichomanes, A. palmeri differs in having generally larger arching fronds, serrulate pinna margins, and flagelliform blade tips. Fisher 35382 (NY, US), cited as this species by Palacios-Rios (1992), appears to be A. resiliens.

52. ASPLENIUM POLYPHYLLUM Asplenium polyphyllum Bertol., Novi Comment. Acad. Sci. Inst. Bononiensis 4: 443. 1840, non C. Presl, 1836, nom. nud., nec C. Presl ex Goldman, 1843. Figs. 33E, F. Asplenium polymeris T. Moore, Index Fil. 154. 1859, nom. superfl. Asplenium bertolonii Donn. Sm., Enum. Pl. Guatem. 4: 189. 1895, nom. superfl. Type. Guatemala. Volca´n Agua, Vellasquez s.n. (BOLO). See R. E. G. Pichi Sermolli & M. P. Bizzarri. 1990–91. Le collezioni pteridologiche conservate nell’erbario di Antonio Bertoloni. Mem. Accad. Lunigianese Sci. “Giovanni Capellini.” 60– 61: 177–232, t. 44. Asplenium trichomanes L. var. viridissimum Christ, Bull. Soc. Roy. Bot. Belgique 35(1): 195. 1896. Asplenium viridissimum J. Bommer ex Christ, Bull. Soc. Roy. Bot. Belgique 35(1): 195. 1896, nom. nud. in syn. Type. Costa Rica. Volca´n Barba, 1890, Tonduz s.n. (Pittier 1937) (BR–photos BM!; isotypes CR, US!, Z).

Roots thin, fibrous, not proliferous; rhizomes suberect; rhizome scales black, clathrate, tips long-attenuate, 4–6 ⫻ 0.8–1 mm, entire; fronds clumped, (20–)30–60(–100⫹) cm long; stipes reddish brown to light brown, lustrous, 1–10(–25) cm ⫻ 1–2 mm, 1⁄20– 1 ⁄4 of frond length, with scattered, hair-like, clathrate scales 1–2 mm at bases, adaxially with tan wings 0.1 mm wide; blades 1pinnate, linear, (9–)20–50 ⫻ 1.5–2.5 cm, apices pinnatifid, proliferous buds subterminal (several pinnae proximal to blade apices); rachises castaneous, lustrous, with rare to sparse clavate hairs 0.3 mm wide, adaxially with brownish wings 0.1–0.3 mm wide; pinnae oblong, 50–80 pairs, 8–10(–22) ⫻ 4–6 mm, articulate, with rachis color extending onto pinna stalks, apices obtuse, margins crenate; veins simple, tips evident adaxially; indument abaxially of scattered tan, appressed, clavate hairs 0.1–0.3 mm long; sori (1–)2–5 pairs per pinna, on both sides of midveins; indusia whitish, 1.5–2.5 ⫻ 1 mm, margins subentire; spores reniform.

Distribution. Terrestrial in cloud forests; (2000–)2450–3050 m. Mexico; Guat, Hond, Salv, CR, Pan; Ven, Ec, Bol. Also reported from Colombia and Peru by Adams (in Davidse et al., 1995). Selected Specimens Examined. Chis (Breedlove 32052, DS; Croat 47297, MO). DF (Lyonnet 905, ENCB, MEXU, MO). Gro (Hinton 11118, NY; Martı´nez S. 884, IEB, MO). Hgo (Sa´nchez M. 701, MEXU). Me´x (Sa´nchez S. 427, US). Mor (Lyonnet 1410, CAS, ENCB, MEXU). Oax (Mickel 4600, 4917, NY, UC, 5270, ENCB, NY, UC, 7061, NY, US).

Clathrate, non-occluded rhizome scales, buds in the axils of distal pinnae, sori regularly on both sides of the midveins, and broader indusia distinguish A. polyphyllum from A. monanthes. Asplenium polyphyllum is also similar to A. castaneum except the latter has more-rounded pinnae, erose indusia, and lacks proliferous rachial buds. Sa´nchez M. 701, from Hidalgo, resembles large forms of A. castaneum in size and shape of the pinnae, but has proliferous rachises. This species was cited for Michoaca´n by Dı´az-Barriga and Palacios-Rios (1992), but the specimen (Dı´az B. & Zamudio 2668, ENCB, IEB) appears to be A. castaneum. This collection number was cited under both species by the authors.

53. ASPLENIUM POTOSINUM Asplenium potosinum Hieron., Hedwigia 60: 247. 1918. Type. Mexico. San Luis Potosı´: Baranca de las Canoas, Pringle 3920 (B; isotypes ARIZ!, DS!, MO!, NY!–3 sheets, US!). Figs. 35F–H. Asplenium regulare Sw. var. latior E. Fourn. ex Hieron., Hedwigia 61: 10. 1919. Asplenium lunulatum Sw. var. latius E. Fourn., nom. nud., Mexic. Pl. 1: 105. 1872. Type. Mexico. Veracruz: Izhuatlancillo, pre`s Orizaba, Bourgeau 2369 (B; isotypes MO, NY!, US!). Asplenium hesperium Mickel & Beitel, Mem. New York Bot. Gard. 46: 59. 1988. Type. Mexico. Oaxaca: Distrito Putla, 3 km N of Putla, Mickel 3973 (NY!; isotypes ENCB!, UC!).

Differs from A. miradorense by: rhizome scales shorter, 2–3.5 mm long, margins toothed; stipes gray-green to brownish gray, dull, narrowly alate (0.1–0.3 mm) to bases; blades each with a proliferous bud at apex; rachises brownish, dull, with green wings 0.5 mm wide, with tan appressed hairs 0.1–0.2 mm; pinnae reduced and sometimes sharply deflexed proximally, 15–25 mm

112

ASPLENIUM

long, median pinnae somewhat falcate, larger, 22–40 mm long, basiscopically excavate in the basal 1⁄3, with mostly simple serrations with sinuses less than 1(–3) mm deep; veins simple or 1forked; sori in larger pinnae (6–)8–10 pairs per pinna, on both veins of a fork at bases of larger pinnae.

Distribution. Terrestrial in montane rain forests; 550–2200. Mexico; Guat. Also cited from El Salvador and Nicaragua by Adams (in Davidse et al., 1995), but his circumscription of A. potosinum included A. standleyi, which is placed under A. miradorense in our treatment. Selected Specimens Examined. Chis (Breedlove 42652, DS, MEXU; Croat 47618, LL, MEXU, MO). Col (Sanders 8548, UC; Va´zquez 413, UC). Gro (Hinton 14705, ARIZ, MO, NY, US). Jal (Wilbur & Wilbur 1464, US). Nay (McVaugh 18984, CAS, IEB, MEXU, NY). Oax (Wendt et al. 4708, CAS, CHAPA, MEXU, MO). SLP (Pringle 3413, NY, UC). Sin (Gentry 5666, ARIZ, MO, NY, US). Ver (Bourgeau 2369, MO, NY, US).

This species is most closely related to A. miradorense, differing in having more evenly serrulate acroscopic pinna margins, more numerous sori, and small-toothed rhizome scales. Some specimens, especially in western Mexico, also differ by the presence of small buds or plantlets arising from axils of subterminal pinnae. This is especially well developed on Hinton 14705 (US), with plantlets 6⫹ cm long. However, buds are absent on most specimens, including all those from Colima, San Luis Potosı´, and Veracruz. Rzedowski & McVaugh 60 (ENCB), from Guerrero, has a plantlet at the tip of one of the blades, but has pinna dissection more like A. miradorense; we tentatively place it under A. potosinum. Other close relatives are A. barbaense, A. harpeodes, and A. pteropus. Asplenium hesperium was said to differ from A. potosinum by the more strongly deflexed proximal pinnae, rounded pinna tips, proliferous rachises, and distribution in western Mexico. These differences do not seem to hold in Chiapas or in western Mexico either. Most specimens we have seen of A. hesperium have nonproliferous blades, including an isotype at UC. Both A. potosinum and A. hesperium agree in having small rhizome scales with occasional teeth, dull gray-green, slightly alate stipes, rather finely serrulate, acroscopic pinna margins, and numerous sori.

54. ASPLENIUM PRAEMORSUM Asplenium praemorsum Sw., Prodr. 130. 1788. Type. Jamaica. Swartz s.n. (S; isotype B-Willd. 19922-2, microfiche UC!). Fig. 31E. Roots coarse, fibrous, not proliferous; rhizomes suberect; rhizome scales dark brown, clathrate, 4–7 ⫻ 0.4–0.6 mm, with long, filiform, uniseriate tips, entire; fronds clumped, 10–50 cm long; stipes black to brown, dull, 10–14 cm ⫻ 1–2(–3) mm, 1⁄4–2⁄3 of frond length, with abundant black-brown lanceolate to hair-like, often twisted scales mostly 2–5 mm long, not winged; blades thick-chartaceous to subcoriaceous, ovate to lanceolate, 2pinnate to 2-pinnate-pinnatifid, 12–45 ⫻ 4–10 cm; rachises black abaxially, green adaxially, with scattered, basally fimbriate or pectinate scales 1–3 mm long, not winged; pinnae lanceolate to deltate, mostly 7–15 pairs, 2–11 ⫻ 1–3 cm, proximal ones stalked 2 mm or more, acute, pinnate, 3–7 segments (pinnules) per proximal pinna, segments obovate or obtriangular, sometimes lacerate, deeply incised, or with fine teeth at apices; veins arising at very narrow angles (5–20⬚) from the main vein of a segment, 4–8 per ultimate segment, tips not evident adaxially, but veins at least faintly visible on both blade surfaces; indument abaxially of dense to scattered narrow scales 1–3 mm long; sori mostly 1–5 per pinna lobe (pinnule); indusia entire, 3–12 ⫻ 0.3–0.5 mm; spores reniform; 2n⫽288 (Jam).

Distribution. Epiphytic or epipetric in mesic to dry pine-oak woods; 700–3000(–3400) m. Mexico; Guat, Hond, Salv, Nic, CR, Pan; Gr Ant; Col, Ven, Ec, Galapagos, Peru, s Braz, Bol, nw Arg. Selected Specimens Examined. Chis (Alava 1260, UC). Col (Va´zquez 653, UC). DF (Orcutt 4297, DS, MO, NY). Dgo (Tenorio L. 6151, MEXU). Gro (Mexia 9044a, UC). Gto (Rzedowski 51235, IEB). Hgo (Sa´nchez Mejorada R. 684, MEXU). Jal (Bueno 17102, UC). Me´x (Moore & Cetto 5461, UC, US). Mich (Pringle 3363, DS, MEXU, MO, NY, UC, US). Mor (Lyonnet 1843, CAS, US). Nay (Dı´az L. 16311, IBUG, UC). Oax (Mickel 3750, UC). Pue (Purpus s.n., UC). Qro (Carranza 1050, ENCB, IEB, MEXU). SLP (Boutin 3397, MEXU, NY, US). Sin (Breedlove 44947, CAS, ENCB). Tam (Sharp et al. 52165, US). Ver (Miller & Torres 2979, MO, UC). Unverified, Doubtful, or Mistaken Reports. Ags (Gonza´lezAdame s.n., HUAA, cited by Siqueiros-Delgado & Gonza´lez-Adame, 2004, but not verified).

ASPLENIUM PSEUDERECTUM

The conspicuously scaly stipes and rachises, 2-pinnate blades, and low angle of minor vein divergence from main veins distinguish A. praemorsum from congeners in Mexico. The close relationship of this taxon with the African A. aethiopicum (Burm.) Becherer (A. furcatum Thunb., A. adiantoides Lam.) needs further study (Morton & Lellinger, 1966; Smith, 1981). Braithwaite (1986) synonymized A. praemorsum under A. aethiopicum, and this judgment was followed by Adams (in Davidse et al., 1995: 295). Most specimens of A. aethiopicum from Africa, especially from South Africa (the type locality) are more dissected (3pinnate), have broader blades, and have narrower ultimate segments (cf., e.g., the illustration in Jacobsen, 1983: 361), than neotropical (especially Mexican and Mesoamerican) material and do not appear to be conspecific. For that reason we maintain both species and use the name A. praemorsum for Mexican plants. Asplenium cuneatum Lam., known from Guatemala (Contreras 3015, 4701, LL), is related to A. praemorsum, and might be expected in Chiapas.

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Distribution. Epipetric on wet cliffs and mossy ledges; 1200– 2300 m; Mexico. Specimens Examined. Chih (Pringle s.n., in 1887, MEXU). Col (Avelar Mun˜oz 16923, UC; Vasquez V. 1565, UC). Jal (Pringle 2769, MO, UC, 4089, MEXU, s.n., MEXU). Nay (Mexia 857, CAS, UC, US).

55. ASPLENIUM PRINGLEI

This species is distinguished by its curved rachises, incised pinnae, and submarginal sori with large, entire indusia. It is closely related to A. sanchezii, A. fibrillosum, and A. blepharodes.

Asplenium pringlei Davenp., Gard. & Forest 4: 449, f. 71. 1891. Type. Mexico. Jalisco: Near Guadalajara, Pringle 1837 (GH?; isotypes CAS!, DS!, NY!–4 sheets, US!, YU!). Fig. 39K.

56. ASPLENIUM PSEUDERECTUM

Roots thin, wiry, not proliferous; rhizomes suberect to erect; rhizome scales dark brown to black, clathrate, ca. 3–4 ⫻ 0.3–0.4 mm, entire; fronds clumped, mostly 10–15 cm long; stipes dark brown to black, lustrous, 1.5–2.5 cm ⫻ 0.7–1 mm, ca. 1⁄10 of frond length, often curved at bases, sparsely scaly, adaxially with wings ca. 0.1 mm wide; blades chartaceous, mostly 8–13 ⫻ 1–1.5 cm, pinnate to pinnate-pinnatifid, linear, lacking buds, rachises black, lustrous, curved, nearly glabrous, adaxially with brownish wings 0.2 mm wide; pinnae oblong, 20–34 pairs, proximal and sometimes middle ones slightly deflexed, 4–5 ⫻ 1.5–3 mm, subsessile, rachis color not running onto the pinna stalks, slightly auriculate acroscopically, apices obtuse, incised 1⁄4–1⁄2 the way to the costae; veins obscure, tips evident adaxially; indument abaxially of scattered appressed, whitish to tan clavate hairs 0.1–0.2 mm long; sori 1–4 pairs per pinna, on both sides of midveins, submarginal on small lobes, 1 sorus per lobe; indusia 0.7–1.2 ⫻ 0.7–1.2 mm, margins entire, often incurved; spores reniform.

Asplenium pseuderectum Hieron., Hedwigia 60: 239. 1918. Lectotype (chosen by Morton & Lellinger, 1966: 17). Puerto Rico. Sintensis 2637 (B; isolectotype US!). Figs. 32E–G. Roots fibrous, not proliferous; rhizomes suberect; rhizome scales generally brown, subclathrate to clathrate, 3–5 ⫻ 0.4–1 mm, entire; fronds clumped, 12–25(–35) cm long; stipes very short to almost obsolete, to 2 cm ⫻ 1 mm, 1⁄10 or less of frond length, glabrous, with green wings to 0.6 mm wide; blades thinly herbaceous, linear-elliptic to oblanceolate, 2.5–5 cm wide near or above the middle, gradually tapering proximally (lowermost pinnae scarcely more than auricles), abruptly caudate-acuminate at apices; rachises greenish to brownish, glabrous, green-winged, the wings 0.2–0.5 mm wide; pinnae narrowly deltate to lanceolate from strongly inequilateral bases, usually somewhat deflexed, mostly 25–35 pairs, 1.5–3 cm ⫻ 4–6 mm at bases, sessile, apices acuminate or somewhat obtuse, margins crenate-dentate on both sides and tips; veins simple except for the basal acroscopic ones, readily visible on both sides, tips evident adaxially; indument abaxially of scattered appressed, whitish to tan clavate hairs 0.1– 0.2 mm long; sori 3–6 pairs per pinna, on both sides of costae; indusia 1–2 ⫻ 0.3–0.4 mm, membranous, margins entire; spores reniform.

114

ASPLENIUM

Distribution. On boulders near water, or mossy tree trunks, in lowland montane rain forests; 150–600 m. Mexico; Guat (Tu¨rckheim s.n. [Donn. Sm. 7808], NY), Bel (Schipp 365, NY), Hond (Wilson 157, NY), Nic (Martı´nez S. 1513, MEXU), CR (Mickel 2900, NY), Pan (Liesner 601, NY); PR. Specimens Examined. Chis (Breedlove 26549, DS, MO, NY, 48393, CAS). Oax (Hallberg 1296, 1311, NY; Herna´ndez G. 1221, CHAPA, NY); Mickel 6454B, NY). Ver (Sinaca C. 1000, IEB, MO).

Proctor (1989) illustrated well the differences between this species and A. pteropus, which is the closest ally. Primarily, A. pseuderectum is distinguished by the proximal pinnae being greatly reduced and by stipes less than 2 cm long. The fronds of A. pseuderectum are also generally about half the size of those of A. pteropus. Adams (in Davidse et al., 1995) did not distinguish these two species in Flora Mesoamericana, so we have cited a representative specimen from each country where A. pseuderectum occurs.

Distribution. Epiphyte in wet forests and rarely on wet rocks; 1000–1600 m. Mexico; Guat, Bel, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Guy, Ec, Peru, s Braz, Bol. Also reported from Honduras and Surinam by Adams (in Davidse et al., 1995). Selected Specimens Examined. Chis (Breedlove 32180, DS, MEXU, 58438, CAS, ENCB, NY; Cabrera 1819, MEXU, MO). Oax (Mickel 1023, 1369A, NY, 5659, ENCB, NY, UC). Pue (Ramı´rez R. 26, XAL). Ver (Ortega O. et al. 1142, ENCB, MEXU, XAL; Va´squez B. & Avendano 1574, XAL; Ventura A. 11613, MEXU, XAL).

This species is closely related to the previous one, A. pseuderectum, and also resembles A. miradorense and A. harpeodes. Asplenium pteropus differs from these by having stipes alate to their bases. It is nearly always epiphytic, or rarely epipetric on mossy rocks, whereas A. miradorense is nearly always terrestrial. A revision of this group (having 1-pinnate proximally reduced blades, winged rachises, and toothed pinnae) is greatly needed. Several specimens identified as this species by Mickel and Beitel (1988) are A. miradorense, e.g., Herna´ndez G. 1323, 1540 (NY).

57. ASPLENIUM PTEROPUS 58. ASPLENIUM PULCHELLUM Asplenium pteropus Kaulf., Enum. Filic. 170. 1824. Type. Brazil. Chamisso s.n. (LE, isotype B). Fig. 34F. Roots fibrous, not proliferous; rhizomes erect; rhizome scales brown to blackish, clathrate, tips twisted and long-attenuate, 6– 9 ⫻ 0.3–0.5 mm, entire; fronds pendent, clumped, 25–45(–60) cm long; stipes stramineous, green or gray-green, 2–8 cm ⫻ 1 mm, ca. 1⁄6–1⁄4 of frond length, glabrous, with green wings 0.5–1 mm wide; blades thin-chartaceous, 1-pinnate with pinnatifid apices, 20–40 ⫻ 5–8 cm; rachises stramineous, dull, glabrous, with green wings 0.5–1 mm wide; pinnae oblong, inequilateral, subfalcate, ca. 25–40 pairs, 3–4 ⫻ 0.7–1 cm, basiscopic bases excavate, subauriculate acroscopically, not overlapping rachises, pinna apices acuminate, margins serrate with rounded teeth, sinuses 1–2 mm deep; veins mostly simple except for basal acroscopic veins, readily visible on both sides, tips expanded adaxially; indument abaxially absent; sori (4–)9–11 pairs per pinna on both sides of midveins, none on basal, forked veins; indusia 1.5–3 ⫻ 0.3–0.6 mm, margins entire; spores reniform; 2n⫽72 (Oax).

Asplenium pulchellum Raddi, Opusc. Sci. 3: 291. 1819. Type. Brazil. Mandiocca, Raddi s.n. (FI; isotypes BR–photos BM, F, UC!, US). Adams (in Davidse et al., 1995: 315) cited the holotype as at PI (photocopy MO!). Fig. 34G. Asplenium pulchellum Raddi var. subhorizontale Rosenst., Repert. Spec. Nov. Regni Veg. 22: 8. 1926. Type. Costa Rica. Rı´o Sumbres, Brade 429 (isotypes NY!, UC!).

Roots thin, fibrous, not proliferous; rhizomes erect; rhizome scales black, clathrate, 1.5–2 ⫻ 0.5–0.8 mm, entire; fronds clumped, 5–12(–18) cm; stipes green with narrow ridges, dull, 1–6 cm ⫻ 0.3–1 mm, 1⁄3 or less of frond length, glabrous or with sparse clavate hairs 0.3 mm long, adaxially with narrow wings. 0.1 mm wide; blades thin-chartaceous, 4–13(–22) ⫻ 1.5–3 cm, 1pinnate, lanceolate, slightly narrowed proximally, apices pinnatifid, not proliferous; rachises green, with sparse, appressed, clavate hairs 0.3 mm long, adaxially with green wings 0.3 mm wide; pinnae oblong-rhombic, falcate, ascending, (5–)7–15 pairs, 0.8–

ASPLENIUM PUMILUM

1.8 ⫻ 0.4–0.8 cm, dimidiate, apices obtuse, toothed on acroscopic margins, sinuses 1 mm or less deep, basiscopic margins entire on proximal 3⁄4; veins forked acroscopically, simple basiscopically, tips evident adaxially; indument abaxially of scattered appressed, tan clavate hairs 0.1–0.2 mm long; sori 1–5 pairs per pinna, on both sides of midveins; indusia 3–5 ⫻ 0.5–0.8 mm, margins entire; spores reniform.

115

with low serrations (sinuses 1 mm or less deep vs. 1–2 mm deep), narrower, more linear blades (vs. ovate), and entire (vs. suberose) indusia. The relationship with A. otites Link, the type of which is a cultivated specimen of uncertain origin (holotype B, isotype BR, photo NY!), is unclear. Morton and Lellinger (1966: 26) circumscribed A. otites in the sense that A. pulchellum is applied here; in this they were followed by Smith (1985) and Mickel and Beitel (1988). According to Morton and Lellinger, A. otites differs from A. pulchellum primarily in its more numerous, often forking veins.

59. ASPLENIUM PUMILUM Asplenium pumilum Sw., Prodr. 129. 1788, non Davenp., 1888. Type. Jamaica. Swartz s.n. (S, photo US!; isotype UPS-THUNB 24835). Figs. 31A, B.

Distribution. Epipetric on shaded limestone bluffs; 200–400(– 1000) m. Mexico; Hond (Thieme 26, UC), Salv, Nic (Proctor et al. 26987, LL), CR, Pan; Col, Ven, Ec, Peru, Braz, Bol. Also reported from Nicaragua, and Paraguay by Adams (in Davidse et al., 1995). Specimens Examined. Chis (Breedlove 39059, DS, NY, TEX). Oax (Herna´ndez G. 1184, CHAPA, ENCB, MEXU, MO, NY, 1584, 1880, CHAPA, MEXU, MO, NY; Mickel 7394, NY, UC).Ver (Chazaro B. et al. 6137, XAL; Ibarra M. 2470, CHAPA, ENCB; Sinaca C. 905, IEB, MO). Unverified, Doubtful, or Mistaken Reports. Tab (reported by Mickel & Beitel, 1988, but not verified and probably based on a misidentification; also reported by Magan˜a, 1992, as A. otites).

Breedlove 39059 differs from other material seen in having larger fronds and reddish brown stipes and rachises, and lacking sharp teeth on the acroscopic pinna margins. Asplenium pulchellum can be confused with several 1-pinnate, green-stiped taxa, especially when these species are precociously fertile. Asplenium pulchellum differs from A. abscissum in smaller frond size (stipes 1–6 cm rather than 13–20 cm, blades usually 4–13 cm rather than 13–20 cm), shorter stipes relative to frond length (1⁄3 or less rather than 1⁄2 of frond length), smaller obtuse pinnae (0.8–1.8 cm long rather than 4.5–8 cm long and acuminate), simply serrate pinna margins (rather than biserrate), and 1-forked acroscopic veins (rather than 2-forked). From A. pteropus, A. pulchellum differs by smaller rhizome scales (1.5–2 mm vs. 6–9 mm long), narrow rachial wings, obtuse, smaller, and fewer pinnae (fewer than 10 vs. more than 30), forked acroscopic veins (vs. simple except for basal vein), and blades only slightly reduced proximally (vs. strongly tapering). Asplenium pulchellum is similar to A. hoffmannii in number of pinnae, rhizome scale size, stipe color and wings, and relative stipe length, but differs in the erect rhizomes, smaller pinnae

Asplenium anthriscifolium Jacq., Collectanea 2: 103, t. 2., f. 3–4. 1789. Type. Martinique. Jacquin s.n. (W, frag. BR, photo US!). Athyrium verapax Christ, Bull. Herb. Boissier, se´r. 2, 6: 296. 1906. Asplenium leucothrix Maxon, Proc. Biol. Soc. Wash. 43: 85. 1930. Type. Guatemala. Alta Verapaz: Cubilguitz, Tu¨rckheim s.n. [Donn. Sm. 8818] (P; isotype US!; frag. NY!).

Roots thin, fibrous, not proliferous; rhizomes erect; rhizome scales black, clathrate, 1–2 ⫻ 0.1–0.3 mm, entire; fronds clumped, 3–20 cm long; stipes castaneous to atropurpureous and sublustrous proximally, greenish or stramineous and dull distally, (3–) 5–15 cm ⫻ 0.4–1 mm, 1⁄2–3⁄4 of frond length, with scattered to dense, white catenate hairs 0.5–0.8 mm long, not winged; blades membranaceous, deltate to pentagonal, mostly (3–)6–12 ⫻ 3–13 cm, pinnate-pinnatifid proximally, apices pinnatifid, not proliferous; rachises green to stramineous, with catenate hairs 0.5–0.8 mm long and greenish ridges or narrow wings 0.1–0.2 mm wide; pinnae deltate, 1–3(–6) pairs, 1–8 ⫻ 0.5–4.5 cm, basal pair largest, greatly exaggerated basiscopically, sessile to stalked to 5 mm, lobed at bases, lobes sometimes nearly free, apices longacuminate, margins crenate-serrate; veins mostly 1–3-forked, evident throughout their length on both sides; indument of both blade surfaces, margins, and veins of whitish catenate hairs 0.3– 0.8 mm; sori 5–20 on the larger pinnae; indusia 3–9 ⫻ 0.5–0.8 mm, margins ciliate; spores reniform; 2n⫽72 (Jam).

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ASPLENIUM

flagelliform, proliferous at tips; rachises castaneous, lustrous, glabrous, with brownish wings 0.1 mm wide; pinnae lanceolate, 12– 25 pairs, 4–8 ⫻ 1.5–2.5 cm, sessile or stalked to 1 mm, pinnate, pinnules oblong to obovate, broadly adnate to cuneate proximally, entire or nearly so, pinna apices acute; veins obscure, tips evident adaxially; indument absent on blades; sori 6–8 pairs on distal 1⁄2 of pinnae, 1–3 pairs per segment (pinnule) in proximal half of pinnate segments; indusia 4–5 mm ⫻ 0.3–0.5 mm, margins suberose; spores reniform; 2n⫽144, 216, 288 (Jam, A. radicans s.l.).

Distribution. Terrestrial and epipetric; 50–1600 m. USA (Fla); Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan (Cornman 574, UC); Gr & L Ant; Col, Ven, Trin, Ec, Galapagos, Peru, Braz, Bol, nw Arg; Africa. Selected Specimens Examined. BCS (Brandegee s.n., UC). Camp (Lundell 1228, DS, MO, NY, UC, US). Chis (Breedlove 47100, CAS, NY). Col (Va´zquez 1170, UC). Gro (Lorea 2540, FCME). Jal (Pringle 2590, UC). Me´x (Hinton 1835, MEXU, US). Mich (Leopo 17082, UC). Mor (Rose & Rose 11078, MEXU, NY). Nay (Jones 23477, CAS, MO, UC). NL (Pringle 2583, CAS, DS, ENCB, MO, NY, US). Oax (Mickel 5821, UC). Qro (Carranza G. 2941, IEB, MEXU). QR (Swallen 2848, US). SLP (Blassingame 1318, US). Sin (Breedlove 35583, CAS, MO). Son (Keil & Canne 8720, NY). Tab (Cowan 1718, CAS, CHAPA, NY). Ver (Dressler & Jones 73, UC). Yuc (Schott s.n., MO, NY).

Asplenium pumilum is closely related to A. minimum, differing by the generally smaller fronds, thinner blade texture, and whitish-hairy blades; the two species agree in having the proximal pinnae more developed basiscopically. A few mixed collections of the two species are known, e.g., Herna´ndez G. 891 (CHAPA). Occasional specimens of A. pumilum are completely glabrous, e.g., Mexia 1182 (MO, UC, US), from Jalisco. Relationships of this species group within Asplenium are uncertain.

60. ASPLENIUM RADICANS Asplenium radicans L., Syst. Nat., ed. 10, 2: 1323. 1759. Lectotype (chosen by Morton & Lellinger, 1966: 37). Jamaica. Browne 92 (LINN 1250.16, right-hand frond). Fig. 38F. Asplenium rhizophyllum L., Sp. Pl., ed. 2, 2: 1540. 1763, hom. illeg., non L., 1753. Type. Same as for A. radicans. Asplenium alleopteron Kunze ex Klotzsch, Linnaea 20: 353. 1847. Syntypes. Guyana. Schomburgk 1150 (B!, frag. NY!), 1206 (B!; isosyntype UC!). Asplenium cyrtopteron Kunze, Linnaea 23: 233, 303. 1850. Lectotype (chosen by Morton & Lellinger, 1966: 37). Cultivated specimen (Hort. berol. ex hb. A. Braun), originally from Venezuela (B!).

Roots wiry, fibrous, not proliferous; rhizomes erect; rhizome scales black, clathrate, 2–3 ⫻ 0.8 mm, entire; fronds clumped, 30– 75 cm long; stipes castaneous to atropurpureous, lustrous, 6–22 cm ⫻ 1–2 mm, 1⁄5–2⁄5 of frond length, glabrous, not winged; blades thick-herbaceous, 15–45 ⫻ 7–15 cm, lanceolate, deeply pinnate-pinnatifid to 2-pinnate, not reduced proximally, apices

Distribution. Terrestrial in wet montane forests; 900–1900 m. Mexico; Guat, Hond, Nic, CR, Pan; Jam, Hisp; Col, Ven, Guy, Peru, s Braz. Also reported from Bolivia by Adams (in Davidse et al., 1995). Selected Specimens Examined. Chis (Breedlove 31318, DS, 32427, DS, MEXU; Ghiesbreght 427, YU; Gittins 4183, NY). Gro (Lorea 2997, FCME). Oax (Hallberg 1553, NY; Mickel 945, NY, UC; Wendt et al. 4643, CHAPA, NY). Pue (Riba s.n., MEXU). Ver (Ramı´rez 654, XAL).

Asplenium radicans is here circumscribed in a narrow sense, comprising only those members of the A. radicans complex that are deeply pinnate-pinnatifid to barely 2-pinnate. To include the entire complex in a single species, from the 1-pinnate A. cirrhatum and A. yelagagense (which see) to 2-pinnate-pinnatifid A. flabellulatum, to fully 3-pinnate A. uniseriale (which see) as varieties of A. radicans (as done by Morton & Lellinger, 1966; Stolze, 1981, Tryon & Stolze, 1993) obscures significant morphological, ecological, and geographical differences between the taxa and gives the impression of having resolved the difficult taxonomic questions raised in this group. Generally, the different species in this group do not grow together. Admittedly, the A. radicans group of terrestrial, flagelliform-tipped species with dark lustrous stipes and rachises is in need of modern biosystematic work. Specimens cited as A. radicans from Veracruz by PalaciosRios (1992) are A. flabellulatum.

61. ASPLENIUM RESILIENS Asplenium resiliens Kunze, Linnaea 18: 331. 1844 [1845], nom. nov. for A. parvulum M. Martens & Galeotti. Fig. 33C.

ASPLENIUM RIPARIUM

Asplenium parvulum M. Martens & Galeotti, Me´m. Foug. Mexique 60, pl. 15, f. 3. 1842, hom. illeg., non Hook., 1840. Type. Mexico. Oaxaca: Environs of Capulalpan and Hacienda del Carmen, Galeotti 6462 (BR!, photos BM, US!; isotypes BR!–photo US!, P!). Roots thin, wiry, fibrous, not proliferous; rhizomes erect; rhizome scales bicolorous, center black, occluded, margins lighter, clathrate, tips uniseriate, 3–5 ⫻ 0.2–0.5 mm, entire; fronds clumped, (3–)8–30(–45) cm long; stipes black to atropurpureous, lustrous, 1–3 cm ⫻ 0.5–1 mm, 1⁄10–1⁄3 of frond length, with scattered, hair-like, clathrate scales 1–1.5 mm long and appressed, tan, clavate hairs 0.1–0.2 mm long, adaxially with whitish to tan wings 0.1–0.2 mm wide; blades subcoriaceous, firm, (5–)9–25 ⫻ 1–2 cm, 1-pinnate, linear-lanceolate, proximal 2–6 pinna pairs often reduced and deflexed, apices pinnatifid, not proliferous; rachises blackish, lustrous, glabrous or with a few appressed, tan clavate hairs, adaxially with whitish to tan wings 0.1–0.3 mm wide; pinnae oblong, 20–40 pairs, (4–)10–20 ⫻ 2–5 mm, sessile, articulate, rachis color not extending onto pinna stalks, bases somewhat auriculate, apices obtuse, margins subentire or crenate, often with revolute margins; veins 1-forked, obscure, tips not evident adaxially; indument abaxially of scattered clavate hairs 0.1 mm long; sori (1–)2–6(–10) pairs per pinna, on both sides of midveins; indusia 0.8–1.5(–2) ⫻ 0.3–0.5 mm, margins erose; spores globose to ovoid; 2n⫽108, apogamous (Jam).

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Specimens cited as this species from Chihuahua by Knobloch and Correll (1962) are A. gentryi (e.g., Correll & Gentry 23032; Gentry 8064). Also, inclusion of Chihuahua and Sonora in the range of A. resiliens by Mickel and Beitel (1988) and by Mickel (1992) was based in part on misidentified specimens, but A. resiliens does occur in both states. Asplenium resiliens appears to be most common in northeastern Mexico, in the states of Nuevo Leo´n and Coahuila. Asplenium resiliens is closely related to A. monanthes, but is distinguished from that, as well as other dark-stiped Aspleniums (except A. heterochroum and A. palmeri), by the lack of conspicuous hydathodes adaxially. It is distinguishable from A. heterochroum by the thicker and more deflexed pinnae, forked fertile veins, and occurrence at higher elevations (usually above 1400 m in A. resiliens). Asplenium resiliens differs from A. palmeri in having nonproliferous blade apices, proximal pinnae deflexed, subentire to weakly crenulate pinna margins, and longer stipes. A few mixed collections with other species in the group have been seen, e.g., Copeland herb. 67, UC, from Morelos, which also contains a frond of A. hallbergii.

62. ASPLENIUM RIPARIUM Asplenium riparium Liebm., Mexic. Bregn. 244 (reprint 92). 1849. Fig. 34H. Asplenium obtusifolium L. var. riparium (Liebm.) Domin, Pterid. Dominica 175. 1929. Lectotype (chosen by Smith, 1981: 51). Mexico. Veracruz: Hacienda de Jovo, Liebmann s.n. [Pl. Mex. 2283, Fl. Mex. 310] (C!; isotype B!).

Distribution. Terrestrial or epipetric, in rock crevices, limestone cliffs, pine-oak forests; (300–)1400–3150 m. S USA; Mexico; Guat; Jam, Hisp; Col, Ven, Ec, Peru, Braz, Bol, n Arg, Uru. Selected Specimens Examined. Chih (Johnston et al. 10766, LL); Chis (Mickel 1210, NY, UC). Coah (Muller 3250, UC, US). DF (Schaffner 109, NY p.p.). Gro (Lorea 2651, FCME). Hgo (Lyonnet 1443, US). Jal (Gonza´lez 18935, GUADA). Mor (Copeland herb. 67 p.p., UC, US). NL (Cowan 3737, UC). Oax (Mickel 3721, NY, UC). Pue (Purpus 3399, MO, US). Qro (Dı´az B. 3840, ENCB, IEB). SLP (Roe & Roe 2199, US). Son (Reichenbacher 1281, ARIZ). Tam (Stanford et al. 649, ARIZ, DS, MEXU, MO, UC). Tlax (Acosta P. 2174, XAL). Ver (Fisher 35382, MO, US). Zac (Johnston et al. 10473, CAS, MEXU). Unverified, Doubtful, or Mistaken Reports. Me´x (McVaugh 4005, ENCB, not seen, cited by Tejero-Dı´ez & Arreguı´n-Sa´nchez, 2004, but not verified).

Roots coarse, fibrous, not proliferous; rhizomes long- to shortcreeping; rhizome scales obscure, red-brown, clathrate, 0.5–2 ⫻ 0.2–0.3 mm, entire; fronds few, 0.5–1 cm apart, 25–60 cm long; stipes brown, dull, 12–30 cm ⫻ 2–3 mm, 1⁄3–1⁄2 of frond length, glabrous or with rare clathrate scales at bases, with green-brown wings 0.3–1 mm wide, flattened and wider proximally (to 4 mm); blades herbaceous, lanceolate, 1-pinnate, 16–35 ⫻ 7–15(–21) cm, reduced proximally, apex of each blade with a large hastate conform terminal pinna, not proliferous; rachises dark green, dull, glabrous, with wings 0.3–1 mm wide; pinnae trapezoidal to lanceolate or falcate, 7–15 pairs, proximal 1–3 pairs slightly reduced, 1–8(–11) ⫻ 1–2 cm, stalked to 4 mm, inequilateral, basiscopic sides excavate, acroscopic sides sometimes auriculate, apices usually acute to acuminate, margins coarsely and irregularly sharply serrate or crenate-serrate; veins mostly 2-forked, visible on both sides of blades, vein ends slightly widened and prominent adaxially; indument abaxially of scattered, appressed, clavate hairs 0.1 mm long; sori 3–11 pairs per pinna, on both sides of midveins; indusia brown, 4–9 ⫻ 0.5–1 mm, margins entire; spores reniform, 64 per sporangium (Murakami & Moran, 1993); 2n⫽78–80 (Chis).

118

ASPLENIUM

Distribution. Terrestrial, or more frequently on rocks in streams in wet forests, especially on the Atlantic slope; 150– 2100 m. Mexico; Guat, Bel (Holst 5332, UC), Hond, Nic, CR, Pan; Col, Ven, Ec.

pureous, lustrous, 1–3 cm long, to 1⁄10 of frond length, with appressed clavate hairs 0.1–0.2 mm long, adaxially with brownish wings 0.1 mm wide; blades thin-herbaceous, 15–35 ⫻ 4–10(–15) cm, 3-pinnate, elliptical, gradually tapered both proximally and distally, apices elongate, naked, proliferous; rachises castaneous to atropurpureous, lustrous, glabrous or with scattered appressed clavate hairs 0.1 mm long, adaxially with wings 0.1–0.3 mm wide; pinnae narrowly deltate, somewhat falcate, 10–24 pairs, 2.5–9 ⫻ 1–2.5 cm, sometimes ending in naked flagelliform apices, sessile, usually 2-pinnate, proximal pinnules essentially opposite, perpendicular to and overlapping rachises, ultimate segments elliptic or obovate, mostly 3–5 ⫻ 1–1.5 mm, bases cuneate, apices acute, margins entire, segments towards apices of pinnae slightly falcate and sometime slightly gibbous on indusia-bearing side; veins solitary in the ultimate segments, tips readily evident adaxially; indument abaxially of appressed clavate hairs 0.1–0.2 mm long; sori 1 per ultimate segment; indusia 1.5–2.5 ⫻ 0.3–0.6 mm, margins minutely erose; spores reniform.

Selected Specimens Examined. Chis (Breedlove 32617, DS, NY, 35410, DS; Matuda 221, MEXU, US). Oax (Herna´ndez G. 1749, CAS, MO; Mickel 5918, NY, UC, 6580, NY, UC). Pue (Campos V. 401, IEB; Riba et al. 318, NY). Ver (Purpus 2846, MO, NY, UC; Ventura A. 3267, CHAPA).

The name A. repandulum, previously used for this species in Mexico (Smith, 1981; Mickel & Beitel, 1988), was restricted to specimens from Ecuador and Peru by Murakami and Moran (1993). They regarded A. riparium as belonging to sect. Hymenasplenium. Morton and Lellinger (1966) reported A. riparium (as A. repandulum) as an epiphyte; however, the Mexican specimens appear to be terrestrial or epipetric in or along streams. This and the closely allied A. obtusifolium L., from Panama, Greater and Lesser Antilles, Colombia, Venezuela, and Trinidad, both grow on wet rocks, are fleshy and dry blackish, have flattened stipes, creeping rhizomes, and somewhat irregular blade dissection. Asplenium obtusifolium has smaller fronds 15–20(–25) cm long, shorter, more obtuse, somewhat flabellate, pinnae 1.5– 3.5 cm long, and proximal 1–4 pinnae each with a free acroscopic basal lobe.

63. ASPLENIUM RUTACEUM Asplenium rutaceum (Willd.) Mett., Abh. Senckenberg. Naturf. Ges. 3: 173. 1859. Fig. 38A. Aspidium rutaceum Willd., Sp. Pl., ed. 4, 5(1): 266. 1810. Lectotype (chosen by Morton & Lellinger, 1966: 36). Plumier, Traite´ Foug. Ame´r., pl. 57, 1705, illustrating a plant from Hispaniola (contrary to Adams in Davidse et al., 1995). Asplenium conquisitum Underw. & Maxon ex Christ, Bull. Herb. Boissier, se´r. 2, 7: 270. 1907. Lectotype (chosen by Maxon, Contr. U.S. Natl. Herb. 10: 488. 1908). Jamaica. Maxon 1558 (P!; isolectotype US!). Additional synonymy given in Morton & Lellinger (1966).

Roots fibrous, not proliferous; rhizomes erect; rhizome scales black, strongly clathrate, 3–4 ⫻ 0.3–0.8 mm, entire; fronds clumped, mostly 20–45 cm long; stipes castaneous to atropur-

Distribution. Terrestrial, or occasionally epipetric or epiphytic in wet montane forests; 600–1800 m. Mexico; Guat, Nic, CR, Pan; Jam, Hisp, PR; Col, Ven, Guy, Sur, Ec, Peru, Bol. Selected Specimens Examined. Chis (Breedlove 29807, DS, 32199, DS, MEXU, 38934, DS; Ghiesbreght 382, K, PH). Oax (Mickel 941, US, 5655, NY, UC, 6549, MEXU, NY, UC; Rzedowski 34077, IEB). Ver (Lira 217, MEXU, XAL; Ramı´rez 782, 848, XAL).

The atropurpureous lustrous stipes and rachises, 3-pinnate blades, proliferous and flagelliform blade apices, and slightly gibbous sorus-bearing segments distinguish this species from the similarly dissected A. cristatum and A. myriophyllum, which are probably not closely related.

64. ASPLENIUM SALICIFOLIUM Asplenium salicifolium L., Sp. Pl. 2: 1080. 1753. Lectotype (chosen by Morton & Lellinger, 1966: 23). Plumier, Descr. Pl. Ame´r., pl. 27. 1693 [⫽ Traite´ Foug. Ame´r., pl. 60. 1705], illustrating a specimen collected by Plumier from either Hispaniola or Martinique, without definite locality. Fig. 36J.

ASPLENIUM SANCHEZII Asplenium bicrenatum Liebm., Mexic. Bregn. 245 (reprint 93). 1849. Type. Mexico. Oaxaca: Dist. Chinantla, Liebmann s.n. [Pl. Mex. 2292, Fl. Mex. 320] (C!, frag. US!).

Roots relatively coarse, fibrous, not proliferous; rhizomes erect; rhizome scales orange-brown, clathrate, 4–7 ⫻ 0.4–0.8 mm, margins with scattered golden hairs 0.8–2 mm long but otherwise entire; fronds clumped, 25–60 ⫻ 10–20 cm; stipes gray-green or brownish green, dull, 9–28 cm ⫻ 2–3.5 mm, ca. 1⁄3–1⁄2 of frond length, glabrous or nearly so, not alate or only obscurely greenwinged; blades thick-herbaceous, 1-pinnate, lanceolate, 16–38 ⫻ 8–20 cm, apices pinnatifid or hastate and Ⳳ confluent, not proliferous; rachises yellowish green to tan, dull, glabrous, with wings 0.1–0.3 mm wide; pinnae lanceolate, falcate, 7–16 pairs, 5.5–9 cm ⫻ 10–15 mm (excluding acroscopic lobe), stalked 2–4 mm, bases inequilateral, acroscopically subcordate with auricles rounded and overlapping rachises, basiscopically excised or cuneate, apices long-attenuate, margins simply or doubly crenate with (17–)25–30 low teeth on acroscopic margins; veins mostly 1–2-forked, obscure, tips evident adaxially; indument abaxially of scattered, appressed, tan, clavate hairs ca. 0.1–0.2 mm long; sori 7–15 pairs per pinna, on both sides of midveins; indusia 5–10 ⫻ 0.5–1 mm, entire; spores reniform; 2n⫽288 (Trin).

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recognized a second variety, var. aequilaterale (Christ) C. D. Adams, confined to Costa Rica and Panama.

65. ASPLENIUM SANCHEZII Asplenium sanchezii A. R. Sm., sp. nov. Type. Mexico. Durango: Camino de Espinazo del Diablo at Huizar, Carr. Durango–Mazatla´n, 2000 m, Sa´nchez S. 749 (holotype US!; isotypes MEXU!, UC!). Figs. 39C, D. Ab A. castaneo pinnis paucioribus 6–15-jugis, frondibus parvioribus, apicibusque marginibus acroscopicis pinnarum perspicue crenulatis, segmento apicali laminarum distinctiore differt; ab A. blepharode et A. fibrilloso indusiis integris differt. (Honoring Jose´ Sa´nchez S., collector of plants in Mexico.)

Roots thin, wiry, fibrous, not proliferous; rhizomes suberect to erect; rhizome scales dark brown to black, clathrate, ca. 3 ⫻ 0.3– 0.4 mm, entire; fronds clumped, 10–12 cm long; stipes castaneous, lustrous, 1–2.5(–6) cm ⫻ 0.5–1 mm, 1⁄10–1⁄2 of frond length, sparsely scaly, adaxially not winged or wings brown, less than 0.1 mm wide; blades chartaceous, 3–10 ⫻ 1–2 cm, 1-pinnate, linear, ending in rather large triangular or hastate terminal segments 5– 10 ⫻ 4–10 mm, not proliferous; rachises castaneous to atropurpureous, lustrous, with a few hair-like scales to 1 mm, adaxial wings 0.1 mm wide or less; pinnae oblong, 6–15 pairs, the proximal ones slightly deflexed, 4–6(–10) ⫻ 2–4(–7) mm, subsessile, rachis color not running onto the pinna stalks, bases not or slightly auriculate, apices obtuse, acroscopic margins crenulate; veins mostly 1-forked, obscure, tips evident adaxially; indument abaxially of scattered, appressed, tan, clavate hairs 0.1 mm long; sori 2–4 pairs per pinna, on both sides of midveins; indusia whitish to brown, 1–2 ⫻ 0.7–0.9 mm, margins entire; spores very large, reniform.

Distribution. Mostly epiphytic in wet montane forests; 100– 1200 m. Mexico; Guat, Bel, Hond, CR, Pan; Gr & L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol. Reported from Nicaragua by Go´mez (1976: 53). Selected Specimens Examined. Chis (Breedlove 32933, DS, MEXU, 41958, DS, MEXU, 52541, CAS, 57628, CAS, ENCB, NY). Oax (Mickel 5847, ENCB, NY, UC, US, 7159, 7422, NY, UC; Torres C. 8664, IEB, MEXU). Ver (Copeland herb. 59, MEXU, UC, US; Finck 46a, US, 142, US).

Asplenium salicifolium is closely related to the much more common A. auriculatum, and differs from that in the larger fronds with more remote and larger pinnae 5.5–9 cm long, more teeth on the acroscopic pinna margins (17–39), often 2-forked veins, and more pairs of sori per pinna (7–15). Proctor (1985: 378) regarded A. salicifolium and A. auriculatum as varieties of a single species. Torres C. 8664, cited above, is somewhat intermediate between the two species. Adams (in Davidse et al., 1995)

Distribution. On rocks and walls of moist cliffs; 1800–2500 m. Mexico. Specimens Examined. Chih (Correll & Gentry 23030, LL). Dgo (Oliver et al. 797, MO; Reeder & Reeder 2493, US). Oax (Frame 312, NY, UC). Sin (Breedlove 35730, CAS, MO; Ownbey & Ownbey 1947, US).

This delicate new hydathodous spleenwort may be most closely related to A. castaneum, from which it differs in the gen-

120

ASPLENIUM

erally fewer pinna pairs and smaller fronds, more decidedly crenulate pinna apices and acroscopic margins, and in the larger, often hastate segments at the blade tips. Asplenium sanchezii differs from A. blepharodes and A. fibrillosum in having entire indusia. From A. gentryi and A. hallbergii, A. sanchezii differs in having smaller fronds and fewer pinnae, 6–15 pairs. From A. soleirolioides and A. stolonipes it differs in the nonproliferous blades. From A. pringlei, it differs in having the sori not on pinna lobes. From A. oligosorum and A. olivaceum it differs in the sori located on both sides of the midveins, usually 2–4 pairs per pinna. Specimens in herbaria have previously been determined as A. castaneum, A. monanthes, or A. pringlei. The sole collection from Chihuahua was cited by Knobloch and Correll (1962: 156) as A. castaneum. The apparent disjunct distribution in Mexico can be expected to be filled by additional collections.

66. ASPLENIUM SCOLOPENDRIUM VAR. AMERICANUM Asplenium scolopendrium L. var. americanum (Fernald) Kartesz & Gandhi, Phytologia 70: 196. 1991 [20 May], comb. superfl. published by Reichstein et al., Biol. Jaarb. 59: 162. 1991 [25 June 1992]. Figs. 35A, B. Phyllitis scolopendrium (L.) Newman var. americana Fernald, Rhodora 37: 220, t. 225, f. 2, 4, 6. 1935. Phyllitis fernaldiana A. Lo¨ve, Svensk. Bot. Tidskr. 48: 214. 1954. Phyllitis japonica Komarov subsp. americana (Fernald) A. Lo¨ve & D. Lo¨ve, Acta Bot. Acad. Sci. Hung. 19: 205. 1973. Type. Canada. Ontario: Grey Co., Inga Falls, Fernald et al. s.n. (GH!). Scolopendrium lindenii Hook., Icon. Pl. 5: t. 488. 1842. Phyllitis lindenii (Hook.) Maxon, Fernwort Papers 2: 30. 1900. Phyllitis scolopendrium (L.) Newman var. lindenii (Hook.) Fernald, Rhodora 37: 221. 1935. Asplenium scolopendrium L. var. lindenii (Hook.) Viane, Rasbach & Reichstein, Biol. Jaarb. 59: 162. 1991 [25 June 1992]. Type. Mexico. Chiapas: “Chamulars” [Chamula], Linden 1543 (K!).

Roots coarse, fibrous, not proliferous; rhizomes erect; rhizome scales blackish to dark brown, clathrate, 1.5–6 ⫻ 1–1.5 mm, entire or weakly short-ciliate; fronds clumped, 9–65 cm long; stipes brownish to stramineous, dull, 1.5–20 cm ⫻ 1.5–5 mm, ca. 1⁄5– 1⁄3 of frond length, with dense, linear to lanceolate, tan, ultimately clathrate scales, not alate; blades thick-herbaceous to subcoriaceous, simple, 7–22(–35) ⫻ 1.8–5 cm, oblong, entire to slightly undulate, bases cordate to subcordate or with two downward-directed lobes, apices acute, proliferous buds absent; rachises brownish at bases, light green to stramineous distally (lighter than the blades), dull, with sparse to dense scales 1–2.5 ⫻ 0.1–0.2 mm, not winged; pinnae absent; veins simple or 1- or 2-forked, obscure, parallel for most of their length, tips evident adaxially; indument abaxially of scattered, linear scales to 0.5–1 ⫻ 0.1 mm, also with sparse catenate hairs to 0.5 mm long; sori paired, facing each other on adjacent veins, usually confined to distal 1⁄2–2⁄3 of blades; indusia (2–)6–15 ⫻ 0.3–0.7 mm, margins entire or repand; spores reniform; 2n⫽144 (Can, USA, Chis, see Viane & Reichstein, 1991: 163).

Distribution. Epipetric on moist calcareous rocks (dolomitic limestone) in sinkholes, at cave entrances, and on cool, moist talus, and on limestone cliffs, occasionally epiphytic on old oaks, in deep shade; 2100–2200 m. Can (Ont), USA (spotty in Ala, Mich, NY, Tenn); Mexico; Hisp (Haiti). Specimens Examined. Chis (Fraser-Jenkins 13213, UC; Garcı´a F. 567, XAL; Ghiesbreght 296, K, 307, K, NY, PH; Mu¨nch 80, DS). NL (Favela L. 210, 215, ENCB, 212, ANSM). Oax (Mickel 5385, NY).

Asplenium scolopendrium var. americanum is extremely rare in Mexico. Plants from southern Mexico have been called Phyllitis lindenii, or Asplenium scolopendrium var. lindenii, which supposedly differs from the eastern North American var. americanum in the less prominently cordate blade bases and more abundantly scaly stipes and rachises. These differences seem part of the normal variation of the species, and the scale difference is probably a function of age of fronds and state of preservation. Specimens from Chiapas have their blade bases exactly like those from further north. Apparently, the two varieties do not differ in chromosome number, as previously supposed (see Viane & Reichstein, 1991). The chromosome count by Lo¨ve and Lo¨ve (1973) of n⫽36 is very likely erroneous (no voucher exists; Brouillet, pers. comm.). Asplenium scolopendrium L. var. scolopendrium [Sp. Pl. 2: 1079. 1753. Phyllitis scolopendrium (L.) Newman, Hist. Brit. Ferns, ed. 2, 10. 1844. Type. Europe (LINN)] occurs in Europe, Macaronesia, and western Asia. The East Asian variant of Asplenium scolopendrium, also tetraploid, is sometimes considered a separate species Phyllitis japonica Kom. or Asplenium komarovii Akasawa, but it seems only weakly, if at all, separable.

67. ASPLENIUM SEILERI Asplenium seileri C. D. Adams, Novon 2: 291. 1992. Type. El Salvador. Laguna Verde, Seiler 957 (F). Figs. 43H, J. Roots thin, fibrous, not proliferous; rhizomes erect; rhizome scales dark brown to black, clathrate, 1.5–2.5(–3) ⫻ 0.4–0.7 mm, entire; fronds densely clumped, 15–20 cm long; stipes brownish, dull, 0.5–4 cm ⫻ 0.6–0.8 mm, 1⁄5 or less of frond length, narrowly green- or brown-winged to bases, glabrous or with sparse

ASPLENIUM SEPTENTRIONALE

clavate hairs 0.3 mm long, wings to ca. 0.2 mm wide; blades thin-chartaceous, 12–18 ⫻ 1.5–2.5(–3) cm, 1-pinnate, linearelliptic, decidedly narrowed proximally, apices pinnatifid, nonproliferous or with small buds at the tips of blades; rachises greenish, dull, with sparse clavate hairs 0.3 mm long, adaxially with greenish wings 0.2–0.3 mm wide; pinnae oblong, spreading, falcate, 30–40 pairs, 0.8–1.5 ⫻ 0.4(–0.6) cm, dimidiate, apices obtuse, margins minutely crenulate acroscopically, sinuses mostly 0.5 mm or less deep, basiscopic margins entire in proximal 3⁄4; veins simple, tips evident adaxially; indument abaxially of scattered, tan, appressed, clavate hairs 0.1–0.2 mm long; sori 3–6 on acroscopic sides of pinnae, 2–4 on basiscopic sides; indusia 1.5–3 ⫻ 0.4–0.5 mm, margins entire; spores reniform.

Distribution. Terrestrial or epiphytic in montane forests; ca. 1700 m. Mexico; Guat, Salv. Also reported from Costa Rica by Adams (in Davidse et al., 1995: 319). Specimens Examined. Chis (Croat 47144, MO, UC; Matuda 186bis, US). Adams (in Davidse et al., 1995: 319) discussed an epipetric specimen from Chiapas, Breedlove 42513, MEXU, as this species.

This species is most similar to A. pulchellum, but differs from that in the more numerous pinnae (ca. 30–40 pairs) and the minutely crenulate pinna margins. In Mexico, it is known from a collection from Mt. Ovando and another from Volca´n Tacana´. The Matuda collection was cited by Smith (1981: 55) as possibly representing an undescribed species. Matuda 186 (MEXU p.p., US), from Pasital, seems to be A. potosinum, vel aff. The Croat collection cited bears small buds at the tips of some blades.

68. ASPLENIUM SEMIPINNATUM Asplenium semipinnatum (Hieron.) A. R. Sm., comb. et stat. nov. Fig. 44B. Asplenium potosinum Hieron. var. semipinnatum Hieron., Hedwigia 60: 247. 1918. Type. Mexico. San Luis Potosı´: Tamasopo Canyon, Pringle 3367 (B; isotypes DS!, MO!, NY!–4 sheets, UC!, US!–2 sheets).

Roots fibrous, not proliferous; rhizomes erect; rhizome scales blackish to dark brown, clathrate, ca. 4 ⫻ 0.8 mm, subentire; fronds clumped, to ca. 65 cm long; stipes dark gray-brown, dull, 8–12 cm ⫻ 1.5–2.5 mm, ca. 1⁄5 of frond length or less, glabrous

121

except for scattered scales at bases, adaxially with green wings 0.3–0.4 mm wide; blades chartaceous, linear-lanceolate, greatly reduced proximally, 40–50 ⫻ 4.5–8 cm, 2-pinnate nearly to the apices, pinnate at apices, rachises terminating in small scaly bud or plantlet with fronds to 4 cm long; rachises brownish, dull, glabrous, with green wings 0.4–0.6 mm wide; pinnae lanceate, 40–50 pairs, 2.5–4 ⫻ 0.8–1.3 cm, sessile or stalked to 0.5 mm, the most developed pinnae with to 8 pairs of oval or elliptic pinnules, the pinnules subentire or toothed near the tips, apices rounded to acutish; veins mostly 2–3 pairs in pinnules, tips evident adaxially; indument abaxially of scattered, appressed, tan, clavate hairs 0.1–0.2 mm long; sori 1–5 per pinnule, on both sides of midveins; indusia mostly 1–2.5 ⫻ 0.5 mm, margins entire or subentire; spores reniform.

Distribution. Terrestrial in wet montane forests; middle elevations. Mexico. Specimen Examined. SLP (Pringle 4074, DS, UC, from the same locality as the type, 4 Dec 1891).

This species, newly elevated to species status here, is fully 2-pinnate. Probably it is related to A. sessilifolium, but differs by the 2-pinnate blades and more robust fronds to 65 ⫻ 8 cm. Spores of Pringle 4074 have been examined and appear well formed. From this same locality, Hieronymus described A. potosinum Hieron. var. incisum Hieron. (Hedwigia 60: 247. 1918), based on Pringle 3968 (B!; isotypes ARIZ!, DS!, K!, LE, MEXU!, MO!, NY!–2 sheets, UC!, US!–2 sheets). This is intermediate morphologically between A. semipinnatum and A. sessilifolium and has malformed sporangia and poorly formed or no spores. We regard it as a probable hybrid. Another collection from the same locality is Pringle 3412 (LL). A third collection of the putative hybrid, from Edo. Hidalgo, is Yatskievych 83-131 (ARIZ).

69. ASPLENIUM SEPTENTRIONALE Asplenium septentrionale (L.) Hoffm., Deutsch. Fl. 2: 12. 1796. Figs. 41G, H. Acrostichum septentrionale L., Sp. Pl. 2: 1068. 1753. Lectotype (chosen by Jonsell & Jarvis, 1994: 150). Europe. Herb. Burser XX: 37 (UPS, microfiche UC!).

122

ASPLENIUM

Roots filamentous, wiry, not proliferous; rhizomes erect, much branched to produce many-stemmed tufts or mats bearing numerous crowded fronds; rhizome scales dark reddish brown to black, clathrate, 2–4 ⫻ 0.2–0.4 mm, entire; fronds clumped, 2– 12 cm long; stipes dark reddish brown proximally, greenish distally, dull, 2–11 cm ⫻ 0.2–0.3 mm, 2⁄3–9⁄10 of frond length, glabrous, not alate but with two thickened adaxial ridges; blades thick-herbaceous, linear, simple to 1-forked or lacerate at the tips, occasionally 1-pinnate with a single lateral pinna, mostly 0.5–2.5 ⫻ 0.1–0.2 cm for unforked blades, to 1 cm if the single pinna strongly diverges, glabrous, bases acute, apices acute, not rooting at tips; rachises green, dull, glabrous; pinnae commonly none or one per frond and then strongly ascending, linear, 5–10 ⫻ 0.75–2 mm, apices acute, margins entire to remotely lacerate; veins 1-forked or absent except for main midrib, obscure, tips not evident adaxially; indument not seen, blades appearing glabrous; sori usually 2⫹ per pinna, parallel to margins and facing each other; indusia whitish to tan, mostly 5–20 ⫻ 0.5 mm, margins entire to subentire; spores reniform, 64 per sporangium; 2n⫽144 (Europe).

s.n. [Langsdorff] (isotypes LE!, B-Willd. 19880!, photo US!). Fig. 31F. Asplenium insigne Liebm., Mexic. Bregn. 246 (reprint 94). 1849, non Blume, 1828, hom. illeg. Lectotype (chosen by Smith, 1981: 52). Mexico. Oaxaca: Between Totontepec and Trapiche de la Concepcio´n, Liebmann s.n. [Pl. Mex. 2293, Fl. Mex. 300] (C!; isolectotype US!). Asplenium progrediens Fe´e, Me´m. Foug. 8: 81. 1857. Syntypes. Mexico. [Veracruz:] “pre`s de Huatusco, 1854,” Schaffner 54 (RB); Orizaba, 1856, Schaffner 449 (P?). Additional synonymy given in Morton & Lellinger (1966).

Roots coarse, fibrous, not proliferous; rhizomes short-creeping to suberect; rhizome scales black, dense, clathrate, tips filiform, 5–7 ⫻ 0.5–1 mm, entire; fronds clumped, (30–)50–150 cm long; stipes castaneous to atropurpureous, dull, 10–40 cm ⫻ 2–8 mm, 1 ⁄3 of frond length, decidedly scaly at bases, glabrous or glabrescent distally, not winged; blades thick-chartaceous to subcoriaceous, 1-pinnate, 24–95 ⫻ 10–25 cm, narrowly deltate, not radicant, each blade with a subconform terminal pinnae; rachises castaneous to atropurpureous, dull or sublustrous, abaxially with scattered, tan, twisted, hair-like scales 1–3 mm long, not winged; pinnae lanceolate, 6–15(–20) pairs, 7–15(–20) ⫻ 1.5–2(–3) cm, stalked to 5 mm, Ⳳ equilateral, exauriculate, bases broadly cuneate to truncate, apices long-attenuate, margins serrate or biserrate; veins mostly 2–3-forked, at 5–30⬚ angle from costae, tips not strongly evident adaxially, but veins visible on both surfaces; indument abaxially of scattered, tan, pectinate or substellate scales 0.1–0.5 mm long; sori 5–15 pairs per pinna, on both sides of and nearly parallel to costae; indusia 5–20 ⫻ 0.8–1 mm, margins entire; spores reniform; 2n⫽144 (Galapagos), ca. 288 (Trin).

Distribution. In volcanic and granitic rock crevices; 2200– 3050 m. W USA (disjunct in W Va); Mexico; Europe, Asia. Specimens Examined. BCN (Brandegee s.n., 18 May 1893, DS, NY, UC, US; Moran & Thorne 14343, ENCB).

Two subspecies are recognized in Europe (Reichstein, 1981); ours is subsp. septentrionale. Asplenium septentrionale is easily distinguished from all other spleenworts in Mexico by the linear, grass-like or distally 1-forked blades, with blade segments (or occasionally pinnae) only ca. 1–2 mm wide. Farther north in the range, in California, this rare species grows in holes or crevices of volcanic boulders, and that also appears to be the favored habitat in the Sierra San Pedro Ma´rtir, the only area where it is known in Mexico.

Distribution. Epiphytic, occasionally terrestrial; 600–2800 m. Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Ec, Galapagos, Peru, s Braz, Bol, Parag, nw Arg, Uru.

70. ASPLENIUM SERRA

Selected Specimens Examined. Chis (Breedlove 22025, DS; Herna´ndez & Ton 532, MO, UC). Gro (Hinton 14279, MO, NY, US; Lorea 3028, FCME). Hgo (Moore 3210, UC). Oax (Mickel 5941, MEXU, NY; Wendt et al. 4659, MEXU, NY). Pue (Arreguı´n 548, CHAPA, ENCB). Ver (Purpus 6434, MO, NY, UC, US; Ventura A. 12354, MO).

Asplenium serra Langsd. & Fisch., Pl. Voy. Russes Monde 1: 16. 1810 (t. 19, 1818). Type. Brazil. Santa Catarina: Krusenstern

Asplenium serra is a member of sect. Sphenopteris, a group characterized by scaly rachises and sori at relatively small, acute angles with respect to the midribs. The most nearly related con-

ASPLENIUM SESSILIFOLIUM

geners in Mexico are A. dissectum and A. praemorsum. From the former A. serra is readily distinguishable by the larger, less incised, much longer, thicker-textured pinnae and non-glandular stipes, from the latter, by the 1-pinnate (vs. 2-pinnate) blades.

71. ASPLENIUM SERRATUM Asplenium serratum L., Sp. Pl. 2: 1079. 1753. Lectotype (chosen by Proctor, Fl. L. Antill. 2: 313. 1977). Plumier, Descr. Pl. Ame´r., pl. 39, 1693, based on material from either Martinique or Hispaniola. Proctor rejected the choice by Morton and Lellinger (1966). Figs. 31H, J. Roots coarse, fibrous, forming a dense mat, not proliferous; rhizomes erect, clothed with dense mass of roots and old leaf bases; rhizome scales black, clathrate, (4–)8–20 ⫻ 1–2 mm, margins with retrorse hairs with attenuate or filiform tips; fronds clumped, (25–)40–100 cm long; stipes blackish or brown to greenish, dull, usually less than 2 cm (occasionally to 10 cm) ⫻ 2–5 mm, glabrous, alate; blades thick herbaceous, simple, unlobed, 35–75(–95) ⫻ (4–)6–10(–14) cm, narrowly oblanceolate, bases long-cuneate, tips acute, margins crenate to finely serrulate, bearing tan scales along both sides of rachises, especially abaxially; rachises greenish, dull, abaxially with scattered scales 1–3.5 mm; pinnae none; veins simple or often 1-forked near their bases, running Ⳳ parallel to each other, usually visible on both sides, tips evident or not adaxially; indument abaxially of scattered, tan pectinate scales 0.3–0.5 mm long on and between veins; sori 100⫹ pairs per blade, parallel and at ca. 70⬚ from the midribs, often confined to the proximal 1⁄3–2⁄3 of veins portion of blades; indusia whitish, (5–)12–25 ⫻ 0.4–0.5 mm, margins entire; spores reniform; 2n⫽144 (Fla), ca. 288 (Trin).

123

Unverified, Doubtful, or Mistaken Reports. QR (Trejo et al. 113, UADY, cited by Palacios-Rios, 2002c, but not verified).

Asplenium serratum is easily distinguished from congeners in Mexico by the simple, nest-forming fronds. The only other simple-bladed asplenioids in Mexico are A. scolopendrium, with sori in pairs facing one another and cordate blade bases, Holodictyum, which has reticulate venation, and Schaffneria, with rounded blades and atropurpureous, lustrous stipes.

72. ASPLENIUM SESSILIFOLIUM Asplenium sessilifolium Desv., Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck. Gesammten Naturk. 5: 322. 1811. Type. “America equinoctiali.” s. coll. s.n. (P!, photos BM, US; isotype B!). Roots wiry, fibrous, not proliferous; rhizomes erect; rhizome scales black to dark brown, clathrate, 2–3(–5) ⫻ 1–1.3 mm, with occasional teeth; fronds clumped, mostly 25–45 cm long; stipes dark reddish brown, dull, 3–10 cm ⫻ 0.5–1 mm, 1⁄6–1⁄4 of frond length, glabrous except for basal scales, with narrow wings 0.1 mm wide; blades herbaceous, mostly 20–40 ⫻ 2.5–5 cm, pinnatepinnatifid, pinnatifid at apices, most blades terminating in small scaly buds, linear-lanceolate, tapering proximally, stalked to 1 mm; rachises castaneous, dull, glabrous, adaxially with wings 0.1– 0.2 mm wide; pinnae 22–50 pairs, 1.3–2.5 ⫻ 0.5–1.5 cm, sessile or stalked ⬍ 1 mm, the most developed each with a single obovate or flabellate, nearly free superior basal segment, apices acute to acuminate, margins bicrenately toothed, crenate-serrate towards tips; veins mostly 5–8 pairs per pinna, simple except in basal acroscopic lobe, tips evident adaxially; indument abaxially of scattered, appressed, tan, clavate hairs 0.1–0.2 mm long; sori 2–8 pairs per pinna, on both sides of midveins, 0–3 on superior basal segments; indusia 1.5–3.5 ⫻ 0.4–0.6 mm, margins entire; spores reniform; 2n⫽72 (Chis, var. sessilifolium).

Key to the Mexican Varieties of Asplenium sessilifolium 1. Pinnae lanceolate, acute to acuminate at tips; pinnae Ⳳ deeply incised along acroscopic margins. ............. 72b. var. occidentale. 1. Pinnae oblong, obtuse at tips; pinnae generally shallowly incised except for a single basal segment. ........... 72a. var. sessilifolium.

Distribution. Epiphytic or epipetric in wet forests of Atlantic slope; 50–1200 m. USA (Fla); Mexico; Guat, Bel, Hond, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol, Parag, Arg. Selected Specimens Examined. Chis (Breedlove 48333, CAS; Martı´nez S. 10669, MO; Stevens & Martı´nez 25893, MO, NY, UC). Oax (Croat & Hannon 63285, MO, UC; Mickel 5845, MEXU, NY).Tab (Cowan 3155, MO; Croat 40101, MO, UC; Davidse et al. 29492, MO, UC). Ver (Finck 166, UC; Nee 30012, MO).

72a. Asplenium sessilifolium var. sessilifolium

Fig. 35E.

Asplenium sessilifolium Desv. var. guatemalense Hieron., Bot. Jahrb. Syst. 34: 461. 1904. Syntypes. Guatemala. Lehmann 1480 (B!, frag. NY!; isosyntypes BM!, US!), Heyde & Lux s.n. [Donn. Sm. 4670, 4671] (B!; isosyntype of 4670 at NY!, US); Costa Rica. Wendland 710 (B!).

Differing from var. occidentale by having oblong pinnae, these generally shallowly incised, (except for a single proximal nearly free segment) and with tips obtuse.

124

ASPLENIUM

Distribution. Terrestrial, epipetric on wet rocks, or epiphytic, in wet montane and cloud forests; (900–)1600–3000 m. Mexico; Guat, Hond, Salv, CR, Pan; Col, Ec, Peru, Bol. Also reported from Hispaniola and Venezuela, by Adams (in Davidse et al., 1995: 320), and from Nicaragua by Go´mez (1976: 53); identification of specimens from some of these areas is suspect. Selected Specimens Examined. Chis (Breedlove 32036, DS, MEXU). Gro (Lorea 1642, FCME). Hgo (Pringle 8975, CAS, MO, NY, UC, US). Me´x (Ferna´ndez N. 508, ARIZ, ASU, CHAPA, MEXU). Mich (Soto s.n., FCME). Oax (Wendt et al. 5024, CAS, CHAPA, MO, NY). Pue (Tenorio L. 12453, IEB, MEXU, XAL). Qro (Rubio 2104, IEB, MEXU). SLP (Rzedowski 10059, ENCB). Ver (Purpus 6054, MO, NY, UC, US).

Asplenium sessilifolium, a widespread species in the Neotropics, is intermediate in dissection between A. potosinum (1pinnate) and A. muenchii (2-pinnate-pinnatifid to 3-pinnate). These three species, and also A. semipinnatum, belong to a natural group that may include also the nonproliferous species A. barbaense, A. miradorense, and A. seileri, as well as others. Asplenium sessilifolium, A. muenchii, and A. potosinum are usually proliferous by the production of buds (and ultimately plantlets) from axils of distal pinnae. However, specimens of A. sessilifolium from Costa Rica and Panama generally lack buds. Some specimens from Guatemala have pinnae more dissected than usual (var. guatemalense). 72b. Asplenium sessilifolium var. occidentale A. R. Sm., var. nov. Type. Mexico. Jalisco: San Sebastian, Segundo Arroyo, Mexia 1557 (holotype UC!; isotypes CAS!, MO!, NY!). Fig. 44A. A var. sessilifolio pinnis lanceolatis, acutis vel acuminatis apice, Ⳳ profunde incisis secus marginem acroscopicum differt. (L. occidentalis, western, referring to the distribution of this variety in western Mexico.)

Distribution. Terrestrial or epipetric on mossy boulders, in wet woods and on cliff faces; (700–)1300–2300 m. Mexico. Specimens Examined. Chih (McGill & Reeves R4937, ASU; Spellenberg & Miller 12029, UC). Dgo (Breedlove 44293, CAS, MEXU; Ownbey & Ownbey 1948, UC; Reeder & Reeder 2482, 2486, US, 2502, MEXU, US; Sa´nchez S. 748, 750, MEXU, US). Jal (McVaugh 14034, MEXU, US, 14245, US; Mexia 1397, MO, US, 1557, IEB, US). Me´x (Hinton 3546, NY, 7360, ARIZ, MO, NY, US; Matuda 30029, US). Mich (Langlasse´ 803, MEXU, US). Oax (Calzada 19762, MEXU, 21352, MEXU, UC). Rev (Felger 15828, UC, US; Jarquı´n s.n., MEXU; Mason 1670, CAS, DS, MO, NY, UC, US; Moran 25510, MEXU). Sin (Breedlove 17106, CAS, NY, 35731, CAS, MO; Correll & Correll 28859, LL, US; Gentry 7275, ARIZ, CAS, DS, NY, UC, US, 7276, NY, UC, US; Lehto 24372, ASU; Ownbey & Ownbey 1940, 1948, US; Roberts & Keil 10218, NY). Son (Ferguson et al. s.n., ARIZ).

73. ASPLENIUM SOLEIROLIOIDES Asplenium soleirolioides A. R. Sm., Proc. Calif. Acad. Sci., ser. 4, 40: 212. 1975. Type. Mexico. Chiapas: Municipio La Grandeza, 10 km E of El Porvenir along road from Huixtla to Siltepec, Breedlove 31820 (DS!). Fig. 37L. Roots filamentous, fibrous, not proliferous; rhizomes erect; rhizome scales black, clathrate, 2–4 ⫻ 0.3–0.5 mm, entire; fronds arcuate to decumbent, mostly 4–10 cm long; stipes castaneous, lustrous, 0.4–2 cm ⫻ 0.3–0.4 mm, 1⁄5 of frond length or usually much less, with rare, hair-like, clathrate scales 0.5–1 mm long and sparse clavate hairs 0.3 mm long, not alate, but with two faint adaxial brown ridges; blades thin-herbaceous, 5–9 ⫻ 0.6– 0.8 cm, 1-pinnate, linear, apices subconform or proliferous, buds along rachises in axils of distal pinnae eventually rooting; rachises castaneous, lustrous, with scattered dark-tipped glandular hairs 0.1–0.2 mm, not winged; pinnae round-obovate to subrhomboid, 8–20(–23) pairs, 2–4 cm ⫻ 0.5–2 mm, articulate, usually alternate, apices rounded, margins entire; veins simple, 2–3 per pinna, tips evident adaxially; indument abaxially of sparse, appressed whitish to tan, clavate hairs 0.1–0.2 mm long; sori 1 or 2 per pinna, the larger (and sometimes only) one along the basiscopic side of midveins; indusia whitish, 1–2 ⫻ 0.5–0.8 mm, margins entire or erose; spores reniform.

ASPLENIUM SPHAEROSPORUM

125

sori single on the ultimate segments; indusia to 1–2.5 ⫻ 0.3–0.4 mm, margins entire; spores reniform.

Distribution. On shaded limestone rocks, in pine and fir forests, by streams; 2100–2900 m. Mexico. Specimens Examined. Chis (only the type). Gro (Lorea 3130, FCME, NY; Reveal et al. 4251, US; Vela´zquez 822, FCME). Hgo (Medina-Cota & Barrios 3730, CAS, NY). Jal (McVaugh 26139, MEXU, MICH). Mich (Carranza & Zamudio 4227, IEB, UC). Oax (Mickel 5386, 7034, NY; Smith 509, UC). Qro (Dı´az B. 4930, IEB, XAL).

Several other small spleenworts have a similar appearance. Among them are Asplenium stolonipes, which is also small in stature, has castaneous lustrous stipes, small indusia, and proliferous from buds in the axils of the proximal pinnae; it differs from A. soleirolioides in having smaller occluded rhizome scales 0.8–1 mm long, shorter fronds 1.7–4 cm long, and occurring on granitic rocks. Asplenium harrisii has similar pinnae except they are deeply lobed and the stipes and rachises are alate and green. Asplenium clutei Gilbert (isotype Clute 118, NY!), from Jamaica and Hispaniola, has lobed to barely 1-pinnate pinnae with marginal glands and rhizome scales with glandular margins.

Distribution. Terrestrial in montane forests; 900–1800 m. Mexico; Guat. Specimens Examined. Chis (Heath & Long 757, CAS; Matuda 2637, ARIZ, F, K, MEXU, MO, NY, UC, US, 4173, LL, MEXU, NY, US).

This species is most closely related to A. excelsum Lellinger from Nicaragua, Costa Rica, and Panama. In Mexico, the closest relative is probably A. achilleifolium, from which it differs in having more dissected blades.

75. ASPLENIUM SPHAEROSPORUM Asplenium sphaerosporum A. R. Sm., Amer. Fern J. 70: 17. 1980. Type. Mexico. Chiapas: SE side of Cerro Baul, 16 km NW of Rizo de Oro, Breedlove 21805 (DS!; isotypes CAS!, NY!). Fig. 36C.

Asplenium solmsii Baker ex Hemsl., Biol. Cent.-Amer., Bot. 3: 639. 1885. Type. Guatemala. “Cuesta de Atitla´n,” Bernoulli & Cario 317 (K, not found). Figs. 40C, D.

Differs from A. auritum by: fronds (20–)30–70 cm long; stipes ⁄3–1⁄2 of frond length; blades deltate-lanceolate, apices acuminate, 2-pinnate proximally, pinnate-pinnatifid distally, (5–)12–18 cm wide; pinnae (12–)25 or more pairs, segments obovate, mostly adnate; spores globose, large, dark brown to blackish, 32 per sporangium.

Roots coarse, fibrous, not proliferous; rhizomes erect; rhizome scales dark brown to blackish, clathrate, but lumina nearly occluded, 6–8 ⫻ 0.3–0.6 mm; fronds clumped, to ca. 50 cm long; stipes brownish gray, dull, to 20 cm ⫻ 3 mm, ca. 1⁄3 of frond length, Ⳳ glabrous, with inconspicuous greenish wings or ridges 0.1 mm wide; blades nearly 3-pinnate, to ca. 35 ⫻ 15 cm; rachises gray, dull, seemingly glabrous, with narrow greenish wings 0.2– 0.3 mm wide, or with a few minute clathrate scales in axils of pinnae; pinnae lanceolate-attenuate, to ca. 15 pairs, 5–8 ⫻ 2–2.5 cm, stalked 1–3 mm, inequilateral, basiscopic pinnules somewhat shorter and more oblique than acroscopic ones, pinnules decurrent on costae, deeply pinnatifid, the largest 24 ⫻ 5–6 mm, the lobes ca. 1 mm wide, oblique, often falcate, apices acute or sometimes bifid; veins only a midvein on the ultimate segments, readily visible on both blade surfaces, tips especially evident adaxially; indument abaxially of sparse, appressed, clavate hairs 0.1 mm;

Distribution. Epiphytic, or rarely terrestrial, in pine-oakLiquidambar forests; 650–2700 m. Mexico; Guat, Hond, Nic,

74. ASPLENIUM SOLMSII

1

126

ASPLENIUM

CR. Also reported from Venezuela by Adams (in Davidse et al., 1995: 321), but specimens not seen from there. Selected Specimens Examined. Chis (Breedlove 64328, CAS, 71365, CAS). Col (Sanders 11707, MEXU, UC). Gro (Mexia 8984, ARIZ, CAS, NY, UC, US). Jal (Dı´az L. 5459, UC). Me´x (Moore & Cetto 5460, US). Mor (Lyonnet 775, MO, US). Oax (Mickel 4775, NY, UC). Qro (Carranza 1051, ENCB, IEB, MEXU). Ver (Ventura A. 9629, ASU, IEB).

See discussion under A. auritum. A possible synonym is Asplenium auritum Sw. var. bipinnatisectum Mett., Abh. Senckenberg. Naturf. Ges. 3: 147. 1859. Lectotype (chosen by Morton & Lellinger, 1966: 20). Mexico. Schiede & Deppe [773] (B-photo US!). This has usually been placed under A. lacerum (e.g., by Mickel & Beitel, 1988: 61), here treated as a synonym of A. cuspidatum, or under A. auritum var. bipinnatifidum Kunze (e.g., by Morton & Lellinger, 1966: 20). Breedlove 24054 (DS, NY), from Chiapas, appears to be a hybrid involving A. sphaerosporum and some other species, possibly A. cuspidatum; the spores are malformed.

76. ASPLENIUM STOLONIPES Asplenium stolonipes Mickel & Beitel, Mem. New York Bot. Gard. 46: 72. 1988. Type. Mexico. Oaxaca: Distrito Ixtla´n, trail N of San Pedro Nolasco to the Llano Verde, at high point above the Llano Verde (Las Cruces), Mickel 5376 (NY!; isotype UC!). Figs. 37K. Roots filamentous, fibrous, not proliferous; rhizomes suberect to erect; rhizome scales black, clathrate, but lumina almost totally occluded, 0.8–1.5 ⫻ 0.2–0.4 mm, entire; fronds clumped, 2–5(– 7) cm long; stipes atropurpureous, lustrous, 4–35 ⫻ 0.2–0.3 mm, 1 ⁄3–1⁄2 of frond length, with sparse hair-like scales 0.5–0.8 mm long, not winged; blades 1-pinnate, 1.5–4(–6) ⫻ 0.5–0.8 cm, often bearing a proliferous bud at the bases of the blades (from each of which may arise a new plant); rachises atropurpureous, lustrous, with scattered dark-tipped glandular hairs 0.1 mm long, lacking wings; pinnae rhomboidal-trapezoidal, imbricate, 5–12 pairs, 2–3(–4) ⫻ 1.5–2 mm, articulate, rachis color not extending into pinna stalks, apices obtuse or rounded, margins entire to undulate on acroscopic sides; veins ca. 3 per pinna, tips evident adaxially, often producing lime dots; indument abaxially of whitish to tan glandular hairs 0.1 mm long; sori 1 or 2 per pinna, on both sides of midveins, or if only one then on the basiscopic side parallel to margin; indusia whitish, 1–2 ⫻ 0.5–0.8 mm, margins entire; spores reniform.

Distribution. Epipetric on granitic rocks in cloud forest of pine, oak, and mixed hardwoods; 2450 m. Mexico. Known only from the type collection. Asplenium stolonipes forms colonies comprising a mass of miniature plants, mostly arising from buds on horizontal stipes. Mickel and Beitel (1988) compared it to A. fragile C. Presl (⫽ A. peruvianum Desv., according to Tryon & Stolze, 1993) and A. tenue C. Presl (⫽ A. triphyllum C. Presl, according to Tryon & Stolze, 1993), both from South America. Asplenium peruvianum differs in having green rachises with prominent wings (0.5 mm) and non-occluded clathrate scales, while A. triphyllum has trifoliate or more dissected pinnae. The South American species are not closely related to A. stolonipes but do agree in the tendency to produce proliferous buds in the axils of proximal pinnae, a feature sometimes also found in the more widespread A. monanthes. This species is actually most like a miniature form of A. monanthes, resembling it in the form of the pinnae and occluded rhizome scales. The combination of small size, dark rachises, proliferous buds at bases of blades, prominent hydathodes, entire indusia, and glandular hairs on the rachises and blades abaxially separate this from others with lustrous, dark rachises. Another close relative is probably A. soleirolioides, which differs in having flagelliform blades and with buds on the distal part of the rachises.

77. ASPLENIUM TENERRIMUM Asplenium tenerrimum Mett. ex Kuhn, Linnaea 36: 97. 1869. Type. Venezuela. [Aragua:] near La Victoria, Fendler 368 (B; isotypes K, MO!, NY!, US!). Fig. 37J. Asplenium cuspidatum Lam. var. tenerrimum (Mett. ex Kuhn) C. V. Morton & Lellinger, Mem. New York Bot. Gard. 15: 29. 1966. Asplenium fournieri Kuhn ex E. Fourn., Bull. Soc. Bot. France 17: 237. 1870. Type. Nicaragua. Isla Omotepe [Omepepe], Le´vy 1159 (P!–2 sheets, photo NY!, UC!; isotypes B!, US!).

Roots filamentous, fibrous, not proliferous; rhizomes suberect; rhizome scales black, clathrate, hair-like, 1.5–2 ⫻ 0.1–0.3 mm, entire; fronds clumped, 4–12 cm long; stipes atropurpureous at bases, lustrous, bicolorous above bases, but often blackish abax-

ASPLENIUM TRYONII

ially, 1–5 cm ⫻ 0.2–0.6 mm, 1⁄3 of frond length, glabrous, with greenish wings 0.1 mm wide; blades membranaceous, 3–5 ⫻ 2– 3.5 cm, 3–4-pinnate, deltate, widest proximally, apices pinnatifid, not proliferous; rachises green or bicolorous for 1⁄2 their length, glabrous, with wings 0.1 mm wide; pinnae deltate, 3–5 pairs, 2–3 ⫻ 0.6–1.5 cm, 2–3-pinnate in larger fronds, ultimate segments linear or narrowly obovate, 0.5–0.8 mm wide, toothed near tips; veins simple, readily visible on both sides, but not ending in widened tips adaxially; indument abaxially of inconspicuous, tan, clavate hairs 0.1 mm long; sori 1 per ultimate segment, at bases of segments near vein fork; indusia 0.8–1.2 ⫻ 0.3–0.4 mm, margins erose; spores reniform.

Distribution. Terrestrial and epipetric along river banks and road banks; 50–900 m. Mexico; Hond, Nic; Col, Ven, Ec. Also cited from El Salvador by Adams (in Davidse et al., 1995: 322). Selected Specimens Examined. Chis (Breedlove 37593, DS; Ramos s.n., MEXU). Gro (Lorea 2579, FCME). Jal (Lott 1386, UC). Me´x (Tejero-Dı´ez 3021, IZTA). Mich (Nakamura 123, IEB). Nay (Ferris 5360, US; Wright 1360, US). Oax (Mickel 6900, NY). Sin (Breedlove 35642, CAS, MO; Mexia 257, CAS, MO, UC).

This small, 3(–4)-pinnate species is similar in dissection and size to Asplenium delicatulum but differs in many details. Asplenium delicatulum has stipes green proximally, entire indusia, sori that are located midway on the ultimate segments, and shorter brown, retiform rhizome scales 0.5–0.8 mm long. It spreads by means of slender, stolon-like roots, which are absent in A. tenerrimum.

78. ASPLENIUM TRICHOMANES Asplenium trichomanes L., Sp. Pl. 2: 1080. 1753. Lectotype (chosen by Viane in Jonsell & Jarvis, 1994: 150). Europe. Herb. Burser XX: 14 (UPS, microfiche UC!). See also Reichstein, 1981; Lovis, 1964). Figs. 42C, D. Roots filamentous, wiry, not proliferous; rhizomes shortcreeping to erect; rhizome scales blackish, clathrate or with central black band and clathrate edges, lumina very small, 1.5–2.5 ⫻ 0.1– 0.3 mm, entire; fronds clumped, 7–20 cm long; stipes casta-

127

neous to atropurpureous, lustrous, 1–4 cm ⫻ 0.5–0.8 mm, 1⁄6– 1⁄4 the blade length, glabrous, with tan inconspicuous adaxial wings ⭐ 0.1 mm wide; blades chartaceous, narrowly linearoblong, 1-pinnate, 6–15 cm ⫻ 4–10 mm, bases slightly narrowed, apices confluent, not proliferous; rachises castaneous, lustrous, glabrous, adaxially with tan wings 0.1–0.2 mm wide; pinnae short-oblong, 10–20(–30) pairs, 4–7 ⫻ 1–4 mm, stalked to 1 mm, ultimately articulate, dimidiate, apices rounded, margins entire to slightly crenulate, sometimes revolute; veins 1-forked, tips not evident adaxially; indument abaxially absent, blades glabrous; sori 2–5 pairs per pinna, on both sides of midveins, occasionally diplazioid; indusia 0.5–1.5 ⫻ 0.2–0.4 mm, margins entire to suberose; spores reniform; 2n⫽72, 144 (USA).

Distribution. In sheltered crevices of ledges and cliffs; ca. 2500 m. Can, USA (Ga, w to Calif and Alaska); Mexico; Eurasia. Specimen Examined. Chih (LeSueur 1130, US).

Presumably, the Mexican specimens are subsp. trichomanes, the diploid cytotype (thus agreeing with specimens from southwestern United States). Tetraploid plants, subsp. quadrivalens D. E. Meyer, are generally more northern in distribution and grow on calcareous (vs. acidic) substrates. Other subspecies, with hexaploid number of chromosomes, are known from Madeira and New Zealand. Asplenium trichomanes is most similar to A. palmeri in Mexico, differing from that in the nonproliferous blades.

79. ASPLENIUM TRYONII Asplenium tryonii Correll, Wrightia 2: 200, f. 36. 1961. Type. Mexico. Chihuahua: Tecolote, 10 mi W of Chinatu´, Correll & Gentry 23000 (LL; isotypes GH!, US!). Figs. 42K–M. Roots thin, wiry, not proliferous; rhizomes suberect to erect; rhizome scales dark brown, clathrate, to 3 ⫻ 0.4 mm, entire; fronds clumped, 4–7 cm long; stipes dull reddish brown, lustrous, ca. 1.5 cm ⫻ 0.5 mm, 1⁄5–1⁄2 of frond length, with hair-like scales, not winged; blades thick-herbaceous, narrowly lanceolate, 1pinnate, bearing buds and rooting at bases of terminal segments or from adaxial surfaces of distal pinnae; rachises dull reddish brown, lustrous, sparsely fibrillose, not winged; pinnae oblong

128

ASPLENIUM

to trullate, essentially opposite, 7–12 pairs, proximal pinnae only slightly reduced, to 5 ⫻ 3.5 mm, sessile, bases broadly cuneate, apices obtuse, margins crenulate or slightly repand on both acroscopic and basiscopic margins, as well as at tips; veins obscure, tips evident adaxially; indument absent, blades glabrous; sori 2– 4 per pinna, on both sides of midveins; indusia 1–2 ⫻ 0.8–1 mm, margins somewhat erose-crenulate; spores not seen.

margins shallowly and simply crenate, crenations often more than 5 mm broad, apices acuminate to long-attenuate; veins 2-forked, faintly visible on both sides, tips prominent and expanded adaxially; indument of tan appressed, clavate hairs 0.1– 0.2 mm long; sori 7–15 pairs per pinna, on both sides of midveins; indusia whitish, 5–12 ⫻ 0.3–0.8 mm, margins entire; spores reniform.

Distribution. In moss on shaded north-facing ledges; 1900– 2200 m. Mexico.

Distribution. Terrestrial or epiphytic in wet montane forests; 200–1400(–2700) m. Mexico; Guat.

Specimen Examined. Chih (Correll & Gentry 23213, LL).

Other Mexican species in the Asplenium trichomanes group that bear blade proliferations are A. palmeri, A. stolonipes, A. soleirolioides, A. polyphyllum, and, rarely, A. monanthes. Asplenium palmeri and A. soleirolioides differ from A. tryonii in having flagelliform apices, generally longer fronds, and many more pinna pairs. Asplenium stolonipes and A. monanthes have buds only at the base of proximal pinnae. Asplenium polyphyllum, with buds at the base of distal pinnae, is a much larger species, with many more pinna pairs and larger pinnae. Asplenium tryonii is one of the rarest Mexican spleenworts, still known only from the type an one additional collection.

80. ASPLENIUM TUERCKHEIMII Asplenium tuerckheimii Maxon, Contr. U.S. Natl. Herb. 13: 15 1909. Type. Guatemala. Baja Verapaz: Panza´l, Tu¨rckheim II 1677 (US!; isotype BR). Fig. 34C. Roots relatively coarse, fibrous, not proliferous; rhizomes erect or suberect; rhizome scales dark brown to blackish, clathrate, lumina occluded, 2–4 ⫻ 0.5–1 mm, entire; fronds clumped, relatively few (2–4), 20–40 cm long; stipes stramineous to green, dull, 10–18 cm ⫻ 1–2 mm, 1⁄2–2⁄3 of frond length, with scattered, clavate hairs 0.1–0.3 mm long, adaxially with narrow wings alongside groove; blades herbaceous, 8–20 ⫻ 7–12 cm, 1-pinnate, ovate, each blade with a conform terminal pinna, not proliferous; rachises greenish to tan, dull, glabrous, adaxially with green wings 0.2–0.3 mm wide; pinnae lanceolate, ascending, often falcate, 3–6 lateral pairs, mostly 8–15 ⫻ 1–2 cm, sessile or stalked 0–2 mm, equilateral, bases cuneate, acroscopically exauriculate,

Selected Specimens Examined. Chis (Breedlove 22299, 31540, DS, 32550, DS, NY, MEXU; Carlson 1779, US; Ton 5749, MO). Oax (Herna´ndez G. 1934, NY; Mickel 7300, ENCB, NY, UC, US). Ver ( Finck 41, 135, US; Purpus 6432, UC, 4349, UC, US).

Asplenium tuerckheimii differs from A. riparium in its clumped fronds, erect rhizomes, blackish rhizome scales with occluded lumina, fewer pinnae per blade (3–6 pairs), equilateral, cuneate pinna bases, more sori (7–15 pairs) per pinna, and whitish indusia. See discussion under A. feei for comparison with that species. Several specimens from northern Chiapas grade towards A. abscissum and are not easy to distinguish from that species (Smith, 1981: 54). Some of these specimens have discrete terminal pinnae, some do not. Additional studies are needed on the relationships and circumscription of these two species.

81. ASPLENIUM ULTIMUM Asplenium ultimum A. R. Sm., sp. nov. Type. Mexico. Nuevo Leo´n: Mpio. Galeana, Cerro El Gallo, 2350 m, Hinton 20142 (holotype TEX!; isotype MEXU!). Figs. 41C, D. A. exiguo gemmis apicibus pinnarum et laminarum carentibus, pinnis infimis pinnas contiguas aequantibus, stipitibus longioribus (plerumque 2.5–4 cm longis) et viridibus praeter bases differt. (L. ultimus, last; this species was the last to be recognized and characterized during the preparation of this treatment of Asplenium.)

Roots filamentous, fibrous, not proliferous; rhizomes not known, probably suberect; rhizome scales not seen, but stipe base scales black, clathrate, 1.5–3 ⫻ 0.2–0.3 mm, entire; fronds presumably clumped, ca. 6–8 cm long; stipes castaneous and lustrous

ASPLENIUM VENTURAE

129

in the proximal 1 cm, greenish distally, dull, (1.5–)2.5–4 cm ⫻ 0.5–1 mm, 1⁄3–2⁄5 of frond length, bearing scattered hair-like scales, not winged; blades mostly 3–4.5 ⫻ 1–2 cm, lanceolate, pinnate-pinnatifid, proximal pinnae not or only slightly reduced, the apices neither proliferous nor flagelliform; rachises brownish, with sparse hair-like scales, not winged; pinnae oblong, 6–9 pairs, 5–10 ⫻ 3–5 mm, pinnatifid to pinnatisect, apices lacking proliferous buds, pinnules oblong to flabellate, margins subentire to toothed; veins obscure, 1–2-forked in larger pinnules, tips not evident adaxially; indument abaxially of scattered red-brown, appressed, clavate hairs to 0.2 mm long; sori to 5 per pinna, on both sides of midveins, mostly one per pinnule; indusia 1–1.8 ⫻ 0.5 mm, margins suberose; spores not seen. Distribution. Terrestrial in wet montane forest; 200–1250(– 2500) m. Mexico; Bel (Holst 5321, UC), Hond, Nic, CR, Pan; Braz. Also reported from Guatemala, Hispaniola, Puerto Rico, Colombia, Venezuela, Ecuador, Peru, and Bolivia by Adams (in Davidse et al., 1995: 323), but his circumscription of the species appears to be broader. Selected Specimens Examined. Chis (Breedlove 33089, 35174, DS; Purpus 6733, PH, UC, US, 6745, MO, NY, UC, US). Oax (Herna´ndez G. 1182, CAS, CHAPA, ENCB, MO, 1472, 1545, CAS, CHAPA, MO, 1672, 1905, CHAPA; Mickel 6404, ENCB, NY, UC, US; Wendt 3632, CAS, CHAPA). Ver (Wendt et al. 3219, CHAPA, ENCB, MEXU, NY).

Distribution. On limestone rock slide, forming colonies; 2350 m. Mexico. Known only from the type collection. This species is probably most closely related to A. exiguum, but differs by having longer stipes (mostly 2.5–4 cm long), greenish stipes (except proximal ca. 1 cm darkened), without reduced proximal pinnae (which are about as long as the longest pinnae), and absence of buds at the tips of the pinnae and blades. The sori are adjacent and at a narrow angle to the costae, except for single sori along the midribs of the basal acroscopic segment of each pinna.

82. ASPLENIUM UNISERIALE Asplenium uniseriale Raddi, Opusc. Sci. 3: 291. 1819. Fig. 38E. Asplenium radicans L. var. uniseriale (Raddi) L. D. Go´mez, Brenesia 8: 53. 1976. Type. Brazil. Raddi s.n. (presumably FI). Asplenium amabile Liebm., Mexic. Bregn. 251 (reprint 99). 1849. Lectotype (chosen by Smith, 1981: 54). Mexico. Oaxaca: Between La Galera and Pochutla, Liebmann s.n. [Fl. Mex. 352] (C!; isotypes B!, K!, US!).

Differs from Asplenium radicans by: rhizome scales clathrate with yellow or brown lumina and sparse, short teeth, 4–5 ⫻ (0.8–) 1–1.5 mm; blades 3-pinnate, basal acroscopic pinnules cut to apices into linear-oblong to obovate segments 4 ⫻ 1 mm; sori 1 per ultimate segment in the most finely divided forms; indusia 2–4 mm long, margins fimbriate, erose, or suberose.

This fully 3-pinnate member of the A. radicans complex is variable in the indusial margins and rhizome scale width (0.8– 1.5 mm). Specimens are similar to A. cristatum in dissection, and some have previously been determined as that species. See A. radicans for more discussion of this complex.

83. ASPLENIUM VENTURAE Asplenium venturae A. R. Sm., sp. nov. Type. Mexico. Veracruz: Mpio. Acajete, Acajete, Ventura A. 11176 (holotype IEB!; isotypes ENCB!, XAL!). Fig. 40A. Ab A. cristato pinnulis tantum dentatis apice, paleis rhizomatum ovatis (vs. lanceatis) apice minus acuminatis, margine dentatis dentibus 0.1–0.2 mm longis, luminibus cellularum fuscatis parvioribus et occlusioribus differt. (Honoring A. Ventura A., avid plant collector in southern Mexico and especially in the state of Veracruz.)

Roots coarse, fibrous, not proliferous; rhizomes erect; rhizome scales blackish, obscurely clathrate, lumina occluded, 2–3 ⫻ 0.5–1 mm, with marginal teeth 0.1–0.2 mm long; fronds clumped, to ca. 30 cm long; stipes dull brown, 5–12 cm ⫻ 1 mm, ca. 1⁄3–2⁄3 of frond length, narrowly green-winged adaxially, with glandtipped hairs 0.2–0.3 mm long especially abaxially, glabrescent, adaxially with wings 0.1 mm wide; blades membranaceous, 2pinnate, 15–20 ⫻ 4–8 cm, lanceolate to deltate-oblong, apices acuminate, pinnatifid, nonproliferous; rachises brownish, glabrous, with narrow wings 0.3 mm wide; pinnae lanceate, spreading, 12–16 pairs, to 4 ⫻ 1.7 cm, sessile, with up to ca. 6 pairs of pinnules per pinna, proximal acroscopic pinnule of each pinna overlapping the rachis, usually with 1 or 2 pairs of slightly re-

130

ASPLENIUM

duced proximal pinnae, pinnules rhombic to oblong, tapering to cuneate bases, 4–12 mm long, toothed toward the tips, proximal pinnules slightly lobed as well as toothed; veins mostly 2– 4 pairs per pinnule, tips evident adaxially; indument abaxially of appressed to spreading gland-tipped reddish brown clavate hairs 0.1–0.2 mm; sori 1–4 per pinnule, on both sides of midveins; indusia 1–2 ⫻ 0.3–0.5 mm, margins entire; spores reniform.

pered, apices acute, not rooting; rachises castaneous, lustrous, with scattered hair-like scales to 1 mm and a few gland-tipped hairs to 0.3 mm, not winged; pinnae oblong, 15–30 pairs, 5–13 ⫻ 3–6 mm, sessile, bases truncate to cuneate, apices obtuse, acroscopic and basiscopic margins shallowly lobed or toothed; veins simple, obscure, tips evident adaxially; indument abaxially of whitish appressed clavate hairs 0.1 mm long; sori 2–6 pairs per pinna, on both sides of midveins; indusia ca. 1–1.5 ⫻ 0.2– 0.5 mm, margins entire; spores reniform, 64 per sporangium.

Distribution. Terrestrial in forests and thickets; 1850 m. Known only from the type collection. This species is most similar to A. cristatum, but differs from that in having pinnules that are merely toothed at the tips and in having rhizome scales more ovate (vs. lanceate), with a less acuminate tips and smaller, more occluded cells, also with marginal teeth 0.1–0.2 mm long. In pinnule shape and toothing, this species is more similar to the 2-pinnate form of A. flabellulatum, but the stipes are dull brown, not lustrous, and the blades are not flagelliform at the tips. The spores appear to be well formed on the type. A somewhat similar specimen from the same locality, Ventura A. 11171 (XAL), has larger, pinnatifid, acroscopic pinnules, reduced proximal pinnae, and narrower, more attenuate blades; the sporangia appear malformed. It could be a hybrid involving A. venturae, possibly with A. sessilifolium, under which it was first identified.

84. ASPLENIUM VESPERTINUM

Distribution. Epipetric on moist, shaded, canyon walls and at bases of boulders; 0–350 m. USA (s Calif); Mexico. Specimens Examined. BCN (Jones 3749, CAS, DS, MO, NY, UC, US; Moran 30526, ARIZ, CAS, ENCB, MEXU, MO, NY).

This species differs from related species of the A. trichomanes group, and in particular from A. trichomanes itself, by having the fertile veins (basal acroscopic ones excepted) usually simple, pinnae mostly 6–10 mm long, with length 2–3 times width, and by the dentate or shallowly lobed acroscopic and often basiscopic pinna margins. Other Mexican species of this group with lobed or incised pinnae include A. blepharodes, A. fibrillosum, and A. formosum. The first two of these both have ciliate indusia; Asplenium formosum has even more deeply incised, almost lacerate pinnae.

85. ASPLENIUM VIRILLAE

Asplenium vespertinum Maxon, Bull. Torrey Bot. Club 27: 197. 1900. Fig. 39F.

Asplenium virillae Christ, Bull. Herb. Boissier, se´r. 2, 4: 1090. 1904. Type. Costa Rica. Rı´o Virilla, Alfaro s.n. [Herb. Costar. 16448] (P!; probable isotype US!). Figs. 43C, D.

Asplenium trichomanes L. var. vespertinum (Maxon) Jepson, Man. Fl. Pl. Calif. 35. 1923. Type. U.S.A. California: San Diego Co., San Miguel Mt., near National City, Kimball s.n. (US!).

Asplenium alfredii Rosenst., Repert. Spec. Nov. Regni Veg. 22: 8. 1925. Type. Costa Rica. Candelaria, A. & C. Brade 424 (S, photo BM!; isotypes K!, NY!, UC!, US!).

Roots filamentous, wiry, not proliferous; rhizomes shortcreeping; rhizome scales blackish or with lighter margins, 2–3 ⫻ 0.2–0.4 mm, entire or denticulate distally; fronds clumped, mostly 10–15(–20) cm long; stipes castaneous, lustrous, 2–5 cm ⫻ 0.5–1 mm, 1⁄5–2⁄5 of frond length, sometimes abruptly curved at bases, glabrous, not winged; blades herbaceous, linear, 1pinnate, mostly 10–15 ⫻ 1–2.5 cm, glabrous, bases gradually ta-

Roots thin, fibrous, not proliferous; rhizomes erect; rhizome scales 2–3 ⫻ 0.4–0.5 mm; fronds clumped, 4–12 cm long; stipes darkened at bases, stramineous to greenish distally, dull, 1–6 cm ⫻ 0.3–0.8 mm, 1⁄3–1⁄2 of frond length, glabrous, not or obscurely alate adaxially; blades herbaceous, with (1–)2–5 pairs of lateral pinnae, stalked 1–2 mm, terminal segments broadly hastatedeltate; rachises greenish, dull, glabrous, lacking wings or these

ASPLENIUM YELAGAGENSE

to ca. 0.1 mm wide; pinnae oblong, 1–5 pairs, 1–2 ⫻ 0.6–1 cm, less than 2 times longer than wide, sessile or stalked to 2 mm, obtuse at apices, margins entire to dentate to weakly lobed acroscopically, entire or nearly so basiscopically; veins mostly 1forked, readily visible on both sides, tips prominent and expanded adaxially; indument abaxially of scattered, tan, appressed clavate hairs 0.1 mm long; sori 2–5 pairs per pinna, on both sides of midveins; indusia tan, 2–4 mm, margins entire; spores reniform.

Distribution. Terrestrial or epipetric, in montane rain forests; 1350–1800 m. Mexico; Salv, CR; Col. Also reported from Guatemala, Honduras, Panama, and Venezuela by Adams (in Davidse et al., 1995: 323), but his circumscription of the species appears to differ from ours. Go´mez (1976: 53) reported this species from Nicaragua. Specimens Examined. Chis (Breedlove 21798, 25070, DS, 56943, CAS).

This species is difficult to distinguish from small, precociously fertile plants of A. abscissum (which see). Asplenium virillae differs by the smaller fronds with fewer pinnae and lateral pinnae rounded or slightly oblong (vs. rhombic, more acute or acuminate at the tips, and more strongly inequilateral in A. abscissum). In addition, specimens of A. virillae seem to dry a much darker green. Both species often occur on wet rocks, and both can become fertile at very small size, the fertile fronds sometimes less than 5 cm long.

86. ASPLENIUM YELAGAGENSE Asplenium yelagagense Mickel & Beitel, Mem. New York Bot. Gard. 46: 74. 1988. Type. Mexico. Oaxaca: Distrito Villa Alta, valley of the Yelagago River, ca. 20 mi NE of Villa Alta, Mickel 1058 (NY!; isotypes NY! UC!). Fig. 34E. Roots fibrous, not proliferous; rhizomes suberect; rhizome scales black, clathrate, 3–4 ⫻ 0.5–0.8 mm, entire; fronds clumped, 20–35 cm long; stipes castaneous to atropurpureous, lustrous, 6– 13 cm ⫻ 1–1.5 mm, 1⁄3–2⁄5 of frond length, glabrous, not winged; blades thick-herbaceous, 15–25 ⫻ 6–10 cm, lanceolate, 1-pinnate,

131

not reduced proximally, apices flagelliform, proliferous; rachises castaneous, lustrous, glabrous, adaxially with tan wings ca. 0.1 mm wide; pinnae lanceolate, 10–15 pairs, 3.5–6 ⫻ 0.8–1 cm (excluding basal auricle), auriculate acroscopically, narrowly cuneate and excised basiscopically, apices acute, margins coarsely serrate with sinuses 0.5–1 mm deep; veins mostly simple, except for acroscopic basal vein, obscure, tips evident adaxially; indument abaxially of scattered, tan, appressed, clavate hairs 0.1–0.2 mm long; sori 7–11 pairs per pinna, on both sides of midveins, each pinna with 1 or 2 diplazioid sori on basal acroscopic vein; indusia 6–9 ⫻ 0.3–0.4 mm, becoming strongly reflexed, margins ciliolate with hairs 0.1 mm long; spores reniform.

Distribution. Terrestrial, in wet montane forests on the Atlantic slope; 1050–1200 m. Known only from the type collection. This species is closely related to A. cirrhatum and similar to it in having proliferous, flagelliform tips and 1-pinnate blades. Asplenium yelagagense is distinct in its larger, dark green, strongly auriculate pinnae 3.5–6 cm long, with margins coarsely serrate (sinuses 0.5–1 mm deep), and also by the longer (6–9 mm long) and more numerous sori (7–11 pairs), some diplazioid on the auricles. More study is needed to establish the relationship between these two species and other 1-pinnate taxa of this complex, including A. acutiserratum (Hieron.) Mickel of Trinidad, A. macrurum Mickel & Stolze of Ecuador, and especially A. karstenianum Klotzsch and A. galipense Hieron. of Venezuela. Unplaced Names Asplenium coriaceum Fe´e, Me´m. Foug. 5: 193. 1852, non Desv., 1827. A. pycnophyllum T. Moore, Index Fil. 159. 1859. Type. Mexico. Galeotti s.n. (RB). Asplenium auritum Sw. var. dissectum E. Fourn., Mexic. Pl. 1: 107. 1872. Syntypes. Mexico. Veracruz: S. Cristo´bal, Botteri 1428; S. Andre´s, Botteri 1429 (both Herb. va. Heurck, presumably BR). Excluded Species Asplenium lividum Mett. ex Kuhn, Linnaea 36: 100. 1869. Type. Venezuela. Aragua: San Carlos River valley, Fendler 156 (B; isotypes F, MO!, US!). This was attributed to Chiapas by Adams (in Davidse et al., 1995) on the basis of Ghiesbreght s.n. (BM), which we have not seen. Otherwise, the species has been reported from Venezuela,

132

ASTROLEPIS

Peru (Tryon & Stolze, 1993), and Africa (Burrows, 1990; Jacobsen, 1983; Schelpe & Anthony, 1986). It is very closely related to A. praemorsum, differing by the less dissected blades (merely pinnate-pinnatifid), fewer scales on the blades, and the scales without a long-attenuate tip. We doubt the conspecificity of the Ghiesbreght collection with the Venezuelan type, and the distinctness of the species is also open to question. Therefore, it is here excluded from the Mexican flora.

1 5. A S T R O L E PI S Astrolepis D. M. Benham & Windham, Amer. Fern J. 82: 55. 1992. Type: Astrolepis sinuata (Lag. ex Sw.) D. M. Benham & Windham [⬅ Acrostichum sinuatum Lag. ex Sw.] Rhizomes compact, horizontal to erect, occasionally branched; rhizome scales stramineous to castaneous, concolorous to weakly bicolorous, linear, ciliate to entire; fronds monomorphic, clumped, 7–80 cm long; stipes 1⁄6–1⁄3 the frond length, castaneous to stramineous, terete, sparsely to densely clothed with white, linear-lanceolate to deeply dissected ciliolate scales; blades linear to linear-oblong, pinnate to pinnate-pinnatifid, coriaceous, abaxially clothed with imbricate, lanceolate to ovate, ciliate scales with underlying scurf of substellate scales, adaxially sparsely to moderately clothed with substellate to long-ciliate scales, often

glabrescent, dull; pinnae 15–42 pairs, short-stalked to subsessile, articulate, ovate to oblong or narrowly deltate, cordate to truncate or rounded at bases, coriaceous; pinna margins undifferentiated, not recurved; veins obscure, free, pinnate; sporangia along distal 1⁄3 of veins; spores tetrahedral, dark brown, rugose; x⫽29. A genus of five species, all occurring in Mexico, and perhaps some additional undescribed species, epipetric or terrestrial in relatively dry regions, California and Oklahoma south to Argentina, disjunct in Georgia and Hispaniola. Previously, the species of Astrolepis have been placed in either Notholaena or Cheilanthes. Benham and Windham (1992) have shown Astrolepis to be a distinct group with a chromosome base number of 29; it is separated from related genera by its pinnate to pinnate-pinnatifid fronds with many pinna pairs, two stipe bundles, imbricate scales abaxially, and substellate to ciliate scales on the adaxial blade surfaces. Molecular data (Gastony & Rollo, 1995) show that Astrolepis is sister to Pellaea and Argyrochosma. References Benham, D. M. 1989. A biosystematic revision of the fern genus Astrolepis (Adiantaceae). Ph.D. dissertation. Northern Arizona University; Benham, D. M. 1992. Additional taxa in Astrolepis. Amer. Fern J. 82: 59–62; Benham, D. M. & M. D. Windham. 1992. Generic affinities of the star-scaled cloak ferns. Amer. Fern J. 82: 47–58; Hevly, R. H. 1965. Studies of the sinuous cloak fern (Notholaena sinuata) complex. J. Arizona Acad. Sci. 3: 205–208; Tryon, R. M. 1956. A revision of the American species of Notholaena. Contr. Gray Herb. 179: 1–106.

Key to the Mexican Species of Astrolepis 1. Pinnae deeply lobed about halfway to the costae. 2. Adaxial laminar scales substellate, ca. 0.5 mm long, the scale body 2–3 cells wide; abaxial surfaces with short red-brown glandular hairs less than 0.1 mm among the scales. ................................................................................................................................. 5. A. sinuata. 2. Adaxial laminar scales extremely long-ciliolate, ca. 1 mm long, creating a web-like appearance when young, the scale body 1–2 cells wide; abaxial surfaces lacking glandular hairs among the scales. ...................................................................................................... 4. A. laevis. 1. Pinnae entire to shallowly lobed generally less than 1⁄3 of the way to the costae, or if more deeply lobed, then with imbricate, persistent adaxial scales 4–7 cells wide. 3. Largest pinnae 4–7 mm long; adaxial laminar scales 0.2–0.5 mm long, the scale body 5–7 cells wide; northern Mexico. ...... 1. A. cochisensis. 3. Largest pinnae (5–)7–20 mm long; adaxial laminar scales ca. 1 mm long; widespread in Mexico. 4. Adaxial laminar scales persistent, imbricate, the scale body 4–7 cells wide; largest pinnae (5–)7–12(–15) mm long. ........ 3. A. integerrima. 4. Adaxial laminar scales generally deciduous, scattered, the scale body 2–3 cells wide; largest pinnae 12–20 mm long. ........ 2. A. crassifolia.

1. ASTROLEPIS COCHISENSIS Astrolepis cochisensis (Goodd.) D. M. Benham & Windham, Amer. Fern J. 82: 57.1992. Figs. 46H, J, K. Notholaena cochisensis Goodd., Muhlenbergia 8: 93. 1912. Notholaena sinuata (Lag. ex Sw.) Kaulf. var. cochisensis (Goodd.) Weath. in I. M. Johnston, J. Arnold Arbor. 24: 314. 1943. Cheilanthes sinuata (Lag. ex Sw.) Domin var. cochisensis (Goodd.) Munz, Aliso 4: 87. 1958. Cheilanthes cochisensis (Goodd.) Mickel, Phytologia 41: 433. 1979. Type. U.S.A. Arizona: Cochise Co., Huachuca Mts, Montezuma Canyon, Goodding 373 (isotype GH!). Astrolepis cochisensis (Goodd.) D. M. Benham & Windham subsp. chihuahuaensis D. M. Benham, Amer. Fern J. 82: 59. 1992. Type. Mexico. Coahuila: 44 km southeast of Saltillo, Benham 1352 (ASC!; isotypes ARIZ!, UC!). Astrolepis cochisensis (Goodd.) D. M. Benham & Windham subsp. arizonica D. M. Benham, Amer. Fern J. 82: 60. 1992. Type. U.S.A. Arizona: Yavapai Co., unnamed tributary of Black canyon, 7 miles N of Black Canyon City, Benham 1312 (ASC!; isotypes ARIZ!, UC!).

Rhizomes short, stout, compact; rhizome scales 5–9 mm, tan or slightly darker near bases, linear, ciliate-dentate to entire; fronds 6–40 cm, clumped; stipes 1⁄6–1⁄3 the frond length; blades linear, pinnate to pinnate-pinnatifid; pinnae 14–45 pairs, ovate to oblong or narrowly deltate, cordate to truncate or rounded at bases, entire to shallowly lobed, less than 1⁄3 of the way to the costa, lobes 1–4, shallow, rounded; largest pinnae 4–7 ⫻ 2.5–4 mm, basal pinnae lacking basiscopic lobes; abaxial surfaces with dense, imbricate, ovate-lanceolate, mostly 0.5–1 mm long, ciliate scales; adaxial surfaces with sparse, deciduous to persistent, substellate to lanceolate, 0.2–0.5 mm long, coarsely ciliate scales, the scale body 5–7 cells wide; sporangia containing 32 or 64 spores; 2n⫽58 (Coah, NL); n⫽2n⫽87, 116 (USA).

ASTROLEPIS INTEGERRIMA

133

ex Sw.) Kaulf. var. pruinosa (Fe´e) E. Fourn., Mexic. Pl. 1: 120. 1872. Type. Mexico. “Valle´es de Toluca et d’Orizaba, Schaffner 167c (P, not found).

Rhizomes short, stout, compact; rhizome scales to 15 mm long, castaneous, linear, entire or only sparsely ciliate, sinuous; fronds clumped, 8–45 cm long; stipes 1⁄6–1⁄4 the frond length; pinnae 10– 33 pairs, each with 0–3 rounded lobes, lobed less than 1⁄3 of the way to costae; largest pinnae 12–20 ⫻ 4–8 mm, basal lobes without basiscopic lobules; adaxial surfaces glabrescent or sparsely clothed with white, ciliate, finely dissected scales ca. 1 mm long, the scale body linear, 2–3 cells wide; abaxial surfaces densely clothed with white to reddish brown, lanceolate, ciliolate scales; n⫽2n⫽87, apogamous (NL).

Distribution. Rocky slopes and cliffs, mostly limestone and other calcareous substrates; 1200–2400 m elevation. Sw USA; Mexico. Selected Specimens Examined. Ags (Rzedowski 25061, NY). BCN (Wiggins 7578A, DS). Chih (Mexia 2624, NY). Coah (Hinton 16512, NY). Dgo (Pennell 18610, NY). NL (Pennell 17036, NY). SLP (Parry & Palmer 991⁄2, NY). Son (Rose 12432, NY). Tam (Gonza´lez Medrano 8626, MEXU). Zac (Stanford et al. 548, NY).

Benham (1992) and Benham and Windham (in FNA Ed. Comm., 1993) recognized three subspecies of A. cochisensis based on spore number per sporangium, size of spore, and plant substrate. Here we treat these as cytotypes, without giving them formal rank. Key to Cytotypes (subspecies of Benham & Windham) of A. cochisensis 1. Spores 64 per sporangium, 39–46 ␮m; diploids of calcareous sites; N Mex, Tex; n Mexico. .......... cytotype chihuahuensis. 1. Spores 32 per sporangium, 59–86 ␮m; apogamous triploids of calcareous or non-calcareous sites. 2. Spores 72–86 ␮m; on non-calcareous substrates; Ariz. ........................................................................ cytotype arizonica. 2. Spores 59–70 ␮m; on calcareous substrates; CalifOkla; n Mexico. ................................................... cytotype cochisensis.

Benham (1989) reported that isozyme analyses suggest that subsp. cochisensis is an autotriploid derivative of the diploid subsp. chihuahuensis. Evidence from isozymes and substrate preferences of subsp. arizonica indicate, however, that it is not a simple autotetraploid and that other taxa remain to be discovered within the Astrolepis cochisensis complex (D. M. Benham & Windham, in FNA Ed. Comm., 1993).

2. ASTROLEPIS CRASSIFOLIA Astrolepis crassifolia (T. Moore & Houlston) D. M. Benham & Windham, Amer. Fern J. 82: 57. 1992. Figs. 45H, J-L. Notholaena crassifolia T. Moore & Houlston, Gard. Mag. Bot. 3: 20. 1851. Cheilanthes crassifolia (T. Moore & Houlston) Mickel & Beitel, Mem. New York Bot. Gard. 46: 111. 1988. Type. Cultivated, Houlston s.n. (BM, photo US). Notholaena pruinosa Fe´e, Me´m. Foug. 8: 78. 1857. Notholaena sinuata (Lag.

Distribution. Moist oak or pine-oak woods, deciduous forests, rocky soils, and crevices, gypsum hills, limestone rocks; 800– 3100 m. Mexico; Guat, Hond. Selected Specimens Examined. Chih (Knobloch s.n., 28 Aug 1938, DS). Chis (Ramı´rez S. 13, ENCB). DF (Lyonnet 44, NY). Dgo (Torrecillas 61, MEXU). Gro (Vela´zquez 1405, FCME, MEXU). Hgo (Lavin 4632, NY). Me´x (Cruz Cisneros 260, MICH, NY). NL (Taylor 118, NY). Oax (Mickel 4506, NY). Pue (Nicolas s.n., 22 Dec 1909, NY). Qro (Sohmer 9241, NY). SLP (Clausen & Edwards 7454, NY). Tam (Stanford et al. 2616, NY). Tlax (Sharp 44437, US). Ver (Bourgeau 355ter, K, frag. NY). Unverified, Doubtful, or Mistaken Reports. Gto (Rzedowski 46922, IEB, cited by Dı´az-Barriga & Palacios-Rios, 1992, but not verified). Mor (reported by Riba et al., 1986, but no specimen cited and report not verified).

Astrolepis crassifolia is distinct by the entire or subentire pinnae and narrow adaxial laminar scales with long cilia.

3. ASTROLEPIS INTEGERRIMA Astrolepis integerrima (Hook.) D. M. Benham & Windham, Amer. Fern J. 82: 57. 1992. Figs. 46A–G, L–O. Notholaena sinuata (Lag. ex Sw.) Kaulf. var. integerrima Hook., Sp. Fil. 5: 108. 1864. Notholaena integerrima (Hook.) Hevly (pro hyb.), J. Arizona Acad. Sci. 3: 207. 1965. Cheilanthes integerrima (Hook.) Mickel, Phytologia 41: 434. 1979. Type. Mexico. Liebmann s.n. (K!, photo GH). Astrolepis windhamii D. M. Benham, Amer. Fern J. 82: 60. 1992. Type. U.S.A. Arizona: Coconino Co., Oak Creek Canyon, 1.7 miles NE of Sedona, Benham 1385 (ASC!; isotypes ARIZ!, UC!).

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ASTROLEPIS

Rhizomes short, stout, compact; rhizome scales 8–14 mm long, castaneous, linear, sparsely toothed, slightly sinuous; fronds clumped, to 35 cm long; stipes 1⁄6–1⁄3 the frond length; blades linear, pinnate to pinnate-pinnatifid; pinnae 20–45 pairs, lobed less than halfway to costae, each with 0–3 pairs of rounded lobes; largest pinnae 7–12(–15) ⫻ 3–8 mm, basal lobes lacking basiscopic lobules; adaxial surfaces clothed with scattered to dense, persistent, imbricate, white, coarsely ciliolate, linear-lanceolate scales, the scale body 4–7 cells wide; abaxial surfaces densely clothed with white to reddish brown, lanceolate, ciliolate scales 1–1.5 mm long with coarse marginal projections; n⫽2n⫽87, apogamous (Chih, sw USA).

Distribution. Dry, rocky slopes, often limestone, scrub vegetation, basalt; 800–2600 m. Sw USA; Mexico; Hisp. Selected Specimens Examined. Ags (Rzedowski 25030, NY). Chih (Pringle 464, MICH, NY). Chis (Breedlove 45965, NY). Coah (Palmer 1406, NY). DF (Ramos 23, ENCB). Dgo (Gentry 8323, MEXU, MICH). Gro (Lorea 1442, ENCB). Gto (Rzedowski 47332, MEXU). Hgo (Fisher s.n., 14 Aug 1937, NY). Me´x (Go´mez M. 4, ENCB). NL (Chase 7770, MICH, NY). Oax (Mickel 3910, NY). Pue (Smith et al. 3672, NY). Qro (Sohmer 9240, MEXU). SLP (Bartholomew 3590, MEXU). Son (White 4585, NY). Tam (Hiriart 982, MEXU). Zac (Lloyd & Kirkwood 24, UC).

thought to be a hybrid between A. cochisensis and the unnamed Mexican species. Astrolepis windhamii is here treated as part of A. integerrima.

4. ASTROLEPIS LAEVIS Astrolepis laevis (M. Martens & Galeotti) Mickel, comb. nov. Figs. 45M–Q. Notholaena laevis M. Martens & Galeotti, Me´m. Foug. Mexique 15: 46. 1842. Notholaena sinuata (Lag. ex Sw.) Kaulf. var. integra Liebm., Mexic. Bregn. 213 (reprint 61). 1849. Type. Mexico. Oaxaca: Sola, Galeotti 6350 (BR!–6 sheets; isotypes P!, K!, frag. NY!). Notholaena deltoidea Hook. & Baker, Syn. Fil., ed. 2, 514. 1874. Cheilanthes deltoidea (Baker) Domin, Biblioth. Bot. 20(85): 133. 1915. Type. Mexico. s. loc., Leibold s.n. (K!). Notholaena sinuata (Lag. ex Sw.) Kaulf. var. madriensis Hevly, J. Arizona Acad. Sci. 3: 208. 1965. Type. Mexico. Me´xico: Temascaltepec, Hinton 1966 (GH; isotype MEXU!). Cheilanthes beitelii Mickel, Mem. New York Bot. Gard. 46: 107. 1988. Astrolepis beitelii (Mickel) D. M. Benham & Windham, Amer. Fern J. 82: 57. 1992. Type. Mexico. Oaxaca: Distrito Sola de Vega, 23–29 km S of Sola de Vega, 27–33 km N of bridge at Juchatengo, Mickel 6316 (NY!).

Rhizomes short, stout, compact; rhizome scales 8–18 mm long, orange-tan to maroon, linear, ciliate-denticulate, sinuous; fronds clumped, 18–80 cm long; stipes 1⁄4–1⁄3 the frond length; blades linear, pinnate-pinnatifid; pinnae 10–32 pairs, short-stalked, lobed ca. halfway to the costae, each with 3–5 pairs of acute lobes; largest pinnae 13–35 ⫻ 8–18 mm, basal lobes with basiscopic lobules; adaxial surfaces sparsely clothed with linear, white, ciliolate scales ca. 1 mm long, the body 1–2 cells wide, the long hair-like arms creating a web-like appearance when young; abaxial surfaces densely clothed with white to reddish brown, linear-lanceolate, ciliolate scales overlying a dense tomentum of finely dissected scales, red-brown glandular hairs absent.

Unverified, Doubtful, or Mistaken Reports. Ver (Calzada & Va´squez 10229, XAL, cited by Palacios-Rios, 1992, but not verified).

This taxon, considered by R. Tryon (1956) to be a variety of Astrolepis sinuata and by Hevly (1965) to be a hybrid between A. sinuata and A. cochisensis, is eglandular, has persistent, broad scales on the adaxial pinna surfaces, and small (8–12 ⫻ 4–5 mm) pinnae, each with 0–4 pairs of rounded lobes. The smaller size, greater number of lobes per pinna, and the imbricate persistent adaxial laminar scales distinguish Astrolepis integerrima from A. crassifolia. There seem to be three major forms placed here, and these may represent distinct taxa: pinnae undivided; pinnae shallowly lobed; and pinnae more deeply lobed, halfway to costa (Figs. 46E, F, & G, respectively). Astrolepis windhamii D. M. Benham (Figs. 46L–O) has been considered by Benham (1989) to be an apogamous allotriploid with one genome each from A. sinuata, A. cochisensis, and an undescribed Mexican taxon related to A. crassifolia. In the shallow pinna lobing and persistent linear-lanceolate adaxial laminar scales, A. windhamii closely resembles A. integerrima, which is

Distribution. Dry, rocky slopes, pine-oak forests, lava fields, road cuts, limestone outcrops; 900–3000 m. Mexico; Guat. Selected Specimens Examined. Ags (Rzedowski 25067, NY). Chih (Herna´ndez M. 8551, MEXU). Chis (Breedlove 21305, NY). Coah (Dorr 2510, MEXU). DF (Schaffner 37, NY). Dgo (Maysilles 8300, MEXU). Gro (Anderson 12873, MICH, NY). Gto (Ventura & Lo´pez 8372, MEXU). Hgo (Sohmer 9302, NY). Jal (McVaugh 15069, MEXU, NY). Me´x (Hinton 15400, NY). Mich (Hinton 12687, NY). NL (Schneider 1079,

ATHYRIUM

135

UC). Oax (Mickel 6344, NY). Pue (Purpus 3144, UC p.p.). Qro (Carranza G. 1817, MEXU). SLP (Pennell 17868, MICH, NY). Tam (Stanford et al. 663, NY). Tlax (Acosta P. 2215, IEB). Ver (Bourgeau 253 bis, NY, P). Zac (Dorado 1595, MEXU p.p.).

The large (13–35 ⫻ 8–18 mm) pinnae with their finely dissected adaxial scales (body only 1–2 cells wide) and the lack of short, glandular hairs abaxially on the blades distinguish this taxon from A. sinuata and A. crassifolia. Astrolepis laevis is distinct from A. crassifolia by the pinnae cut halfway to the costa with 3–5 pairs of lobes (vs. shallowly or not at all cut) and the adaxial laminar scales skeletonized and appearing web-like, the scale body 1–2 cells wide (vs. 2–3 cells wide and not appearing weblike).

5. ASTROLEPIS SINUATA Astrolepis sinuata (Lag. ex Sw.) D. M. Benham & Windham, Amer. Fern J. 82: 56. 1992. Figs. 45A–G. Acrostichum sinuatum Lag. ex Sw., Syn. Fil. 14. 1806. Notholaena sinuata (Lag. ex Sw.) Kaulf., Enum. Filic. 135. 1824. Gymnogramma sinuata (Sw.) C. Presl, Suppl. Tent. Pterid. 219. 1836. Cheilanthes sinuata (Lag. ex Sw.) Domin, Biblioth. Bot. 20(85): 133. 1913. Type. Peru. Lagasca s.n. (MA or S). Notholaena chalcolepis A. Braun ex Kunze, Linnaea 13: 135. 1839. Type. Mexico. 1827, Karwinski s.n. (B; isotype P!). Notholaena sinuata (Lag. ex Sw.) Kaulf. var. pinnatifida Farw., Amer. Midl. Naturalist 12: 283. 1931. Type. U.S.A. New Mexico: October 1880, Rusby s.n. (isotype NY!). Notholaena sinuata (Lag. ex Sw.) Kaulf. var. robusta Hevly, J. Arizona Acad. Sci. 3: 208. 1965. Type. Mexico. Michoaca´n: Zita´cuaro, Hinton et al. 13265 (ARIZ; isotypes NY!–2 sheets). Astrolepis sinuata subsp. mexicana D. M. Benham, Amer. Fern J. 82: 59. 1992. Type. U.S.A. Texas: Jeff Davis Co., Davis Mountains, Little Aguja canyon, D. M. Benham 1331 (ASC; isotype UC!).

Rhizomes short, stout, compact, horizontal; rhizome scales 5– 9 ⫻ 0.1 mm at bases, castaneous, often brown- to black-centered with age, linear, ciliate-denticulate, sinuous; fronds clumped, to 72(–100) cm long; stipes less than 1⁄4 the frond length, stout, terete, castaneous, densely clothed with white linear-lanceolate to deeply dissected-ciliolate scales; blades linear, pinnatepinnatifid; pinnae (14–)25–42 pairs, short-stalked to 3 mm, articulate, coriaceous, oblong, lobed ca. halfway to costae, each with 4–6 pairs of acute lobes; largest pinnae (near middle of blade) 12–20 ⫻ 6–10 mm, basal lobes with small basiscopic lobules; adaxial surfaces sparsely clothed with white, ciliolate to finely dissected substellate scales 0.5 mm long, the scale body 2–3 cells wide; abaxial surfaces densely clothed with white to reddish brown, linear-lanceolate, ciliolate scales overlying a dense tomentum of finely dissected hair-like scales and numerous short red-brown glandular hairs less than 0.1 mm; 2n⫽58 (USA); n⫽2n⫽87 (Jal, Zac, USA).

Distribution. Exposed limestone rocks and slopes, subdeciduous forests, granite boulders, thorn scrub, oak forests, crevices in lava; (400–)1450–2700 m. Sw USA; Mexico; Guat, Hond, Pan, Hisp, PR; Col, Ven, Ec, Peru, Bol, Parag, Arg. Selected Specimens Examined. Ags (Rzedowski 25028, MEXU). BCN (Franklin 5800, NY). BCS (Carter 5743, MEXU). Chih (Freytag & Baxter 87, NY). Chis (Breedlove 26963, NY). Coah (Stanford et al. 105, NY). DF (Bourgeau 255, NY). Dgo (Correll & Johnston 20050, NY). Gro (Hinton 9724, NY). Gto (Rusby 6, NY). Hgo (Fisher 35368, NY). Jal (Palmer 633, NY). Me´x (Rzedowski 30734, ENCB). Mich (Hitchcock & Stanford 7162, NY). Mor (Dunn & Dunn 18662, NY). Nay (Forman 1822, MICH). NL (Hinton 20074, MEXU). Oax (Mickel 4559, NY). Pue (Arse`ne 2051, MEXU). Qro (Rose 11532, NY). Sin (Breedlove 16433, NY). SLP (Pennell 17600, NY). Son (Gentry 1195, NY). Tam (Stanford et al. 663, NY). Ver (Purpus 9043, MO). Zac (Anderson 12614A, NY).

The inconspicuous dense glandular hairs of Astrolepis sinuata s.str., as well as the persistent, short, substellate scales of the adaxial pinna surfaces, distinguish this widespread species from the eglandular A. laevis, A. integerrima, and A. crassifolia. Specimens with very sparse glandular hairs may represent hybrids with these other three taxa (although parentage of the putative hybrids is unclear). Benham and Windham (in FNA Ed. Comm., 1993) distinguished subsp. mexicana (western Texas and southeastern New Mexico, Mexico, and Central America) from subsp. sinuata in having 64 rather than 32 spores per sporangium, and averaging 37–44 ␮m rather than 50–65; these are here treated as cytotypes. A very large specimen from Guerrero (Fonseca & Vela´zquez 2475, MEXU, FCME, NY) reaches 1 m in length, and the pinnae are deeply cut and with 7 pairs of lobes.

16. ATHYRIUM Athyrium Roth, Tent. Fl. Germ. 3: 31, 58. 1799. Lectotype (chosen by J. Smith, Hist. Fil. 327. 1875): Athyrium filix-femina (L.) Roth [⬅ Polypodium filix-femina L.]. See Pic. Serm. (Webbia 8: 437–442. 1952), for discussion of place of publication of Roth’s names. Terrestrial; rhizomes short-creeping or erect, scaly; fronds medium-sized, erect; stipes stramineous to red, sparsely scaly to glabrous, with 2 crescent-shaped vascular bundles at bases; blades

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ATHYRIUM

1-pinnate to bipinnate-pinnatifid, deltate to linear-lanceolate, laminae thin, glabrous; veins free, ending short of margin; sori elongate along veins, oblong to lunate or hooked over the vein to equally extended on both sides of a single vein, indusiate (ours) or not; spores bilateral, with folded to winged perispore; x⫽40. Athyrium is a genus of about 100 species, most of eastern Asia. Most species occur in temperate climates or high elevations of tropical regions. Athyrium skinneri occurs at lower elevations in Mexico and Central America. Athyrium is distinguished by the sori elongate along the veins with the indusia often acroscopically hooking over the vein. The genus appears to be related most closely to Diplazium, onocleoid ferns, blechnoid ferns, and thelypteroid ferns, but the exact relationships are still uncertain. Athyrium filix-femina is considered cosmopolitan, but has not been well studied. In Mexico there are two extremes of this complex, plus three members of the A. skinneri complex, the latter having creeping rhizomes, thinner blade texture, and occurring at lower elevations.

Athyrium pumilio Christ, Bull. Herb. Boissier, se´r. 2, 3: 147 1903. Type. Mexico. Chiapas: Near San Cristo´bal, Mu¨nch 91 (P!, photos NY!, UC!; isotypes DS!, US!, frag. NY!). Athyrium barnebyanum Mickel & Beitel, Mem. New York Bot. Gard. 46: 77. 1988. Type. Mexico. Me´xico: 5–6 mi NE of Amanalco on Rte 56, ca. 26 mi from jct with Rte 15, Mickel 657 (NY!; isotype ISC!).

Rhizomes erect; rhizome scales 6–8(–10) ⫻ 0.8–1 mm, brown, some with black streaks or slightly bicolorous, lanceolate, entire; fronds clumped; stipes 20–55 cm long, 1⁄2 the frond length, stramineous, glabrous except with scattered scales at bases; rachises stramineous, glabrous; blades bipinnate-pinnatifid, 24–40 ⫻ 15– 44 cm, deltate-lanceolate, basal pinnae not deflexed, (6.5–)8–17 ⫻ (2.5–)3.5–8 cm, apices pinnatifid, basal basiscopic pinnule short, 0.6–1.8 cm long, largest basiscopic pinnules 1.5–4 cm long, basiscopic pinnules larger than acroscopic ones, costae and pinnule surfaces glabrous or with scattered black hair-like scales, wings along costae and segment midribs conspicuous adaxially, often with a tooth 0.5–0.8 mm long at vein branching, pinna margins serrately incised to laciniate; sori linear to hooked; indusia 0.5–0.8(–1) ⫻ 0.1–0.3(–0.5) mm, entire or with occasional fimbriae.

Key to the Mexican Species of Athyrium 1. Rhizomes erect; fronds clumped, bipinnate-pinnatifid; 1800–3600 m elevation. 2. Fronds linear to linear-lanceolate, 7–18 cm wide; basal pinnae (2.5–)6–9.5 ⫻ (1.5–)2.5–3 cm, often deflexed; basiscopic pinnules of proximal pinnae roughly equal to acroscopic ones; rhizome scales pale brown. ........................................................................ 2. A. bourgeaui. 2. Fronds deltate-lanceolate, 15–44 cm wide; basal pinnae (6.5–)8–17 ⫻ (2.5–)3.5–8 cm, spreading or ascending; basiscopic pinnules of the proximal pinnae generally longer than acroscopic ones; rhizome scales pale brown or dark brown to black. ...................................................................................... 1. A. arcuatum.

1. Rhizomes creeping, apex protruding 2–6 cm past stipe bases; fronds distant, pinnate-pinnatifid to tripinnatepinnatifid; 100–1450 m. 3. Blades pinnate-pinnatifid, ovate-lanceolate; largest pinnae 3.5–9 ⫻ 1.5–2.5 cm; 100–1500(–2300) m. .... 4. A. skinneri. 3. Blades 2–3-pinnate-pinnatifid, deltate; largest pinnae 8–17 ⫻ 3.5–9 cm; 400–2300 m. 4. Blades bipinnate-pinnatifid. ................................ 3. A. palmense. 4. Blades tripinnate-pinnatifid. ................................... 5. A. tejeroi.

1. ATHYRIUM ARCUATUM Athyrium arcuatum Liebm., Mexic. Bregn. 278 (reprint 126) 1849. Type. Mexico. Puebla: Barranca de Chinautla, Liebmann s.n. [Pl. Mex. 2303, Fl. Mex. 863] (C!–2 sheets; isotype US!, frag. NY!). Figs. 48A, 49A, B, E–G. Athyrium galeottii Fe´e, Me´m. Foug. 5: 187. 1852. Asplenium martensii Mett., Abh. Senckenberg. Naturf. Ges. 3: 244. 1859, non Asplenium galeottii Fe´e, 1857. Athyrium martensii (Mett.) T. Moore, Index Fil. 186. 1860, nom. superfl. Type. Mexico. [Veracruz:] Pic d’Orizaba, Galeotti 6269 (P!; isotype K!, frag. NY!).

Distribution. Terrestrial in moist woods and along streams, deciduous forests, montane cloud forests; 1800–3200 m. Mexico; Guat. Selected Specimens Examined. Chis (Breedlove 27215, NY). DF (Rzedowski 34731, IEB). Gro (Lorea 4103, ENCB). Gto (Ventura & Lo´pez 9177, IEB). Hgo (Medina-Cota & Barrios 3728, NY). Jal (Rzedowski 27527, ENCB, NY). Me´x (Molseed 545, UC). Mich (Rzedowski 46181, IEB). Mor (Lyonnet 2508, MEXU). Nay (Te´llez 12776, MEXU). Oax (Mickel 4696, NY). Pue (Weber 781, ENCB). Qro (Dı´az B. & Carranza 7339, IEB). Ver (Ventura 9013, CAS, DS, ENCB, IEB, MO, NY, UAMIZ, XAL).

Two members of the Athyrium filix-femina complex occur in Mexico, and they have been considered as distinct species by Mickel and Beitel (1988). The name Athyrium arcuatum has been applied to large plants with deltate blades and basal pinnae largest, arcuate upwards at their distal ends. In contrast, the name Athyrium bourgeaui has been applied to plants with blades at least slightly reduced proximally. Unfortunately, there seem to be no characters that distinguish the two other than blade shape, and there are many intermediate

ATHYRIUM PALMENSE

specimens. Specimens from northwestern Mexico with reduced blade bases may be the same as what is called Athyrium filixfemina var. californicum Butters or var. cyclosorum Rupr. and may well be the same as A. bourgeaui since there seem to be no characters to distinguish them. Similarly, the same extremes in blade form seem to occur farther south in Central and South America. Much more work is needed on this complex in Latin America before we can apply specific names with confidence.

2. ATHYRIUM BOURGEAUI Athyrium bourgeaui E. Fourn., Mexic. Pl. 1: 102. 1872 [“bourgaei”]. Type. Mexico. “In sylvis vallis Mexicensis pr. San Nicolas,” Bourgeau 1042 (P!–2 sheets, frag. NY!, photos NY!, GH!, UC!, US!; isotype K!). Figs. 48B, 49C, D, H, J-L. Athyrium paucifrons C. Chr., Repert. Spec. Nov. Regni Veg. 9: 371. 1911. Type. Mexico. Michoaca´n: “A` l’ouest du Zapote, pre`s Morelia,” Arse`ne 25 (P!; isotypes BM!, P!–2 sheets, frag. NY!, US!).

Similar to A. arcuatum except: rhizome scales pale brown, 9–11 1–1.5 mm; stipes 7–47 cm long, 1⁄3–1⁄2 the frond length; blades 18–55 ⫻ 7–18 cm, linear to linear-lanceolate, basal pinnae often deflexed; basal pinnae (2.5–)6–9.5 ⫻ (1.5–)2.5–3 cm, basal basiscopic pinnule short, 0.6–0.8 cm long, largest basiscopic pinnule 1.6–2.5 cm long, basiscopic and acroscopic pinnules roughly equal in length, costae abaxially with sparse to scattered pale brown, hair-like scales 1–1.5 ⫻ 0.1–0.3 mm, pinna margins slightly recurved, serrate; wings along costae and segment midribs conspicuous adaxially, with short teeth 0.1 mm at vein branchings; indusia 0.5–0.8 mm long, entire or sparingly fimbriate. ⫻

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love 44488, CAS, MEXU). NL (Rodrı´guez L. 135, ANSM). Oax (Mickel 7433, NY, UC). Pue (Flores O. 122, ENCB). Qro (Dı´az-Barriga & Carranza 7285, NY). Son (Van Devender 97-850, ARIZ). Ver (Johnson s.n., 18 Sep 1906, NY).

This taxon was referred by Smith (1981) and Stolze (1981) to the north temperate species Athyrium filix-femina (L.) Roth, sensu lato, which is in need of modern taxonomic monographic work. We can find no distinct break between A. bourgeaui of Mexico and A. filix-femina vars. californicum and cyclosorum of the western United States, but until the relationships of the North American and European species are understood, we prefer to use the earliest Mexican name for this taxon. A similar case is made for using the earliest Mexican name (Athyrium arcuatum) for the taxon referred to by Smith (1981) as A. dombei Desv., a species described from Peru. Athyrium bourgeaui has lanceolate fronds, narrower than those of A. arcuatum, basal pinnae smaller and more deflexed, longer and broader rhizome scales, acroscopic and basiscopic pinnules roughly equal in size, and sparse to scattered, hair-like scales along the costae. Athyrium arcuatum has deltate-lanceolate fronds broader than those of A. bourgeaui, larger ascending to spreading basal pinnae, short and narrow rhizome scales, basiscopic pinnules longer than acroscopic pinnules, and glabrous costae. Athyrium barnebyanum, with scattered black, hair-like scales on the abaxial laminar surfaces, costae, and rachises, and dark brown to black scales 2.5–3 mm wide on rhizomes and stipe bases, resembles A. arcuatum in blade size and shape, and is here included in the latter species.

3. ATHYRIUM PALMENSE Athyrium palmense (Christ) Lellinger, Proc. Biol. Soc. Wash. 89: 708. 1977. Figs. 47E–G. Cystopteris fragilis (L.) Bernh. var. palmensis Christ, Bull. Herb. Boissier, se´r. 2, 7: 264. 1907. Type. Costa Rica. San Jose´: La Palma, Werckle´ 17054 (P!).

Similar to A. skinneri except: rhizome and stipe base scales 4– 5 ⫻ 0.5–1 mm; stipes 23–25(–30) cm long, 1⁄2 the frond length; blades bipinnate-pinnatifid, 23–28 ⫻ 12–28 cm, deltate; rachises with rare to sparse hair-like scales; largest pinnae 8–17 ⫻ 3.5–9 cm, wings along adaxial costae and segment midribs conspicuous, 0.1 mm wide, along almost entire length of axes; sori mostly hamate to round; indusia 0.5–0.8(–1) ⫻ 0.3–0.5 mm, coarsely dentate.

Distribution. Terrestrial in moist woods and along streams, pine forests, oak forests, evergreen cloud forests, montane cloud forests; 1850–3600 m. Mexico; Guat, Salv. Selected Specimens Examined. BCS (Brandegee s.n., 3 Oct 1899, UC). Chih (Correll & Gentry 23079, NY, UC). Chis (Breedlove 21998, NY). DF (Lyonnet 2682, MEXU). Dgo (Benı´tez P. 2282a, CHAPA, MEXU). Gto (Rzedowski 44805, IEB, MEXU). Hgo (Moore 5378, UC). Jal (Equipo 4-15475, GUADA, UC). Me´x (Rzedowski 30903, NY). Mich (Bartholomew 2862, CAS, NY). Mor (Sandoval 46, MEXU). Nay (Breed-

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ATHYRIUM

Distribution. Terrestrial in moist woods on Pacific side, mostly middle elevations, pine forests, cloud forests, pine-oak forests; 400–2300 m. Mexico; Salv, CR.

Distribution.Terrestrial in moist woods on Pacific side, mostly low elevations, tropical subdeciduous forests; 100–850(–1050) m. Mexico; Guat, Salv, Nic, CR, Pan; Col, Ec.

Selected Specimens Examined. Col (Va´zquez & Phillips 942, UC, UCR). Dgo (Ibana Garcı´a 647, US). Gro (Lorea 2536, MEXU). Jal (McVaugh 16203, US). Me´x (Hinton 4941, NY, US). Mich (Pringle 13706, US). Mor (Lyonnet 2578, MEXU, US). Nay (McVaugh 16345, MEXU, NY, US). Oax (Mickel 6124, NY, UC). Qro (Carranza G. 2266, IEB). Sin (Ortı´z C. et al. 257A, MEXU).

Selected Specimens Examined. Chis (Breedlove 37774, NY). Col (McVaugh 15823, US). Gro (Mexia 8859, B, NY, UC, US). Jal (McVaugh 19951, CAS, IEB, NY). Me´x (Hinton 2649, NY, US). Mich (King & Soderstrom 4926, NY, UC). Nay (Feddema 816, MEXU; Te´llez & Chiang 9712, MEXU). Oax (Mickel 5979, NY, UC). Sin (Salazar 689, US). See A. palmense and A. tejeroi for comparison and discussion.

Athyrium palmense is found primarily at higher elevations than is A. skinneri, though it occupies roughly the same geographical distribution; the mixed collections in Oaxaca (Mickel 3946, 6124, 6992, NY) are from 750–850 m. The deltate blades, larger pinnae, shorter sori, and distinctive costal wings appear to correlate with the bipinnate-pinnatifid nature of the fronds, and separate A. palmense from A. skinneri.

4. ATHYRIUM SKINNERI Athyrium skinneri (Baker) Diels in Engler & Prantl, Nat. Pflanzenfam. 1(4): 224. 1899. Figs. 47A–D. Asplenium skinneri Baker in Hooker & Baker, Syn. Fil. 226. 1867. Type. Guatemala. Skinner s.n. (K!, frag. NY!, UC! photo ex K).

Rhizomes creeping, with apices protruding 2–4 cm beyond stipe bases, sparsely scaly; rhizome scales 3–4 ⫻ 0.3–0.5 mm, light brown to brown, linear-lanceolate to narrowly deltate, slightly twisted; fronds distant; stipes 9–22 cm long, 1⁄3–1⁄2 the frond length, pink to rarely stramineous, glabrous except for scattered scales (similar to rhizome scales) at bases, broadly grooved to somewhat flattened; rachises pink and flexuous, broadly grooved, very narrowly winged; blades pinnate-pinnatifid, 12–20 ⫻ 6–15 cm, ovate-lanceolate, free pinnae 4–10 pairs, stalked, apices pinnatifid, largest pinnae 3.5–9 ⫻ 1.5–2.5 cm, laminae membranaceous, abaxially glabrous or with rare hair-like scales on costae and pinna bases, adaxially glabrous with papillae 0.1 mm in pinna axils; pinna margins serrate, wings along adaxial costae and segment midribs absent or minute, not conspicuous and not continuous; sori mostly linear to hamate, some diplazioid or round; indusia deeply laciniate, 0.5–1(–2) ⫻ 0.5–0.8 mm, coarsely dentate.

5. ATHYRIUM TEJEROI Athyrium tejeroi Mickel & Tejero, Brittonia 56: 199. 2004. Athyrium tripinnatum Mickel & Tejero, Brittonia 56(2): 115, nom. illeg., non Tagawa, 1937. Type. Mexico. Me´xico: Mpio Tejupilco, km 38 a Nanchititla (El Peparo), 28 Sep–1 Oct 1988, Tejero-Dı´ez 2919 (IZTA!; isotype NY!). Figs. 47H, J, K. Rhizomes short-creeping, 3 mm diam.; rhizome scales light brown, narrowly deltate, with long-acuminate apices, the margins entire, to 4 ⫻ 0.5 mm; fronds approximate (2–4 mm distant) to 60 cm long; stipes ca. 1⁄3 frond length, stramineous, adaxially grooved, with sparse linear scales similar to those of rhizomes and appressed septate hairs 0.2–0.5 mm long (mostly on abaxial surface); blades tripinnate-pinnatifid, broadly deltate, to 30 cm wide, with 15–18 pairs of free pinnae, the blade apices shortly attenuate; rachises subflexuous, stramineous, slightly alate, with scattered hairs similar to those of stipes mainly on abaxial surface; pinnae alternate, narrowly triangular, ascending, the basal ones to 19 ⫻ 10 cm, texture herbaceous, abaxial surface with appressed hairs 0.1–0.2 mm long, adaxial surface glabrous; veins pinnate, ending in the teeth; sori one per ultimate segment, 1– 1.5 mm long, elongate, J-shaped, or round; indusia 0.2–0.4 mm wide, whitish, coarsely toothed.

AZOLLA

Distribution. Terrestrial in oak woods; 1500 m. Known only from the type collection. This species is similar to A. palmense in the slender rhizomes, the thin blade texture, and the indusium deeply lacerated into 2–4 teeth, but differing in the blades being 3-pinnate-pinnatifid rather than 2-pinnate-pinnatifid; from A. skinneri differing in pinnate-pinnatifid blades. With increasing dissection, there is increasing frond size: A. skinneri to 44 cm long with 5–7 pairs of pinnae, A. palmense to 53 cm long with 12–15 pairs, and A. tejeroi to 60 cm with 15–18 pairs.

17. A Z O L L A Azolla Lam., Encycl. 1: 343. 1783. Type: Azolla filiculoides Lam. Heterosporous, minute, free-floating or on mud; stems slender, branching, with unbranched roots and two rows of alternate, imbricate leaves; leaves bilobed, with an upper green lobe and colorless lower lobe; cavities in the leaves contain a nitrogenfixing cyanobacterium, Anabaena azollae; sporangia in nut-like unisexual sporocarps; microsporangiocarps globose with numerous microsporangia; megasporocarps each with single megasporangium containing a single megaspore, the microspores in 4 clumps (massulae) per sporangium, each massula with anchorshaped glochidia, the megaspore apparatus with 3 or 9 flotation lobes; x⫽22. Azolla is a genus of aquatic, leptosporangiate, heterosporous ferns, and comprises perhaps six species, two widely distributed in the New World, ranging from the United States and Canada to Argentina. Four species are often cited in neotropical floras, e.g., Flora Mesoamerica (Moran in Davidse et al., 1995) and Peru (Tryon & Stolze, 1994). All these belong to sect. Azolla, characterized by the megasporocarps having three floats and the roots lacking root hairs; section Rhizosperma (Meyen) Mett. is apparently known natively only in the Old World and comprises two additional species (Saunders & Fowler, 1992). Especially within sect. Azolla, species boundaries are unclear, and for Mexico, we choose to recognize only two of the four commonly accepted

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species, regarding the other two as synonymous (at least as to the types) with the two we do treat. Identification of species is based largely on microscopic characters: whether the glochidia on the microsporangia are septate or not (Svenson, 1944); megaspore surface ornamentation; zonation of the perine (Perkins et al., 1985); and number of cells in epidermal trichomes (Pieterse et al., 1977). Because the majority of specimens are sterile, reproductive character states often cannot be evaluated, and so identification is uncertain. Even if fertile material (indicated by “f” in the citation of specimens below) is available, species identification may not be possible, even with a dissecting microscope (40 times magnification); a compound microscope and even the scanning electron microscope may be necessary. The duckweed genera Spirodela, Lemna, Wolffia, and Wolffiella (Lemnaceae) often occur mixed with Azolla, sometimes with 5 or 6 species on one sheet. Additional records for Azolla can often be found by checking herbarium specimens of these other small aquatics. All these genera have great proclivity for distribution by waterfowl and by humans. Various species of Azolla have been used as a green manure, especially in rice paddies. As the plants dry and decay, they release nitrogen from the nitrogen-fixing cyanobacterial colonies of Anabaena azollae, contained in the leaf cavities of the Azolla. Azolla is related to Salvinia, but the origins of both genera and their ties with the other ferns are remote and not easily discernible. Marsilea and allies have recently been shown to be the sister group to Salvinia and Azolla (Hasebe et al., 1995; Pryer, 1993; Pryer et al., 1995; Rothwell & Stockey, 1994) based on both paleontological and molecular evidence. The spores of extant Azolla appear to be relatively unchanged from the earliest known fossils of Azolla from the Lower Cretaceous. References Derrick, L. N., A. C. Jermy & A. M. Paul. 1987. Checklist of European pteridophytes. Sommerfeltia 6: ix–x; Dunham, D. G. 1986. Taxonomic re-evaluation and species recognition of Azolla Lam., with particular reference to section Azolla. Ph.D. thesis, Portsmouth Polytechnic, Portsmouth, UK; Fowler, K. & J. Stennet-Willson. 1978. Sporoderm architecture in modern Azolla. Fern Gaz. 11: 405–415; Lumpkin, T. A. 1993. Azollaceae. Pp. 338–342 in Flora of North America, north of Mexico. Vol. 2, ed. Flora of North America Editorial Committee; Perkins, S. K., G. A. Peters, T. A. Lumpkin & H. E. Calvert. 1985. Scanning electron microscopy of perine architecture as a taxonomic tool in the genus Azolla Lamarck. Scan. Electron Microscop. 4: 1719–1734; Pieterse, A. H., L. de Lange & J. P. Van Vliet. 1977. A comparative study of Azolla in the Netherlands. Acta Bot. Neerl. 26: 433–449; Pryer, K. M. 1993. Ontogeny and phylogeny in the aquatic fern family Marsileaceae: cladistic analysis of morphological and molecular data. Amer. J. Bot. 80 (Suppl.): 172; Reid, J. D. 2002. Systematics of the genus Azolla (Azollaceae). Botany 2002, abstracts: 102; Rothwell, G. W. & R. A. Stockey. 1994. The role of Hydropteris pinnata gen. et sp. nov. in reconstructing the cladistics of heterosporous ferns. Amer. J. Bot. 81: 479–492; Saunders, R. M. K. & K. Fowler. 1992. A morphological taxonomic revision of Azolla Lam. section Rhizosperma (Mey.) Mett. (Azollaceae). Bot. J. Linn. Soc. 109: 329–357; Saunders, R. M. K. & K. Fowler. 1993. The supraspecific taxonomy and evolution of the fern genus Azolla (Azollaceae). Pl. Syst. Evol. 184: 175–193; Svenson, H. K. 1944. The New World species of Azolla. Amer. Fern J. 34: 69–84; Zimmerman, W. J., R. Quintero Lizaola & R. FerreraCerrato. 1993. The genetic identification of species of agronomic Azolla Lam. indigenous to Mexico. Amer. Fern J. 83: 97–104; Zimmerman, W. J., I. Watan-

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abe, T. Ventura, P. Payawal & T. A. Lumpkin. 1991. Aspects of the genetic and botanical status of neotropical Azolla species. New Phytol. 119: 561–566.

Key to the Mexican Species of Azolla 1. Plants pinnately branched, 2–6 cm long; megaspores warty with raised angular bumps; largest hairs on upper leaf lobes unicellular. ...................................................... 1. A. filiculoides. 1. Plants Ⳳ dichotomously branched, 0.5–2 cm long; megaspores pitted or not, lacking raised angular bumps; largest hairs on upper leaf lobes with 2 or more cells. ........................................................................................ 2. A. microphylla.

1. AZOLLA FILICULOIDES Azolla filiculoides Lam., Encycl. 1: 343. 1783. Type. Straights of Magellan. Commerson s.n. (P-Lam!). Figs. 50D–H, J. Azolla caroliniana Willd., Sp. Pl., ed. 4, 5(1): 541. 1810. Type. “Habitat in aquis Carolinae.” Richard s.n. (B-Willd. 20260!). A synonym of A. filiculoides according to Dunham, as reported by Jermy in Derrick et al., 1987.

probably a misidentification). Tab (reported by Magan˜a, 1992, as A. caroliniana, but probably based on a misidentification).

2. AZOLLA MICROPHYLLA Azolla microphylla Kaulf., Enum. Filic. 273. 1824. Type. U.S.A. California: Chamisso s.n. (LE?). Figs. 50A–C. Azolla mexicana C. Presl, Abh. Ko¨nigl. Bo¨hm. Ges. Wiss., ser. 5, 3: 150. 1845, non Schltdl. & Cham., Linnaea 5: 625. 1830, nom. nud. Type. Mexico. [Veracruz:] “inter Serpillo et Estero,” Schiede & Deppe [839] (B!).

Plants flattened, Ⳳ dichotomously branched, 0.5–2 cm diam.; upper leaf-lobes imbricate, rhombic-ovate to suborbicular, ca. 0.5–0.7 mm long, broadly rounded to obtuse at apices, papillose on upper surfaces, lower lobe larger than upper, the margins purplish-tinged or hyaline; largest papillae on upper leaf lobes 2–3-celled; microsporocarps globose, ca. 1 mm diam.; glochidia of massulae 0–6-septate; megasporocarps 0.25–0.3 mm long; megaspores pitted on basal portion.

Differs from A. microphylla by: plants elongate, pinnately branched, frequently 2–6 cm long; dorsal (upper, photosynthetic) leaves larger, to 1 mm long, more prominently papillose, papillae 1-celled; megaspores warty with raised angular bumps, with relatively sparse filamentous hairs on the surface.

Distribution. Floating in water, marshes, stagnant streams; (0–)1400–2950 m. Canada (Brit Col), USA; Mexico; Hond, Nic; Gr Ant; Col, Ven, Fr Gui, Ec, Peru, s Braz, Bol, n Arg, Uru; naturalized in Europe and elsewhere?

Distribution. Floating in water, marshes, stagnant streams; 1500–2550 m. Alaska, w USA; Mexico; Guat, Hond, Nic; Col, Ven, Bol, Chile, Arg, Uru; e Asia, Hawaii, New Zealand, Australia, naturalized in Europe (and elsewhere?). Specimens Examined. Chih (Hartman 614, NY). Me´x (Pringle 5222, NY). Unverified, Doubtful, or Mistaken Reports. Ags (Siqueiros 3350, HUAA, cited by Siqueiros-Delgado & Gonza´lez-Adame, 2004, but not verified). DF (reported by Zimmerman et al., 1993, but not verified). Hgo (reported by Zimmerman et al., 1993, but not verified). Mor (Bonilla 64, HUMO, cited by Riba et al., 1996, but we regard this number, also at UAMIZ, as A. microphylla; Campos F. 12, HUMO, also cited by them). Oax (reported by Zimmerman et al., 1993, but not verified). Pue (reported by Zimmerman et al., 1993, but not verified). Qro (Pineda s.n., ENCB, QMEX, cited by Arreguı´n et al., 2001, but not verified). Sin (reported by Vega A. et al., 1989, but

Selected Specimens Examined. Ags (Rzedowski & McVaugh 734, ENCB). BCN (Thorne 57742, UC, f). Chis (Breedlove 38372, 34927, DS). DF (Rzedowski 21992, ENCB, NY). Gro (Fonseca 1375, 1441, 1590, FCME). Gto (Rzedowski 47246, IEB, UC). Hgo (Rzedowski 32098, IEB). Jal (Mexia 1857, NY, UC). Me´x (Rzedowski 31984, 33730, ENCB). Mich (Dı´az B. 6388, 6467, IEB, UC). Mor (Bonilla 64, UAMIZ). Nay (Jones 23461, NY, UC). Oax (Camp 2764, ENCB, NY, UC). Pue (Arse`ne 1208, 2363, US, both f). Sin (Alava 1421, UC, f). SLP (Schaffner s.n., “in 1876 et 81,” NY, US). Tab (Lot & Novelo 1099, MEXU). Ver (Gutie´rrez B. 1222, IEB, NY, XAL). Unverified, Doubtful, or Mistaken Reports. Chih (Knobloch 370, MSC, cited by Knobloch & Correll, 1962, but not verified). Tlax (Zimmerman et al., 1993, but not verified).

There is disagreement over the typification of A. microphylla. Svenson (1944) applied the name primarily to material from South America, but the type was said by Kaulfuss to have been collected in California. Most authors have followed Svenson in excluding California from the range, implying that they thought

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the provenance of the type was incorrect; however, there is no firm evidence to support this contention, and at least two recent authors have stated otherwise (Tryon & Stolze, 1994; Moran in Davidse et al., 1995). Even Svenson himself stated that “if the Chamisso specimen, which was fruiting, is ever examined, I suspect it will turn out to be either A. mexicana or A. filiculoides.” Most specimens of Azolla from California appear to be the same as Mexican material, and so we apply the older of the two names. Since Svenson’s (1944) revision of the group, A. mexicana has usually been distinguished from A. microphylla by the branching pattern of the plants (said to be dichotomous in A. mexicana, pinnate in A. microphylla) and megaspore ornamentation (pitted in A. mexicana, smooth in A. microphylla). Other stated differences are not clearcut or the character states are strongly overlapping. Zimmerman et al. (1993) reported A. microphylla to be genetically similar to A. mexicana (as A. caroliniana). Reid (2002) used cladistic analysis of DNA sequence data from three noncoding regions (ITS, atpB-rbcL IGS, and trnL-trnF region) to reconstruct phylogenetic relationships among species of Azolla. She interpreted her results to indicate that the species of sect. Azolla form a monophyletic group. However, A. microphylla and A. mexicana formed a single evolutionary lineage, and multiple accessions sampled from these putative species were interdigitated within the clade. She suggested that this lineage arose through hybridization and homoploid speciation involving A. caroliniana and A. filiculoides, an intriguing hypothesis that needs further testing. For this reason, we choose to treat A. mexicana and A. microphylla as conspecific. Azolla filiculoides Lam. differs from A. microphylla in having more elongate, pinnately branched stems and generally larger leaves with broader, whitish margins. Most specimens previously identified as A. filiculoides from Mexico are probably A. microphylla, as judged by their dichotomous branching pattern and small, narrowly marginate or emarginate leaves. Specimens of A. microphylla in Mexico and elsewhere have sometimes been identified as A. caroliniana Willd., but Jermy (in Derrick et al., 1987) reported that the type of that name was shown by Dunham (1986) to be a synonym of A. filiculoides Lam. Svenson (1944) applied the name A. caroliniana in a strict sense, restricting it to specimens from the eastern United States and perhaps the Greater Antilles. Azolla mexicana is regarded by some as only a minor variant of A. caroliniana sensu Svenson (non Willd.) (as, for example by Zimmerman et al., 1993). The same species is also naturalized in the Old World, e.g., in Europe (Pieterse et al., 1977, as A. caroliniana).

1 8. B L E C H N UM Blechnum L., Sp. Pl. 2: 1077. 1753. Lectotype (chosen by J. Smith, Hist. Fil. 300. 1875): Blechnum occidentale L. Struthiopteris Weiss, Pl. Crypt. Fl. Gott. 286. 1769. Type: Struthiopteris spicant (L.) Weiss [⬅ Blechnum spicant (L.) Sm.]. Lomaria Willd., Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck. Gesammten Naturk. 3: 160. 1809. Type: Lomaria nuda (Labill.) Willd. [⬅ Onoclea nuda Labill.] ⬅ Blechnum nudum (Labill.) Luerss.

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Parablechnum C. Presl, Epimel. Bot. 109. 1849 [1851]. Blechnum sect. Parablechnum (C. Presl) T. Moore, Index Fil. xxv. 1857. Type: Parablechnum procerum (G. Forst.) C. Presl [⬅ Osmunda procera G. Forst.] ⬅Blechnum procerum (G. Forst.) Sw. Lomaridium C. Presl, Epimel. Bot. 154. 1849 [1851]. Lomaria sect. Lomaridium (C. Presl) J. Sm., Hist. Fil. 303. 1875. Type: Lomaridium plumieri (Desv.) C. Presl [⬅ Lomaria plumieri Desv.] ⫽ Blechnum binervatum (Poir.) C. V. Morton & Lellinger.

Terrestrial, hemiepiphytic or epiphytic; rhizomes longcreeping to compact and erect, sometimes forming trunks, scaly, often stoloniferous and forming large colonies; fronds monomorphic or dimorphic, small to medium-sized (to large); stipes adaxially grooved, stramineous to atropurpureous, glabrous to sparsely or moderately scaly, non-articulate to rhizomes; fronds mostly pinnatifid to pinnate, rarely undivided, margins entire to serrulate, young expanding fronds usually reddish; blades glabrous, occasionally with small scales on costae and veins, chartaceous to coriaceous; veins of sterile blades free, parallel, simple or 1–2-forked, running to or nearly to margins; sori costal, on commissural veins (connecting adjacent veins) parallel and close to costae, indusiate; indusia directed inwardly, opening toward costae, entire to laciniate; spores bilateral, perispore inconspicuous to winged; x⫽28, 29, 31, 32, 33, 34, 36. Blechnum is the largest genus of the family Blechnaceae and is related to Woodwardia. The nearly 200 species are primarily tropical, with four boreal-temperate species and several australtemperate species. There are 15 species in Mexico; most are widespread, extending into Central and South America. The genus has sometimes been subdivided on the basis of monomorphic (Blechnum) vs. dimorphic fronds (Lomaria), but there are several groups of dimorphic species, each of which could be justified as a distinct genus: sect. Lomariocycas (J. Sm.) C. V. Morton, with stout trunks (Costa Rica to Bolivia); sect. Parablechnum (C. Presl) T. Moore, with stout, short, erect rhizomes (e.g., B. schiedeanum); an unnamed group, with erect, slender trunks, pinnatifid sterile blades, and reduced proximal pinnae (e.g., B. divergens, B. lehmannii, B. wardiae); and sect. Lomaridium (C. Presl) comb. ined., with climbing rhizomes and toothed rhizome scales (e.g., B. fragile, B. ensiforme). The monomorphic species seem closely allied to each other, with the exception of B. serrulatum (in sect. Diafnia (C. Presl) J. Sm.). Recent molecular work by Cranfill (2001) has demonstrated that a monophyletic Blechnum can be obtained only by the recognition of many smaller genera, e.g., Lomaria, Lomaridium, Lomariocycas, and Parablechnum, mentioned above. Such a re-circumscription would restrict the application of Blechnum to the group of monomorphic species allied to B. occidentale. The Mexican species of Blechnum are characterized by adaxially grooved stipes and rachises, with the grooves usually extending into the costae, except in smaller species like B. stoloniferum. Most Mexican species have enlarged vein endings on the blade surfaces adaxially, except B. fragile and B. ensiforme, which have vein endings visible only on the abaxial surfaces, and B. serrulatum, in which the veins run into cartilaginous pinna margins.

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Among the monomorphic blechnums, hybrids appear to be numerous, as judged by intermediate morphology and malformed spores. Many of these species and hybrids in this group are relatively common in disturbed and semi-disturbed habitats, such as along roadsides and trails. Definitive assignment of parentage of putatively hybrid plants must await further morphological, cytological, and electrophoretic studies, in many cases.

References Broadhurst, J. 1912. The genus Struthiopteris and its representatives in North America. Bull. Torrey Bot. Club 39: 257–278, 357–385; Cranfill, R. B. 2001. Phylogenetic studies in the Polypodiales (Pteridophyta) with an emphasis on the family Blechnaceae. Ph.D. thesis, University of California, Berkeley, California; Morton, C. V. & D. B. Lellinger. 1967. Notes on the ferns of Dominica and St. Vincent. Amer. Fern J. 57: 66–77; Murillo, M. T. 1968. Blechnum subge´nero Blechnum en Sur Ame´rica, con especial referencia a las especies de Colombia. Nova Hedwigia 16: 329–366.

Key to the Mexican Species of Blechnum 1. Fronds monomorphic, the fertile ones not contracted. 2. Pinnae articulate at rachises, margins serrulate, each tooth involving 1–4 veins; enlarged vein endings not readily visible; sterile fronds with pale brown scales (1 mm long) on abaxial costae. ......................................................................................................... 13. B. serrulatum. 2. Pinnae not articulate, margins entire to obscurely serrulate, each tooth a modified marginal cell; enlarged vein endings visible adaxially; sterile fronds glabrous or with septate hairs, without scales. 3. Blades with a conform terminal pinna; lateral pinnae usually 2–8 pairs; costae and rachises glabrous abaxially. 4. Lateral pinnae usually 1⁄3–2⁄3 the length of the terminal pinna, usually 2–4(–8) pairs; terminal pinna subcordate or with confluent or separate semicircular lobes proximally; lateral pinnae with broadly cuneate to rounded bases. .................... 6. B. gracile. 4. Lateral pinnae Ⳳ equalling the terminal pinna in length, usually 2–3 pairs; terminal pinna cuneate proximally; lateral pinnae narrowly cuneate at bases. .............................................................................................................................................. 8. B. meridense. 3. Blades with gradually tapering, pinnatifid apices; lateral pinnae 10 or more pairs, adnate or short-stalked with rounded to cordate bases; costae and rachises glabrous or hairy abaxially. 5. Blades linear, with much reduced, adnate, deltate proximal pinnae; costae not usually grooved adaxially. ............. 11. B. polypodioides. 5. Blades ovate to deltate, slightly or not at all reduced at bases, proximal pinnae stalked, with cordate bases; costae grooved adaxially. 6. Rachises sparsely to usually densely hairy with septate hairs 0.3–0.8 mm long; blades 7–23 ⫻ 3.5–8(–11) cm; stipes 3.5–21 cm long; largest pinnae 2.3–4(–6.5) cm long; 700–2550 m. ...................................................................... 1. B. appendiculatum. 6. Rachises glabrous; blades 17–40 ⫻ (8.5–)11–17(–22) cm; stipes 17–30 cm long; largest pinnae (4.5–)6–9(–12.5) cm long; 50–900 m. .................................................................................................................................................................. 9. B. occidentale. 1. Fronds dimorphic, the fertile ones strongly contracted as compared to the sterile ones. 7. Sterile blades pinnate, contracted at bases, at least the proximal pinnae not adnate; plants terrestrial; rhizome scales ovate to lanceolate, entire, concolorous; vein endings enlarged and visible on adaxial blade surfaces. 8. Pinna margins regularly serrate; costae of sterile and fertile pinnae with dense scales 2–3 ⫻ 1–1.5 mm abaxially; rachises stramineous. ......................................................................................................................................................................................... 4. B. falciforme. 8. Pinna margins entire or occasionally serrulate, often crispate; costae of sterile and fertile pinnae abaxially with scattered scales 1–3 ⫻ 0.5–0.8 mm; rachises stramineous or purplish. ................................................................................................... 12. B. schiedeanum. 7. Sterile blades pinnatifid to pinnatisect, all segments adnate; plants terrestrial or epiphytic; rhizome scales lanceolate and concolorous or linear and bicolorous; vein endings enlarged on abaxial or adaxial blade surfaces. 9. Plants hemiepiphytic or epiphytic, rhizomes scandent with distant fronds; rhizome scales linear, bicolorous with curved marginal teeth; enlarged vein endings on abaxial surfaces. 10. Sterile pinnae 10–15 cm long; sterile blades (17–)22–30 cm wide; proximal pinnae abruptly reduced to auricles, median and proximal pinnae distant, 4–30 mm apart; fertile pinnae shorter than sterile pinnae; rhizome scales dark brown with darker central stripe, curved marginal teeth small (0.1 mm or less long); costae deeply grooved, elevated and of lighter color than laminae. .......................................................................................................................................................... 3. B. ensiforme. 10. Sterile pinnae 1.5–4 cm long; sterile blades (2–)3.5–4.5(–8) cm wide; proximal pinnae gradually reduced to auricles, basal auricles distant or not; fertile pinnae equal to or longer than sterile pinnae; rhizome scales pale brown with central dark stripe and curved marginal teeth 0.1–0.3 mm long; costae shallowly grooved and same color as laminae. .. 5. B. fragile. 9. Plants terrestrial, rhizomes erect with clumped fronds or short-creeping, sometimes stoloniferous; rhizome scales ovatelanceolate, concolorous with entire or ciliate margins; enlarged vein endings on adaxial surfaces. 11. Proximal pinnae of sterile blades abruptly reduced to mere vestigial pinnae a few mm long; sterile blades (16–)24–30 cm wide, pinnae 10–16 cm long. ................................................................................................................................................ 2. B. divergens. 11. Proximal pinnae of sterile blades gradually reduced to green, semicircular lobes, lacking vestigial pinnae; sterile blades 2–15 cm wide, pinnae 1–8 cm long. 12. Sterile blades 40–80 ⫻ 12–15 cm, pinnae 5.5–8 cm long; semicircular lobes running down almost to rhizomes; rhizome scales 10–20 mm long. ........................................................................................................................................... 15. B. wardiae. 12. Sterile blades 8–27 ⫻ 2–5(–9) cm, pinnae 1–5 cm long; semicircular lobes running down almost to rhizomes, or stipes 1⁄4 the frond length; rhizome scales 3–5 mm long. 13. Sterile blades (3–)3.5–9 cm wide; pinnae (1.5–)2–4.5 cm ⫻ 1–1.5 mm; rhizomes erect, non-stoloniferous; 1350– 1750 m. ........................................................................................................................................................................... 7. B. lehmannii.

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13. Sterile blades 2–3.5 cm wide; pinnae 1–2 cm ⫻ 5–8 mm; rhizomes short-creeping, stoloniferous; 1600–3250 m. 14. Fertile pinnae 30⫹ pairs, surcurrent at bases, arcuate, conduplicate, not apiculate at apices; fertile blades parallel-sided for most of their length; sterile blades subcoriaceous; costae of sterile pinnae grooved adaxially. ......................................................................................................................................................... 10. B. penna-marina. 14. Fertile pinnae 15–20 pairs, equilateral (equally surcurrent and decurrent) at bases, falcate-ascending, not conduplicate, apiculate at apices; fertile blades tapering from blade middle to apex; sterile blades chartaceous; costae of sterile pinnae not grooved adaxially. ...................................................................... 14. B. stoloniferum.

1. BLECHNUM APPENDICULATUM Blechnum appendiculatum Willd., Sp. Pl., ed. 4, 5(1): 410. 1810. Type. Venezuela. Caracas, Bredemeyer s.n. (B-Willd. 20038photo GH; microfiche UC!). Fig. 54D. Blechnum glandulosum Kaulf. ex Link, Enum. Hort. Berol. Alt. 2: 462. 1822. Type. Described from cultivated material, Brazil, Otto s.n. (B!). Blechnum meridionale C. Presl in J. Presl & C. Presl, Delic. Prag. 1: 186. 1822. Blechnum occidentale L. var. β intermedium E. Fourn., Mexic. Pl. 1: 112. 1872. Type. Brazil. Rio de Janeiro, Pohl s.n. (PR, frag. NY!). Blechnum occidentale L. var. minor Hook., Sp. Fil. 3: 51. 1860. Lectotype (chosen by Proctor, 1985: 289). Mexico. Liebmann s.n. (K!; isolectotype GH) [probably Veracruz: Mirador, labelled as B. glandulosum by Liebmann at C]. Blechnum occidentale L. var. pubirhachis Rosenst., Hedwigia 46: 94. 1906. Type. Brazil. Santa Catarina: Blumenau, Haerchen 18.1 (S; isotype NY!).

Similar to Blechnum occidentale except: stipes (3.5–)5–25 cm long, 1⁄3–1⁄2 the frond length; blades 7–23 ⫻ 3.5–8(–11) cm; rachises moderately to densely hairy with sticky, septate hairs 0.2– 0.8 mm long; largest pinnae (1–)2–4(–8) cm ⫻ (4–)6–8 mm, abaxially glabrous or with hairs at pinna bases; 2n⫽124 (Fla, Oax, Chis, Jam).

The non-adnate proximal pinnae, which are only slightly reduced, pinnatifid apices, and moderately to densely hairy rachises distinguish this species in the B. occidentale complex. See discussion under B. gracile, B. occidentale, and B. polypodioides for further differences. Several putative hybrids appear to involve B. appendiculatum. Blechnum appendiculatum ⫻ gracile: Chis (Breedlove 22456, DS, NY); Oax (Hallberg 1292, NY). In Mexico, this hybrid occurs at low elevations (300–700 m). Hybrids between B. appendiculatum and B. gracile have fertile fronds much narrower than the sterile, subconform blade apices, 8–10 pinna pairs, and malformed spores; they are known from Mexico, Central America (e.g., Guatemala: Bernoulli 456, NY), and South America, and occur at generally lower elevations than B. appendiculatum. Mickel and Beitel (1988) and Mickel (1992) cited several collections as representing the hybrid Blechnum appendiculatum ⫻ polypodioides: Jal (Mexia 1336, NY); Gro (Langlasse´ 671, US); Mor (Rose & Painter 7270, NY); and Ver (Purpus 1969, NY). At least some of these now seem to us like merely small specimens of B. appendiculatum, e.g., Mexia 1336 and Rose & Painter 7270. The Mexia collection in various herbaria has mostly well formed spores. Another collection that may indeed be this hybrid is Breedlove 44609(CAS), from Nayarit, 2100 m. It has 2–3 pairs of reduced proximal pinnae and non-sulcate costae adaxially, like B. polypodioides, but the pinnae are closely placed (even overlapping) and rounded at apices, more like B. appendiculatum; there is also a high percentage of malformed spores, 50%. Chromosome counts for B. appendiculatum indicate that it, like B. occidentale, is tetraploid. Both Mexican vouchers reported by Smith and Mickel (1977) as B. occidentale, from Chiapas and Oaxaca, we now identify as B. appendiculatum. See under B. occidentale for discussion of putative hybrids with that species.

2. BLECHNUM DIVERGENS Distribution. Terrestrial along roadsides and stream banks, in montane rain forests, pine-oak forests; (400–)700–2550 m. USA (Fla, Tex); Mexico; Guat, Hond, Salv, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Ec, Peru, Braz, Bol, Parag, Arg. Selected Specimens Examined. Chis (Breedlove 23310, DS, MEXU, NY). Col (Sanders 10366, UC). DF (Lyonnet 2125, DS, UC). Gro (Mexia 9047, CAS, UC). Hgo (Arreguı´n 1032, ENCB, MEXU). Jal (Iltis & Guzma´n M. 29082, UC). Me´x (Hinton 2572, NY). Mich (Hinton 13443, CAS, NY). Mor (Copeland herb. 48, UC). Nay (Ferris 5979, DS). Oax (Mickel 3745, NY, UC). Pue (Ramos & Ferna´ndez R-3, NY). Qro (Zamudio 6925, IEB). SLP (Harriman 12207, NY). Tam (Dressler 2020, UC). Tlax (Vela´zquez s.n., 9 Nov 1977, ENCB, IEB, MEXU). Ver (Purpus 1969, NY, UC).

Blechnum divergens (Kunze) Mett., Ann. Sci. Nat. Bot., se´r. 5, 2: 255. 1864. Fig. 54B. Lomaria divergens Kunze, Linnaea 9: 57. 1834. Type. Peru. Pampayaco, Poeppig s.n. [Diar. 1141] (LZ, destroyed). Lomaria exaltata Fe´e, Me´m. Foug. 11: 10, t. 3. 1866. Struthiopteris exaltata (Fe´e) Broadh., Bull. Torrey Bot. Club 39: 264. 1912. Type. Guadeloupe. 1861, L’Herminier s.n. (P; isotypes BM, NY!, P!–3 sheets).

Rhizomes erect, usually trunk-like, to 40 cm tall; rhizome scales brown to golden brown, concolorous, lanceolate, 7–15 ⫻ 1.5–2 mm, margins entire to ciliate; fronds strongly dimorphic, clumped; stipes 20–30 cm ⫻ 4–6 mm, dark red-brown to atro-

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purpureous, papillate, with matted, twisted, appressed hairs; sterile blades 50–80 ⫻ (16–)24–30 cm, pinnatifid, abruptly reduced at bases to vestigial lobes, apices tapering gradually, proximal pinnae contiguous to slightly distant; rachises tan to sometimes atropurpureous in the proximal portion; sterile pinnae adnate, falcate, 10–16 ⫻ 1.5–2 cm, apices acute, margins entire, slightly revolute, abaxially with narrow twisted tan scales and hair-like scales 1–2 mm long on costae and veins, adaxially glabrous; veins simple or 1-forked, vein endings enlarged and readily visible on adaxial surfaces; fertile pinnae 8–16 cm ⫻ 1.5–4 mm; indusia 0.5 mm wide, erose.

Rhizomes long-creeping, 5–10 mm diam., climbing; rhizome scales bicolorous, reddish brown, with narrow black central stripe, linear-lanceolate, 5–12 ⫻ 0.1–0.5 mm, margins with curved teeth 0.1 mm long or less, teeth directed both forward and backward; fronds dimorphic, distant, 80–120 cm long; stipes 1 ⁄5–1⁄3 the frond length, 4–22 cm ⫻ 3–5 mm long, glabrous, bicolorous, atropurpureous proximally, often becoming stramineous distally; sterile blades (40–)55–100 ⫻ (17–)22–30 cm, pinnatisect (entire to lobed in juvenile forms), largest proximal pinnae not connected to adjacent ones, blade bases abruptly reduced with 6⫹ pairs of greatly reduced auricles or vestigial pinnae, blade apices entire and with an acuminate tip; rachises bicolorous to concolorous, often atropurpureous in proximal part, glabrous, lacking aerophores at pinna bases; sterile pinnae 20–30 pairs, adnate, slightly falcate, (10–)12–20 ⫻ 1–1.8 cm, bases adnate their width or more, decurrent and surcurrent, apices long-acuminate, margins entire, often recurved, both sides glabrous, adaxially with deeply grooved and elevated costae distinct from laminae; veins distinct, simple or 1-forked, with enlarged vein endings visible abaxially; fertile pinnae 20–30 pairs, 8–14(– 20) cm ⫻ 2–4 mm, indusia 1 mm wide, entire.

Distribution. Terrestrial in wet forests at middle elevations; 1150–2300 m. Mexico; Guat, Hond, CR, Pan; Gr & L Ant; Col, Ven, Ec, Peru, Braz, Bol. Selected Specimens Examined. Chis (Breedlove 21456, DS, NY, 21512, DS, MEXU, NY, 23215, 31854, DS, 33698, DS, MEXU, MO, 32391, DS, MEXU). Gro (Lorea 2343, IEB, NY). Oax (Mickel 963, MEXU, NY, US).

This is likely to be confused only with B. lehmannii and B. wardiae in Mexico. Blechnum divergens has broader blades and longer pinnae, matted or scattered hair-like scales on the stipes and costae, and blade bases that are abruptly reduced (with vestigial pinnae below that only 1–2 mm long). Blechnum lehmannii has proximal pinnae that taper gradually to semicircular lobes at the blade bases, smaller, entire scales, and narrower blades. See B. wardiae for comparison with that species. Moran (in Davidse et al., 1995) mentioned a hybrid between B. divergens and B. wardiae from Costa Rica. Blechnum jamaicensis (Broadh.) C. Chr., from the Greater Antilles, has more gradually tapering pinnae proximally, generally larger fronds, and distant median and proximal pinnae.

3. BLECHNUM ENSIFORME Blechnum ensiforme (Liebm.) C. Chr., Index Filic. 153. 1905. Figs. 52B, 54C. Lomaria ensiformis Liebm., Mexic. Bregn. 234 (reprint 82). 1849. Struthiopteris ensiformis (Liebm.) Maxon, Contr. U.S. Natl. Herb. 13: 17. 1909. Type. Mexico. Oaxaca: Dist. Chinantla [Dist. Choapan], Teotalcingo, Liebmann s.n. [Pl. Mex. 2342] (C!–2 sheets; isotype US!).

Distribution. Hemiepiphytic or rarely epiphytic in wet forests at middle elevations, montane cloud forests; (600–)1000–2800 m. Mexico; Guat, Bel (Davidse & Holland 36781, UC), Hond, Salv, Nic, CR, Pan; Col, Ven, Guy, Ec, Peru, Bol. Selected Specimens Examined. Chis (Breedlove 22662, DS, MEXU, NY, 31708, DS, MEXU; Matuda 2913, K, MEXU, NY, US; Purpus 6741, NY, UC, US). Gro (Hinton et al. 14512, NY, US; Lorea 2994, IEB, MEXU). Oax (Hallberg 1715, ENCB, NY, UC; Mickel 4822, ENCB, NY, UC, US, 5651, MEXU, NY). Ver (Riba et al. 1173, ENCB, MEXU).

Blechnum ensiforme is most closely related to B. fragile, from which it differs in having much broader blades that are abruptly reduced proximally. Tryon and Stolze (1993) subsumed both B. ensiforme and B. fragile, as well as the related South America B. acutum (Desv.) Mett., under B. binervatum (Poir.) C. V. Morton & Lellinger, from the Lesser Antilles and South America, but we regard all four of these as distinct at species rank. Mickel and Beitel (1988: 82) cited several collections from Mexico (Mickel 991, 1090, 1442, 6561; Hellwig 345; Ventura A. 9365; all from Oaxaca except the last, all NY) and Central America as putative hybrids between B. ensiforme and B. fragile. Most

BLECHNUM FRAGILE

of these specimens now appear to us to be juvenile, sterile collections of B. ensiforme, which often has a very elongate terminal segment on juvenile fronds. The sole fertile collection of those they cited (Mickel 1090), and the one most likely of hybrid origin, is somewhat intermediate between the two species, but perhaps more like B. ensiforme in sterile blade shape (compare Mickel & Beitel, 1988, f. 123B with 125C); this collection does indeed have partially malformed spores, but a significant proportion of the spores appear normal.

4. BLECHNUM FALCIFORME Blechnum falciforme (Liebm.) C. Chr., Index Filic. 154. 1905. Figs. 52F–H. Lomaria falciformis Liebm., Mexic. Bregn. 234 (reprint 82). 1849. Struthiopteris falciformis (Liebm.) Broadh., Bull. Torrey Bot. Club 39: 365. 1912. Lectotype (chosen by Smith, 1981: 58). Mexico. Oaxaca: Cerro de Sempoaltepec, Liebmann s.n. [Pl. Mex. 2343, Fl. Mex. 775] (C!). Lomaria deflexa Liebm., Mexic. Bregn. 236 (reprint 84). 1849. Type. Mexico. Oaxaca: “Inter Totontepec et Trapiche de la Concepcio´n,” Liebmann s.n. [Pl. Mex. 2341, Fl. Mex. 770] (C!, frag. US!).

Differs from B. schiedeanum by: stipes with dense undercoating of matted, twisted, hair-like scales; rachises with scattered, brown scales 4–6 ⫻ 0.5–2 mm and dense undercoating of hairlike, fimbriate scales; sterile pinnae abaxially with dense scales 2–3 mm long and pale, hair-like scales on costae, also with appressed, twisted hair-like scales 4–5 mm long on veins and laminae abaxially, pinna margins cartilaginous, regularly serrulate, sinuses 0.5 mm deep.

145

Unverified, Doubtful, or Mistaken Reports. Tam (reported by Mickel & Beitel, 1988, but not verified).

See B. schiedeanum and key for comparisons.

5. BLECHNUM FRAGILE Blechnum fragile (Liebm.) C. V. Morton & Lellinger, Amer. Fern J. 57: 68. 1967. Fig. 52A. Lomaria fragilis Liebm., Mexic. Bregn. 232 (reprint 80). 1849. Blechnum binervatum (Poir.) C. V. Morton & Lellinger subsp. fragile (Liebm.) R. M. Tryon & Stolze, Fieldiana, Bot. 32: 64. 1993. Lectotype (chosen by Smith, 1981: 59). Mexico. Veracruz: Barranca de Huitamalco, Liebmann s.n. [Pl. Mex. 2321, Fl. Mex. 773] (C!; isolectotypes NY!, US). Osmunda polypodioides Sw., Prodr. 127. 1788. Blechnum onocleoides Sw., J. Bot. (Schrader) 1800(2): 75. 1801, nom. superfl.. Lomaria polypodioides (Sw.) Desv., Me´m. Soc. Linn. Paris 6: 288. 1827, hom. illeg., non Gaudich., 1825. Lomaria onocleoides (Sw.) Spreng., Syst. Veg., ed. 16, 4: 62. 1827. Blechnum polypodioides (Sw.) Kuhn, Filic. Afr. 92. 1858, hom. illeg., non Raddi, 1819. Type. Jamaica. Swartz s.n. (S; isotypes B, BM!, photo NY!). Lomaria mexicana Fe´e, Me´m. Foug. 8: 70. 1857. Syntypes. Mexico. Galeotti 6465 (P? not seen); Totutla, Mirador, Huatusco, Schaffner 100 (P? not seen).

Rhizomes long-creeping, climbing; rhizome scales bicolorous, uniformly pale brown or with a narrow, dark, central stripe, linear-lanceolate, 7–12 ⫻ 0.5–0.8 mm, with curved marginal teeth 0.1–0.3 mm long, directed both forward and backward; fronds strongly dimorphic, distant, 25–100 cm long; stipes 7–15(– 28) cm ⫻ 1–2 mm, 1⁄6–1⁄4 the frond length, atropurpureous proximally, sometimes becoming stramineous distally, glabrous; sterile blades chartaceous to subcoriaceous, pinnatisect, 40–65 ⫻ (2–)3–5(–8) cm, bases gradually reduced to deltate or semicircular lobes, blade apices pinnatifid to entire, acuminate; rachises atropurpureous to stramineous abaxially, glabrous or nearly so, lacking aerophores at pinna bases; sterile pinnae adnate, (1.5–)2–4 ⫻ 1 cm, obliquely triangular to falcate, apices acute, margins entire, glabrous on both sides; veins distinct, simple or 1-forked, with enlarged vein endings readily visible abaxially; fertile pinnae 4–6(–9) cm ⫻ 1–2 mm, indusia 0.5 mm wide, entire; 2n⫽58 (Jam).

Distribution. Terrestrial in wet forests at middle to upper elevations in Atlantic side, montane cloud forests, oak and beech forests, pine-oak forests; 1050–3150 m. Mexico; Guat, Hond, Salv, CR, Pan; Col, Ec. Selected Specimens Examined. Chis (Breedlove 21990, 22665, DS, MEXU, NY; Purpus 6739, NY, UC, US). Gro (Hinton et al. 14304, NY, US; Lorea 2785, IEB, MEXU). Hgo (Rzedowski 30211, ENCB, NY). Me´x (Hinton 3470, NY, UC, US). Oax (Mickel 3755, ENCB, MEXU, NY, UC; Pringle 5599, GH, LL). Pue (Lyonnet 470900009, MEXU). Ver (Ventura 7727, 9012, ENCB, MEXU, NY).

Distribution. Hemiepiphytic (or epiphytic) in wet forests of Atlantic side; 900–1950 m. Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Gr Ant; Col, Ven, Trin, Ec, Peru, Bol.

146

BLECHNUM

Selected Specimens Examined. Chis (Breedlove 23120, DS, MEXU, NY, 32220, 32602, DS, MEXU). Oax (Mickel 4304, MEXU, NY, 5691, MEXU, NY, UC; Torke et al. 518, ENCB, NY). Pue (Riba 17, ENCB, UAMIZ; Ventura A. 390, ENCB, NY). Ver (Copeland herb. 51, B, BM, MEXU, MICH, UC, US; Finck 101, UC).

The scandent rhizomes with distant, dimorphic fronds, rhizome scales with curved teeth, and abaxially enlarged vein endings distinguish B. fragile and B. ensiforme from other species of Blechnum in Mexico. The two species can be separated by the characters in the key. The B. fragile/ensiforme complex in the Neotropics needs reexamination. Blechnum fragile from Central America has bicolorous rhizome scales as does some material corresponding to B. fragile from the West Indies, whereas other specimens (from Puerto Rico and Hispaniola especially) have concolorous scales. The relationship of this to B. binervatum (Poir.) C. V. Morton & Lellinger of the Lesser Antilles and South America, which has concolorous rhizome scales, distant pinnae, wider blades, and abruptly reduced proximal pinnae, requires clarification, but we regard them as distinct species, contrary to Tryon and Stolze (1993).

6. BLECHNUM GRACILE Blechnum gracile Kaulf., Enum. Filic. 158. 1824. Type. Brazil. Otto s.n. (isotype B!). Fig. 53H. Blechnum intermedium Link, Hort. Berol. 2: 75. 1833. Type. Hort. Lips. (B!; isotype NY!). Blechnum subdimorphum Copel., Univ. Calif. Publ. Bot. 19: 288, t. 38. 1941. Type. Mexico. Veracruz: Co´rdoba, Copeland herb. 50 (MICH!–photos UC!, US!; isotypes P!, UC!, US!). Blechnum fraxineum auct., non Willd., Sp. Pl., ed. 4, 5(1): 413. 1810.

Rhizomes erect, short-creeping to ascending, stoloniferous; rhizome scales concolorous, brown to dark brown, narrowly lanceolate, 2–5 ⫻ 0.5–0.8 mm, margins with rare teeth; fronds monomorphic, the fertile often somewhat longer and more erect than sterile, clumped; stipes 1.8–22 cm ⫻ 1–2 mm, 1⁄3–2⁄3 the frond length, stramineous, smooth to obscurely papillate, with sparse scales orange or orange-brown scales at bases, these to 6 ⫻ 1.5 mm; blades 8–25 ⫻ 3–12 cm, 1-pinnate; rachises stramineous, glabrous, lacking aerophores at pinna bases; pinnae 2–4(–8) pairs, distant, proximal pinnae slightly reduced, apices conform, 1.5–3 times length of lateral pinnae, bases of terminal pinna often with 0–2 semicircular confluent to separate lobes (4–6 mm long), largest lateral pinnae 2–8 ⫻ 0.8–1.5 cm, bases rounded to broadly cuneate, terminal pinna 6–15 cm ⫻ 10–20 mm, pinna margins entire or nearly so, sometimes incurved, glabrous on both sides, costae grooved adaxially; veins simple to 1(–2)-forked, enlarged vein endings visible on adaxial surfaces; indusia tan, 0.3–0.5 mm wide, entire or subentire.

Distribution. Terrestrial on banks and along streams in wet forests of low to middle elevation, seasonal evergreen forests, oak forests; 20–920(–1150) m. Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Gr Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol, Parag, Arg. Selected Specimens Examined. Chis (Breedlove 31587, DS, MEXU, NY; Purpus 6738, NY, US). Gro (Hinton 11039, NY; Lorea 2416, IEB, MEXU, UC). Jal (McVaugh 26402, IEB, MEXU, MICH, NY). Nay (Te´llez 12849, MEXU). Oax (Mickel 6085, NY, UC, US, 7381a, MEXU, NY). Tab (Becerra s.n., 8 Apr 1914, MEXU). Ver (Purpus 2376, BM, MO, NY, UC; Ventura A. 12501, IEB, MEXU). Unverified, Doubtful, or Mistaken Reports. Qro (Ferna´ndez 4208, ENCB, cited by Arreguı´n et al., 2001, but not verified).

Among the monomorphic taxa, B. gracile is characterized by 1–3(–4) lateral pinna pairs, conform terminal pinnae 1.5–3 times the length of the lateral pinnae, blade apices often with subcordate bases and/or 1–2 semicircular lobes below the terminal pinna, glabrous rachises, and distant, glabrous pinnae with cuneate or rounded bases. Morphologically, B. gracile matches the tetraploid B. fraxineum sensu Walker (1966, 1973, 1985), who analyzed this entity cytologically from Trinidad. The name B. fraxineum Willd. has often been used for specimens of B. gracile in Mexico and elsewhere in the range, but in the strict sense the name B. fraxineum is believed to apply to sterile triploids between B. meridense Klotzsch (syns: B. longifolium Willd., non Cav.; B. schlimense Fe´e), a diploid taxon, and B. gracile (Mickel & Beitel, 1988). Blechnum meridense differs from B. gracile in having fertile and sterile fronds similar in size and shape (vs. narrower, longer fertile fronds in B. gracile), terminal pinna similar in size and shape to lateral pinnae (vs. much shorter and often rounded lateral pinnae in B. gracile), and terminal pinna cuneate proximally (vs. subcordate, with rounded lobes in B. gracile). Blechnum gracile evidently hybridizes with several other species in the complex. Some of these putative hybrids, as judged by their intermediate morphology between the suspected parents, are discussed below; see also under B. appendiculatum for discussion of hybrids with that species. Blechnum gracile ⫻ occidentale (⫽ B. ⫻caudatum; Fig. 55C): Chis (Breedlove 34096, DS; Martı´nez S. 8273, 8300, 8349, MEXU;

BLECHNUM MERIDENSE

Matuda 3612, LL, MEXU; Purpus 6738, UC). Gro (Langlasse´ 934, US). Oax (Cruden 1115, MEXU, UC; Herna´ndez G. 1986, CAS, CHAPA, MEXU, MO; Mexia 9181, CAS, LL, MO, NY, UC, 9181b, NY, UC; Mickel 6086b, NY, 6867b, 7381b, NY, UC). Pue (Lopez-Forment Conradt 119, MEXU). Ver (Calzada 918, CAS, MEXU; Ventura A. 15691, MEXU, 18933, ENCB, MEXU, XAL). Range (partially reported by Mickel & Beitel, 1988): s Mexico, Guat., Bel, Hond, CR; Cuba; Ven, Trin. In Mexico, this hybrid grows at low elevations (150–500 m) along both the Atlantic and Pacific slopes. It occurs in mixed collections with B. gracile (Mickel 7381b, 6086b), and with B. occidentale s.str. (Mexia 9181b, NY, UC; Mickel 6867b). Normal spore production has not been seen in any of the specimens cited, and in many of the putative hybrids the sporangia seem collapsed (e.g., Cruden 1115, UC; Mexia 9181, UC); Calzada 918 has the spores strongly malformed and collapsed. Hybrids between B. gracile and B. occidentale have fertile fronds narrower than the sterile. The name B. caudatum Cav., pro sp., Descr. Pl. 262, 1801, lectotype (designated by Christensen, 1937) from Ecuador, Ne´e s.n. (MA) (see discussion by Moran in Davidse et al., 1995: 326) apparently applies to this hybrid. See discussion under B.meridense for another hybrid, B. antillanum, of similar blade dissection but abaxially hairy along the rachises.

7. BLECHNUM LEHMANNII Blechnum lehmannii Hieron., Bot. Jahrb. Syst. 34: 473. 1904. Fig. 51D. Blechnum lherminieri (Bory) C. Chr. subsp. lehmannii (Hieron.) Lellinger, Amer. Fern J. 93: 146. 2003. Type. Colombia. Rı´o Timbiquı´, Lehmann 8928 (B; isotypes F, K!–3 sheets, US). Struthiopteris maxonii Broadh., Bull. Torrey Bot. Club 39: 268, t. 22. 1912. Blechnum maxonii (Broadh.) C. Chr., Index Filic., Suppl.1: 16. 1913. Type. Panama. Chiriquı´: Cerro de la Horqueta, Maxon 5415 (US!; isotype NY!).

Rhizomes ascending to usually erect; rhizome scales 4–6 ⫻ 1– 1.5 mm, brown, concolorous, lustrous, broadly deltate or ovate, entire; fronds strongly dimorphic, clumped; stipes 5–7 cm long, 1⁄8–1⁄4 the frond length, stramineous to dark purple or bicolorous, sometimes with dark brown scales resembling those of rhizomes (perhaps caducous), papillate; sterile blades thickherbaceous, 17–27 ⫻ 4–6(–9) cm, narrow, pinnatifid to pinnatisect, tapering gradually proximally, 2–3 proximal pinna pairs rounded, lacking vestigial pinnae below, blades becoming lobed to entire at the acuminate apices; rachises papillate, lacking aerophores at pinna bases; sterile pinnae 15–25 pairs, adnate, falcate, 1.5–5(–7) cm ⫻ 7–15 mm, apices obtuse, margins entire, sometimes slightly revolute, both surfaces glabrous; veins simple or 1forked, enlarged vein endings readily visible on adaxial surfaces; fertile pinnae 2–5(–7) cm ⫻ 1–2 mm; indusia tan to brown, ca. 0.5 mm wide, erose; 2n⫽396 (Hond).

147

Distribution. Montane cloud forests, in ravines, terrestrial and on summit of ridges; 1350–1750 m. Mexico; Guat, Hond, Salv, Nic, CR, Pan; Col, Ven, Ec, Peru, Braz, Bol. Selected Specimens Examined. Chis (Breedlove 22604, DS, MEXU, NY, 31854, DS, MEXU, 32391, DS, NY; Broome 713, DS). Gro (Lorea 3006, 3031, IEB). Oax (Hallberg 1538, NY; Torres C. et al. 11906, MEXU; Wendt et al. 4637, CAS, CHAPA, MEXU, NY).

Several recent authors (Mickel & Beitel, 1988; Moran in Davidse et al., 1995) have synonymized B. lehmannii under B. l’herminieri (Bory) C. Chr., type from Guadeloupe. The latter species has wider blades and the proximal pinnae more abruptly reduced. For now, we maintain the two as distinct. There is large variation in spore size in this species throughout its range; we suspect polyploid variation, and the single chromosome count, perhaps dodecaploid, is among the highest known for leptosporangiate ferns. Of Mexican species, B. lehmannii is most similar to B. divergens and differs from that in the much narrower blades with gradually reduced proximal pinnae. The pinnae are generally rounded at the tips (vs. usually acute in B. divergens). Breedlove 31854 was determined (by Smith, 1981) as atypical B. divergens, but is instead rather atypical B. lehmannii. Blechnum wardiae is also similar to B. lehmannii, but differs by having wider blades.

8. BLECHNUM MERIDENSE Blechnum meridense Klotzsch, Linnaea 20: 349. 1847. Type. Colombia [Venezuela]. Me´rida: Moritz 242 (B!, tracing NY!; isotypes BM!, NY!). Fig. 55A. Blechnum longifolium Humb. & Bonpl. ex Willd., Sp. Pl., ed. 4, 5(1): 413. 1810, hom. illeg., non Cav., 1802. Type. Venezuela. [Monagas.] Caripe, Humboldt 464 (B-Willd. 20045, fiche UC!; isotypes NY!, P).

Differing from B. gracile in the lateral pinnae Ⳳ equalling the terminal pinna in length, usually 2(–3) pairs, lateral and terminal pinnae narrowly cuneate proximally, sessile or slightly adnate, lacking confluent or separate semicircular lobes at the base of the terminal pinna; 2n⫽62 (Trin).

148

BLECHNUM

Distribution. Terrestrial in lower montane and montane forests; 800 m. Mexico; Pan; L Ant; Col, Ven, Trin.

Rhizomes erect, stoloniferous; rhizome scales 4–7 ⫻ 0.8–1 mm, bicolorous, central portion dark brown, outer margins pale brown, lanceolate, margins minutely serrulate or ciliate near apices; fronds clumped; stipes 17–28 cm, ca. 1⁄2 the frond length, stramineous, papillate, glabrous with scattered scales at bases; blades 17–28 ⫻ (8.5–)11–17(–22) cm, pinnate, lanceolate to ovatelanceolate, slightly or not at all reduced at bases, pinnae distant in proximal 2⁄3 of blade, apices gradually pinnatifid; rachises stramineous, papillate, glabrous, lacking aerophores at pinna bases; pinnae (4.5–)6–9(–12.5) ⫻ 0.7–1.8 cm, linear-lanceolate, subfalcate, bases subcordate to cordate and petiolate in proximal pinnae, apices acute, margins entire, both sides glabrous, adaxially with a row of minute intramarginal teeth, costae grooved; veins simple or 1-forked, enlarged vein endings visible adaxially; indusia tan, 0.8 mm wide, entire to minutely erose-ciliate; 2n⫽124 (Jam, Trin).

Specimens Examined. Chis (Breedlove 32982, 42005, DS, MEXU).

The two collections cited, especially Breedlove 42005, are indistinguishable from many specimens seen from Venezuela. Of Mexican species, B. meridense most closely resembles B. gracile and the hybrid between B. gracile and B. occidentale. On each blade there are two (to three) pairs of lateral pinnae and the terminal pinna is about the same size as the lateral ones. All the pinnae are narrowed and more or less narrowly cuneate at their bases. Blechnum meridense has usually been called B. fraxineum Willd., but as Mickel and Beitel (1988) discussed, the name fraxineum applies to the hybrid between B. gracile and B. meridense, and is not yet known from Mexico. Diploid B. meridense, although apparently absent itself in Jamaica, forms sterile triploid hybrids with B. occidentale s.l. in that country, and Trinidad, according to Walker (1966, 1973, 1985); these triploids can form large populations through vegetative reproduction by stolons. If this is true, the situation begs explanation. Hybrids between B. meridense and B. occidentale are distinguishable by having an intermediate number of pinna pairs (ca. 5–8), subconform blade apices almost as long as the lateral pinnae, fully pinnate blade bases, and distant median pinnae (thus eliminating B. polypodioides as a possible parent). These hybrids occur in South America and Costa Rica, whereas the pubescent hybrid B. appendiculatum ⫻ meridense (⫽ B. ⫻antillanum Proctor) occurs in Jamaica and South America.

9. BLECHNUM OCCIDENTALE Blechnum occidentale L., Sp. Pl. 2: 1077. 1753 (as B. orientale L., a species from southeast Asia), a transposition of names corrected in ed. 2 of Sp. Pl., 1763). Type. Based in part on an illustration in Petiver, Pter. Amer. t. 3, f. 9, 1712, which is a redrawing of t. 29B in Plumier, Traite´ Foug. Ame´r., 1705, illustrating a collection by Plumier from Hispaniola. LINN 1247.1 is this species, but according to Jackson (1912) was added to LINN after 1755. Petiver and Morrison illustrations were also cited. Figs. 53B,C, 54E.

Distribution. Terrestrial in wet forests, roadside banks, along streams in low to middle elevations, pine-oak forests, 50–1200 m. Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol, Parag, n Arg. Selected Specimens Examined. Chis (Breedlove 21722, CAS, MEXU, NY; Martı´nez & Ramos 26211, MEXU. Gro (Lorea 2807, MEXU). Mich (Hinton 16264, NY, UC). Nay (McVaugh 12131, MEXU, MICH). Oax (Mickel 5180, 6084, 6840, NY, UC). Pue (Go´mez-Pompa JB-753, MEXU; Sa´nchez-Mejorada 44, 622, MEXU). SLP (Rzedowski 10051 p.p., ENCB). Tab (Cowan 3068, ENCB, MEXU, NY). Ver (Purpus 1968, UC; Ventura 4998, ENCB, MEXU, NY). Unverified, Doubtful, or Mistaken Reports. Col (McVaugh 16096, NY, cited by Mickel, 1992, but not verified). Jal (cited by Va´zquez G. et al., 1995, but not verified, possibly misidentified). Qro (Rzedowski 42867, IEB, cited by Arreguı´n et al., 2001, but not verified). Tam (reported by Sharp, 1954, but the specimens are probably B. appendiculatum, as here circumscribed).

The B. occidentale complex, in Mexico comprising B. appendiculatum, B. gracile, B. meridense, B. occidentale s.str., and B. polypodioides, is a most difficult group in the Neotropics, both systematically and nomenclaturally. Walker (1966, 1973, 1985) has shown in Jamaica and Trinidad that numerous sterile hybrids exist, but “unlike the situation in most other genera, sterile interspecific hybrids are not always easy to detect on herbarium material using the presence of abortive spores as a criterion. The

BLECHNUM PENNA-MARINA

spores in this complex seem to be more or less colorless and the abortive ones do not seem to collapse to the extent found in many other fern hybrids.” Many authors (e.g., Murillo, 1968; Stolze, 1981) interpret species in this complex more broadly and as being extremely variable, failing to recognize the occurrence of hybrids. In light of cytological work by Walker in Jamaica and Trinidad, a narrower species concept seems more useful. However, acceptance of hybridization raises new questions— how to recognize and separate possible primary hybrids, allopolyploids, and backcross hybrids. By recognizing B. appendiculatum and B. occidentale as separate species and presenting some new interpretations in the B. gracile-meridense group, we provide hypotheses for further testing. The numerous names and types in this group make it difficult to assign the oldest name to the taxonomic entities, and some changes are likely, once taxonomic circumscription and relationships are understood better. In Mexico, B. occidentale s.str. occurs at 50–900 m, whereas B. appendiculatum occurs at 900–2550 m; this elevational difference is also true in Central America and probably in other parts of the ranges as well. The differences in size (as enumerated in our key and descriptions) appear correlated with differences in elevation and pubescence, and there are relatively few specimens that are difficult to place in one species or the other. Blechnum appendiculatum is much more common in the central, western, and northwestern parts of Mexico. Both B. occidentale and B. appendiculatum are characterized by fully pinnate proximal pinnae, distant pinnae in at least the proximal half of the blades, almost no or only slight reduction in the proximal pinnae, and pinnatifid blade apices. Hybrids with B. gracile can be detected by the smaller number of pinnae and the subconform terminal pinna, whereas hybrids with B. polypodioides (⫽ B. ⫻confluens; Fig. 55B) have proximal pinnae reduced and partially adnate (acroscopic portion of proximal pinna adnate but the basiscopic portion cordate), falcate pinnae that are adnate over the distal 2⁄3 of the blades, and narrow, linear blades. These hybrids and their ranges are discussed further under B. gracile and B. polypodioides. Hybrids between B. occidentale and B. polypodioides (Fig. 55B) have linear blades approaching those of B. polypodioides, but blade bases that are intermediate between the putative parents, broader (5–15 cm wide), pinnatifid blade apices, acuminate and falcate pinnae that are distant only in the proximal third of the blades, and adnate pinnae, except at the blade bases where pinnae are adnate on the acroscopic side and free and cordate on the basiscopic side (fully adnate in polypodioides, fully free and cordate in occidentale). Rachises are glabrous in the hybrids, a character by which they can be distinguished from B. appendiculatum ⫻ occidentale, discussed below. Specimens corresponding to B. occidentale ⫻ polypodioides are the following: Gro (Lorea 1905, MEXU; Lorea 4607, MEXU, UC); Nay (Alava & Cook 1619, UC; Jones 23478, UC; Lamb 591, DS; McVaugh 18848, NY; Mexia 535, CAS, DS, NY, UC); Ver (Bourgeau 1438, B, NY; Bourgeau 3601, UC p.p.; Copeland herb. 47, UC; Fisher 35373, NY; Lamb 591, DS; Purpus 6431, NY, UC, US). Some specimens of this hybrid appear to have well formed spores: Mexia 535. Mickel and Beitel (1988: 87) cited several additional collections

149

of this putative hybrid, some of which we now regard as other species or other hybrid combinations. Copeland herb. 47 (UC), also B. occidentale ⫻ polypodioides, was determined by Rojas as B. flaccisquama A. Rojas and said to occur from southern Mexico to Panama (type from Costa Rica). Two names based on Mexican types appear to be applicable to this hybrid: Blechnum ⫻aduncum Liebm., Mexic. Bregn. 1: 237 (reprint 85). 1849, pro sp. Type. Mexico. Veracruz: Huatusco, Liebmann s.n. (C); and Blechnum ⫻confluens Schltdl. & Cham., Linnaea 5: 613. 1830, pro sp. Type. Mexico. [Veracruz:] Hacienda de la Laguna, Schiede 779 (B!–2 sheets, photos US!, frag. NY!). Hybrids between B. occidentale and B. appendiculatum are also suspected, and Mickel and Beitel cited several collections from Oaxaca (Mickel 1020a, 4103, 5059, 5023, 6127, NY), and Chiapas (Gittins 4206, NY; Breedlove 29103, NY). Most of these are now judged to be B. appendiculatum because they bear a few hairs on the rachises abaxially, have relatively narrow blades, and grow at middle elevations, above 1350 m. Two abortive-spored specimens, one from Chiapas (Ghiesbreght 277, NY) and one from Guerrero (Hinton 14049, NY), are unusual in having very wide blades (22–24 cm), like B. occidentale, and densely hairy rachises, like B. appendiculatum. Blechnum ⫻falciculatum C. Presl (Epimel. Bot. 106. 1849 [1851]); Type. Mexico. Veracruz: Galeotti 6397 (BR!–3 sheets, photos BM, GH, US!; isotype K!) may apply to this broad-bladed, hairy form.

10. BLECHNUM PENNA-MARINA Blechnum penna-marina (Poir.) Kuhn, Filic. Afr. 92. 1868. Figs. 54F–H. Polypodium penna-marina Poir. in Lamarck, Encycl. 5: 520. 1804. Type. Straits of Magellan [Argentina? Chile?]. Commerson s.n. (P-Desf., photo GH).

Differs from B. stoloniferum by: fronds (both sterile and fertile) stiffly erect; blades subcoriaceous; sterile pinnae with costae grooved adaxially; fertile pinnae 30⫹ pairs, inequilateral and surcurrent at bases, arcuate, conduplicate, not apiculate at apices, of relatively uniform length from near blade bases to just short of blade apices.

Distribution. Terrestrial on seepage slopes along streams, conifer forests, pine-oak forests; 2300–2600 m. Mexico; Guat (Bea-

150

BLECHNUM

man 4203, UC); Peru, se Braz, Bol, Chile, Arg.; circumsubantarctic islands; se Australia, New Zealand. Specimens Examined. Chih (Correll & Gentry 23093, UC, US). Dgo (Maysilles 7925, NY).

This startling discovery, reported for the first time here, is remarkable for the nearly 4000 mile disjunction between southern Peru and northwestern Mexico. Previously, the specimen from Chihuahua had been regarded by everyone as B. stoloniferum (Knobloch & Correll, 1962; Smith, 1981; Mickel & Beitel, 1988). However, the two Mexican collections cited above differ from that in characteristics mentioned in the diagnosis above. The illustration of B. stoloniferum in Knobloch and Correll (1962: 134) is an excellent depiction of B. penna-marina, showing the numerous, spreading non-apiculate, conduplicate fertile pinnae. The specimens we cite from Durango and from Guatemala are sterile, and so many of the diagnostic characters are not determinable; however, in features of the sterile blades, they appear to be B. penna-marina, not B. stoloniferum. One can only ponder the means by which the Mexican populations became established.

11. BLECHNUM POLYPODIOIDES Blechnum polypodioides Raddi, Opusc. Sci. 3: 294. 1819, non (Sw.) Kuhn, 1868. Type. Brazil. Corcovado, Raddi s.n. (presumably FI–photo US; isotypes B, K, photos US; Tryon & Stolze, 1993, indicated the type might be in PI). Fig. 53D. Asplenium blechnoides Lag. ex Sw., Syn. Fil. 76. 1806. Blechnum unilaterale Sw., Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck. Gesammten Naturk. 4: 79, t. 3, f. 1. 1810, nom. superfl. Blechnum blechnoides (Lag. ex Sw.) C. Chr., Index Filic. 151. 1905, hom. illeg., non Keyserl., 1873. Type. Peru. Ne´e s.n. (S). Blechnum scabrum Liebm., Mexic. Bregn. 236 (reprint 84). 1849. Lectotype (chosen by Smith, 1981: 60). Mexico. Oaxaca: Teotalcingo [Choapan], District Chinantla, Liebmann s.n.[Pl. Mex. 2305, Fl. Mex. 272 (C!–photo NY!, frag. US!]. Blechnum angustifrons Fe´e, Me´m. Foug. 7: 25, t. 9, f. 2. 1857. Type. Mexico. [Oaxaca: Dto. Ixtla´n] Llano Verde, Galeotti 6383 (P?; isotypes BM, BR), the number cited by Martens and Galeotti (1842). Fe´e (Me´m. Foug. 7: 25. 1857), in error, cited Galeotti 6284 and 6440 as type numbers (corrected in Me´m Foug. 9: 4. 1857), but these two specimens are B. appendiculatum.

Rhizomes erect or ascending, stoloniferous; rhizome scales bicolorous, central portion or apices lustrous red-brown, outer or basal part pale brown, 3–5(–9) ⫻ 0.8–1 mm, lanceolate, margins entire; fronds 10–55 cm long, clumped; stipes stramineous, 2–10(– 15) cm ⫻ 1–2 mm, 1⁄10–1⁄3 the frond length, smooth to papillate, at bases with scattered pale brown scales 2–3 mm long; blades thin-chartaceous, 14–30(–40) ⫻ (3–)4–6(–10) cm, pinnatisect, narrowly elliptic, proximal pinnae partially or completely adnate, often deflexed, distant and irregularly deltate, blade apices pinnatifid; rachises stramineous, with sparse scales 0.8–13 ⫻ 0.1– 0.3 mm, otherwise either glabrous or with dense, sticky, septate hairs 0.3–0.5 mm long, lacking aerophores at pinna bases; pinnae 18–40 pairs, 1.5–3(–5) ⫻ 0.6–1 cm at bases, 3–4(–6) mm wide in

middle, linear-oblong, straight to often falcate, adnate, surcurrent and auriculate, apices acute, often apiculate, margins entire, abaxially the costae glabrous or hairy with sparse to dense septate hairs 0.1–0.3 mm, adaxially glabrous, with an intramarginal row of minute teeth, costae grooved to ungrooved; veins simple or 1-forked, with slightly enlarged vein endings visible adaxially; indusia red-brown, 0.5 mm wide, entire to erose; 2n⫽124 (Jam, Trin, Galapagos).

Distribution. Terrestrial or, rarely, epiphytic, on open or lightly shaded banks at middle elevations, pine-oak forests; 50– 1450(–3000) m. Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Galapagos, Peru, Braz, Bol, Parag, Arg. Selected Specimens Examined. Chis (Breedlove 32486, DS, ENCB, MEXU, NY; Purpus 6740, MO, UC, US). Gro (Lorea 2840, FCME, IEB, NY). Jal (s. coll. 21287, NY). Me´x (Sa´nchez 8, FCME). Nay (Breedlove 44308, CAS, MEXU; Te´llez 9430, ENCB, MEXU). Oax (Mickel 1096, ENCB, MEXU, NY, 3989, MEXU, NY). Pue (Rzedowski 29976, ENCB). Sin (Mexia 424, B, CAS, UC, US). Ver (Copeland herb. 49, BM, MEXU, MICH, NY, UC, US).

Blechnum polypodioides is distinguished from other monomorphic taxa by its narrowly elliptic blades that gradually taper at both ends, pinnatifid blade apices, greatly reduced, adnate, proximal pinnae, and relatively thin-chartaceous blades; a few proximal pinna pairs are distant and irregularly deltate. Stipes are short, mostly 2–10(–15) cm long, about 1⁄3–1⁄10 the frond length. Blades are variably pubescent, totally glabrous on the Atlantic slope and usually with hairy rachises and sometimes also laminar surfaces abaxially on the Pacific slope. The nomenclature of this species, confused with Blechnum polypodioides (Sw.) Kuhn [a synonym of B. fragile (Liebm.) C. V. Morton & Lellinger], was clarified by Morton and Lellinger (1967). Blechnum polypodioides is part of the Blechnum occidentale complex, and hybrids are suspected with both B. appendiculatum and B. occidentale (which see for discussion). Walker (1966, 1973, 1985), in studies in Jamaica and Trinidad, found the species (as B. unilaterale) to be tetraploid and to hybridize with B. occidentale s.l., producing sterile tetraploids that formed large colonies by vegetative reproduction.

BLECHNUM SERRULATUM

151

Blechnum asplenioides Sw., a close relative in South America, has even narrower blades and lacks stipes altogether, or nearly so; pending further study, it appears sufficiently distinct to maintain at species rank.

12. BLECHNUM SCHIEDEANUM Blechnum schiedeanum (Schltdl. ex C. Presl) Hieron., Hedwigia 47: 239. 1908. Figs. 52C–E. Lomaria schiedeana Schltdl. ex C. Presl, Suppl. Tent. Pterid. 143. 1836. Lomaria spectabilis Liebm., Mexic. Bregn. 235 (reprint 83). 1849, nom. superfl. Type. Mexico. Veracruz: Jalapa, Schiede [781] (isotypes B, HAL–photo K, frag. NY!, US!). Lomaria longifolia Schltdl. ex M. Martens & Galeotti, Me´m. Foug. Mexique 49. 1842, non Kaulf. (1824), hom. illeg. Type. Mexico. Veracruz: Totutla, Galeotti 6406 (BR, photos US; isotype K, photo US). Lomaria danaeacea Kunze, Linnaea 18: 326. 1844. Blechnum danaeaceum (Kunze) Christ in Pittier, Prim. Fl. Costaric. 3(1): 22. 1901. Syntypes. Mexico. [Veracruz:] Leibold 121 (isosyntype P!–photo UC!, US) and Leibold 122 (not seen). Lomaria acrodonta Fe´e, Me´m. Foug. 8: 70. 1857. Type. Mexico. [Veracruz:] “a` Huatusco et Totutla,” [1854] Schaffner 102 (P? not seen). Lomaria varians E. Fourn., Mexic. Pl. 1: 113. 1872. Blechnum varians (E. Fourn.) C. Chr., Index Filic. 161. 1905. Syntypes. Mexico. Veracruz: Orizaba, Botteri 1420 in Herb. Van Heurck (AWH?); “in Valle Cordovensi,” Bourgeau 1826 (BR!, K!, P!–6 sheets, NY!–2 sheets!).

Rhizomes large, woody, suberect; rhizome scales 10–30 ⫻ 2–10 mm, light brown, ovate to lanceolate, entire; fronds strongly dimorphic, to 200 cm long, clumped; stipes (0.5–) 25–80 cm ⫻ 3–7 mm, 1⁄3–1⁄2 the frond length, stramineous or less often purplish or dark-streaked, usually smooth, at bases with scattered scales resembling rhizome scales; sterile blades subcoriaceous to coriaceous, 26–100 ⫻ (10–)20–40 cm, 1-pinnate, proximal pinnae not reduced, terminal pinna conform; rachises stramineous, papillate or corrugate, bearing sparse ovate to lanceolate or hair-like, twisted scales 5–7 ⫻ 0.3–0.5 mm, tuberculiform aerophores at pinna bases; sterile pinnae (13–)18–30 pairs, free, straight to slightly falcate, 11–25(–36) ⫻ 1.8–3.2 cm, bases rounded to broadly cuneate, the largest stalked to 4 mm, apices longacuminate, margins entire to slightly undulate, serrulate at tips, recurved, abaxially with sparse to moderate tan scales 1–2 ⫻ 0.5– 1 mm and twisted, hair-like or linear scales on costae, adaxially glabrous; veins simple or 1-forked, enlarged vein endings visible adaxially; fertile pinnae 8.5–18 ⫻ 0.2–0.5 cm, abaxially with sparse scales 1–3 mm on costae; indusia brown, lacerate at maturity, ca. 1 mm wide.

Distribution. Terrestrial in montane forests, near streams; (350–) 750–1600(–2300) m. Mexico; Guat, Hond, Salv, Nic, CR, Pan. Selected Specimens Examined. Chis (Breedlove 22646, 31636, DS). Gro (Lorea 2103, IEB, MEXU). Hgo (Martı´nez 8, US). Me´x (Hinton 5252, NY, UC, US). Oax (Mickel 6167, ENCB, NY, UC, 6546, NY, UC). Pue (Rzedowski 32426, ENCB, IEB; Ventura A. 404, ENCB, IEB, MEXU, NY). Ver (Ventura 89, ENCB, NY, 9366, ENCB, IEB, MEXU, NY). Unverified, Doubtful, or Mistaken Reports. Jal (a specimen labelled as from Jalisco, Espero´n M. 16722, UC, is mistakenly labelled, and actually from Edo. Veracruz).

Blechnum schiedeanum and B. falciforme are occasionally difficult to separate. The main distinguishing characters are the presence in B. falciforme of cartilaginous serrations along the whole pinna margins, dense scales 2–3 ⫻ 1–1.5 mm along the costae of the sterile and fertile pinnae, and appressed, twisted, hair-like scales 4–5 mm long on veins and laminae. Blechnum varians, often regarded as a distinct species (e.g., by Smith, 1981, Mickel & Beitel, 1988; Moran in Davidse et al., 1995), is virtually identical to B. schiedeanum except that the stipes and rachises are dark purplish. It seems likely that pigmentation on the axes of this species is a response to increased exposure to light, and the pigmented variants seem to occur sporadically throughout the range of B. schiedeanum. Blechnum cordatum (Desv.) Hieron., widespread and common in South America and with many synonyms, is very closely related to B. schiedeanum and perhaps not distinguishable. Monographic study is needed on this group of Blechnum before names can be applied with confidence.

13. BLECHNUM SERRULATUM Blechnum serrulatum Rich., Actes Soc. Hist. Nat. Paris 1: 114. 1792. Type. French Guiana. Leblond s.n. (P, photo US). Figs. 53E–G. Rhizomes creeping or the apices ascending, 5–10 mm diam., lacking stolons; rhizome scales concolorous and dark brown, to somewhat bicolorous with blackish central streaking, lanceolate,

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BLECHNUM

4–8 ⫻ 0.3–0.8 mm, margins entire; fronds monomorphic or weakly dimorphic (pinnae of fertile blades slightly narrowed), clumped; stipes (10–)20–40 cm ⫻ 3–5 mm, 1⁄4–1⁄2 the frond length, tan, smooth, glabrous; blades chartaceous to subcoriaceous, 50–120 ⫻ 10–30 cm, oblong to ovate, bases truncate and lacking vestigial pinnae, subabruptly or abruptly tapered at the conform apices, 1-pinnate, terminal pinna similar in shape to lateral pinnae; rachises tan to stramineous, smooth, glabrous, lacking aerophores at pinna bases; pinnae 25–40 pairs, linearoblong, 5–15 cm ⫻ 5–15 mm, sessile or stalked to 2 mm, not adnate, equilateral, articulate (distinct articulate line at bases) and deciduous, apices acute, margins serrulate, not revolute, abaxially with small tan scales 0.5–1 ⫻ 0.5–8 mm long on costae, adaxially glabrous; veins simple or 1-forked, vein endings not enlarged or conspicuous, fusing with strongly cartilaginous whitish margins; indusia 0.5 mm wide, entire to erose.

14. BLECHNUM STOLONIFERUM Blechnum stoloniferum (Mett. ex E. Fourn.) C. Chr., Index Filic. 154. 1905. Figs. 51C, 54J–L. Lomaria stolonifera Mett. ex E. Fourn., Mexic. Pl. 1: 113. 1872. Struthiopteris stolonifera (Mett. ex E. Fourn.) Broadh., Bull. Torrey Bot. Club 39: 277. 1912. Types. Mexico. Veracruz: Orizaba, Mu¨ller 61.1491, Meissner Herbarium (isotypes B!, K!, NY!). Lomaria ghiesbreghtii Baker in Hooker & Baker, Syn. Fil., ed. 2, 481. 1874. Blechnum ghiesbreghtii (Baker) C. Chr., Index Filic. 154. 1905. Type. Mexico. Chiapas: Ghiesbreght 207 (K!; isotypes BM!, K!, NY!, PH!, NY!).

Rhizomes horizontal, compact and stoloniferous; rhizome scales 3–4 ⫻ 0.8–1 mm, light brown, deltate, entire; fronds strongly dimorphic, clumped, 10–35 cm long, fertile fronds taller than sterile; stipes of sterile fronds 1–6(–12) cm ⫻ 1 mm, 1⁄10–2⁄5 the frond length, atropurpureous, glabrous or the bases sparsely scaly, smooth to papillate; sterile blades pinnatifid to pinnatisect, 8–22 ⫻ (1–)2–3.5(–5) cm, chartaceous, narrowly elliptic, bases gradually reduced to distant semicircular lobes, apices gradually tapered and pinnatifid; rachises smooth or papillate, lacking scales, lacking aerophores at pinna bases; sterile pinnae 20–27 pairs, broadly adnate, oblong, mostly 1–2(–3) ⫻ 0.6–0.8 cm, somewhat falcate, apices rounded or infrequently acute, margins entire or faintly crenulate to undulate, often recurved, glabrous on both sides; veins simple to 1-forked, vein endings enlarged and readily visible on adaxial surfaces; fertile pinnae 8–18 ⫻ 0.8–2 mm; indusia 0.6–0.8 mm, subentire to erose or minutely ciliate.

Distribution. Terrestrial, wet ditches, swamps at low elevations; 0–900 m. USA (s Fla); Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Bah, Gr & L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol, Parag, n Arg; Malesia, Australia. Selected Specimens Examined. Chis (Breedlove 22219, DS, MEXU, NY; Miranda 8497, MEXU). QR (Castillo Rivero 201, XAL; Goldman 625, US). Tab (Cowan 2503, CAS, ENCB, MEXU, NY; Gliessman et al. 1700, ENCB, NY; Matuda 3088, MEXU, US; Rovirosa 433, NY, US. Ver (King 985, ENCB; Lot 1594, ENCB, MEXU, MO, UAMIZ, UC).

This species differs markedly from other Mexican species of Blechnum in its lack of visibly enlarged vein endings (fusing with the cartilaginous margins instead), articulate pinnae, narrowly spaced veins, and relatively widely spaced teeth. The sori are often confined to the distal pinnae. Mickel and Beitel (1988: 89) cited B. serrulatum from Yucata´n, perhaps on the basis of Goldman 625, labelled “Plants of Yucatan, Mexico, Puerto Morelos”; however, this locality is in Quintana Roo. The name Blechnum indicum Burm. f., in recent years considered to apply to Asplenium longissimum Blume, has been recently neotypified by Chambers and Farrant (in McCarthy, 1998: 710) by Coveny 4712 (NSW; isoneotype UC!, among others). With this typification, it is very closely related to B. serrulatum. The differences were discussed by Chambers and Farrant (in McCarthy, 1998: 374). Its range is in Southeast Asia, Australia, and Polynesia.

Distribution. Terrestrial in montane cloud forests, evergreen cloud forests, pine-oak forests; 1600–3250 m. Mexico; Guat, Hond, Salv, CR. Selected Specimens Examined. Chis (Breedlove 32053, DS, MEXU, NY). Dgo (Bashor 695, NY). Gro (Lorea 1156, FCME, MEXU). Hgo (Pringle 8752, MEXU, NY, UC, US). Me´x (Hinton 7222, ENCB, NY, US). Mich (Rzedowski 46193, ENCB, IEB, MEXU). Mor (Lyonnet & Cha´vez 3376, MEXU, US). Oax (Mickel 4661, ENCB, MEXU, NY, UC). Pue ( Pringle s.n., 6 Oct 1909, CAS, US). Ver (Herna´ndez M. 1571, ENCB, MEXU).

The small frond size, stoloniferous nature, and occurrence at higher elevations distinguish B. stoloniferum from the other members of sect. Lomaria in Mexico (B. divergens, B. lehmannii, and B. wardiae).

BOLBITIS

15. BLECHNUM WARDIAE Blechnum wardiae Mickel & Beitel, Mem. New York Bot. Gard. 46: 89. 1988. Type.Mexico. Oaxaca: Distrito Ixtla´n, 8 km S of Vista Hermosa, 70 km N of Ixtla´n de Jua´rez on Rte 175, Mickel 6749 (NY!; isotypes ENCB!, NY!, UC!, US!). Figs. 51A,B, 54A. Rhizomes erect, trunk-like, to 30 cm tall; rhizome scales concolorous, brown, lanceolate, 10–20 ⫻ 2–4 mm, margins entire; fronds strongly dimorphic, clumped; stipes 1–4 cm ⫻ 2–4 mm, atropurpureous, papillate; sterile blades 40–80 ⫻ 12–15 cm, pinnatifid, gradually reduced at bases to semicircular lobes, proximal pinnae contiguous, blade apices tapering gradually; rachises papillate, lacking aerophores at pinna bases; sterile pinnae adnate, slightly falcate, 6–8 cm ⫻ 12–15 mm, medial pinnae acute, proximal pinnae and lobes rounded, margins entire, glabrous on both sides; veins simple or 1-forked, vein endings enlarged and conspicuous on adaxial surfaces; fertile pinnae 5–10 cm ⫻ 1–1.5 mm; indusia 0.5 mm wide, entire.

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between these two species, but B. stoloniferum is a much smaller species and seems less intimately related, so this hypothesis seems unlikely. Breedlove 32391 from Chiapas was previously cited as B. divergens by Smith (1981). Unplaced Names Blechnum triangulare Link, Hort. Berol. 2: 76. 1833. Type. Mexico. Hort. berol., authentic specimens in GH, L–photo GH, and US, fide Murillo (1968). Murillo (1968) placed this in synonymy under B. confluens, which we regard as the probable hybrid Blechnum occidentale L. ⫻ polypodioides Raddi.

19. BOLBITIS Bolbitis Schott, Gen. Fil., t. 14. 1834. Lectotype (chosen by Christensen, Index Filic. XXVI. 1906): Bolbitis serratifolia (Kaulf.) Schott [⬅ Acrostichum serratifolium Kaulf.]. For other synonyms, see Hennipman (1977).

Terrestrial or hemiepiphytic; rhizomes creeping, scaly; fronds medium-sized, erect, dimorphic, the fertile with contracted pinnae or lobes and proportionately longer stipes; stipes glabrous or sparsely scaly, not articulate to rhizomes; blades simple to pinnate, less commonly to bipinnatifid, thin to subcoriaceous, glabrous; veins areolate, with or without included free veinlets; proliferous buds present at bases or tips of blades in some species; sporangia usually covering the abaxial surfaces of fertile blades; indusia and paraphyses absent; spores bilateral, each with winged or rugose perispore; x⫽41.

Distribution. Terrestrial in wet forests; (800–)1500–2300 m. Mexico; Salv, Nic, CR, Pan; w Col, w Ec. Selected Specimens Examined. Chis (Breedlove 32391, DS, MEXU, 49315, CAS, ENCB, NY). Oax (Hallberg 1542, NY, UC; Mickel 4262, ENCB, MEXU, NY, UC, US, 5578, ENCB, 6749, ENCB, NY, UC). Pue (Lyonnet 550500004, MEXU). Ver (Bohs et al. 1698, XAL; Riba 518, XAL; Ventura A. 7418, ENCB, MEXU, NY).

This species is distinct by the combination of relatively large size (sterile fronds 12–15 cm wide, pinnae 6–8 cm long, blades 40–80 cm long), gradual reduction of the proximal pinnae to semicircular lobes that run down nearly to the rhizomes, and relatively large rhizome scales 10–20 mm long. Blechnum wardiae resembles most closely B. divergens and B. lehmannii. Mickel and Beitel (1988) postulated that it was intermediate between B. divergens and B. stoloniferum, and possibly of allopolyploid origin

There are about 45 species in the genus, which is pantropical; about a third of the species occur in the Neotropics. Eight species are known in Mexico, and all, except B. portoricensis, are restricted to southern Mexico. Bolbitis species in Mexico occur in lowland to middle elevation wet forests and are uncommon or rare. The genus is distinct in its areolate venation, dimorphic fronds, and sporangia covering the abaxial surfaces of fertile blades. Unpublished molecular data by Cranfill indicate that Bolbitis is a member of the dryopteroid clade, with chromosome base number of x⫽41, and probably allied to Elaphoglossum and several other genera that show tendencies to be epiphytic or hemiepiphytic, e.g., Lomagramma, Cyclodium, Maxonia, Polybotrya, and Olfersia. Reference Hennipman, E. 1977. A monograph of the fern genus Bolbitis (Lomariopsidaceae). Leiden Bot. Ser. 2: i–xii, 1–331.

Key to the Mexican Species of Bolbitis 1. Plants hemiepiphytic, rhizomes climbing on trees; rhizome scales denticulate. ..................................................................................... 2. B. bernoullii. 1. Plants terrestrial, rhizomes not climbing on trees; rhizome scales entire. 2. Blades pinnatifid, with subterminal bulblets on the pinnae; free included veins present. .................................................................. 3. B. hastata.

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BOLBITIS 2. Blades pinnate (at least at bases), nonproliferous or with bulblets at intervals along rachises or with flagelliform, rooting apices; free included veins present or absent. 3. Blade apex similar to lateral pinnae. 4. Blades proliferous with a bud at base of each terminal pinna. .................................................................................................. 4. B. hemiotis. 4. Blades nonproliferous, buds absent. 5. Venation pattern with both recurrent and excurrent included free veins in vein areoles. ...................................... 5. B. pergamentacea. 5. Venation pattern with only recurrent included free veins in the vein areoles. .............................................................. 7. B. serratifolia. 3. Blade apex pinnatifid, dissimilar to lateral pinnae. 6. Free included veins present, areoles small, numerous; blade apices nonproliferous or with bulblets at intervals along rachises of flagelliform apices. ................................................................................................................................................... 6. B. portoricensis. 6. Free included veins absent, areoles large, few; blade apices nonproliferous. 7. Pinna margins lacking teeth in the sinuses (if present); free pinnae usually 1 pair per blade; blades drying dark green or blackish. ..................................................................................................................................................................................... 1. B. aliena. 7. Pinna margins with teeth in sinuses; free pinnae to 5 pairs per blade; blades green when dried. ................................. 8. B. umbrosa.

1. BOLBITIS ALIENA Bolbitis aliena (Sw.) Alston, Kew Bull. 1932: 310. 1932. Figs. 57A–C. Acrostichum alienum Sw., J. Bot. (Schrader) 1800(2): 13. 1801. Gymnopteris aliena (Sw.) C. Presl, Suppl. Tent. Pterid. 244. 1836. Leptochilus alienus (Sw.) C. Chr., Bot. Tidsskr. 26: 285. 1904. Type. Jamaica. Collector not stated [Swartz s.n.] (S; isotype S). For additional synonymy, see Hennipman (1977: 132).

Rhizomes long-creeping, mostly 3–4 mm diam.; rhizome scales brown, lanceolate, 4–7 ⫻ 1–1.5 mm, entire; fronds distant, 25– 65(–90) cm long; stipes 15–45 cm ⫻ 1.5–3 mm, those of sterile fronds ca. 1⁄2–2⁄3 the sterile frond length, those of fertile fronds ca. 3⁄4 the fertile frond length; sterile blades thin-chartaceous, deltate, 15–45(–60) ⫻ 15–30 cm, pinnate, apices pinnatifid, nonproliferous; sterile pinnae 1–2 free pairs ⫹ 2–7 adnate and strongly decurrent pairs, margins crenate-serrate or shallowly lobed, sometimes cut 1⁄2 way to costa, basiscopic side exaggerated on proximal pair, regularly lobed or few conspicuous lobes; veins areolate, areoles lacking included veinlets or with only few, short, simple, free veinlets; fertile blades pinnate, 10–16 ⫻ 5–12(–20) cm; sporangia nearly covering the abaxial surfaces of fertile blades; 2n⫽164 (Jam).

Distribution. Terrestrial on rocks or ground in wet forest; 700– 1000 m. Mexico; Hond, Nic, CR, Pan; Gr & L Ant; Col,

Ven, Trin, Ec, Peru, Bol. Also cited from El Salvador by Seiler (1982: 389). Unverified Specimens. Chis (Ghiesbreght 409, K, cited by Hennipman, 1977, but not verified; Ghiesbreght 409, GH, is cited by Hennipman, 1977 as Bolbitis serratifolia). Ver (Bourgeau 2147, L, not seen, cited by Hennipman, 1977, but not verified).

The nonproliferous pinnatifid blade apices and large areoles without free included veins (or a few, short, unforked included veins) distinguish this species from B. portoricensis, with which it has been confused. An even closer relative is B. umbrosa, which differs by the presence of a small tooth in sinuses of sterile blades. The apparent mixed collection of Ghiesbreght 409, as cited by Hennipman (1977), suggests that the identity of specimens of this number in herbaria needs to be re-examined. Breedlove 21621, 22483, and 30866, cited by Smith (1981: 62) as this species, are all B. umbrosa.

2. BOLBITIS BERNOULLII Bolbitis bernoullii (Kuhn ex Christ) Ching in Christensen, Index Filic., Suppl. 3: 47. 1934. Figs. 58F, G. Acrostichum bernoullii Kuhn ex Christ, Bull. Soc. Roy. Bot. Belgique 35(1): 244. 1896. Gymnopteris bernoullii (Kuhn ex Christ) Diels in Engler & Prantl, Nat. Pflanzenfam. 1(4): 201. 1899. Leptochilus bernoullii (Kuhn ex Christ) C. Chr., Bot. Tidsskr. 26: 296. 1904. Lectotype (chosen by Christensen, Bot. Tidsskr. 26: 296. 1904). Guatemala. Costa Grande, between Escamillas and Palahucco, Bernoulli & Cario 382 [282] (isotype B, frags. BM, NY!). Gymnopteris tuerckheimii Christ, Bull. Herb. Boissier, se´r. 2, 6: 290. 1906. Leptochilus tuerckheimii (Christ) C. Chr., Index Filic., Suppl. 1: 48. 1913. Type. Guatemala. Alta Verapaz: Cubilquitz, Tu¨rckheim 8831 (US!; isotype US!). Gymnopteris donnell-smithii Christ, Bull. Herb. Boissier, se´r. 2, 6: 289. 1906. Leptochilus donnell-smithii (Christ) C. Chr., Index Filic., Suppl.1: 48. 1913. Bolbitis donnell-smithii (Christ) Ching in C. Christensen, Index Filic., Suppl. 3: 48. 1934. Type. Guatemala. Alta Verapaz: Cubilquitz, Tu¨rckheim 8830 (US; frag. NY!).

Rhizomes long-creeping, climbing, to 3 mm diam.; rhizome scales dense, brown, 10–20 ⫻ 0.5–1.5(–3) mm, denticulate; fronds distant or subdistant, 60–135 cm long, distant or subdistant; stipes 14–40 cm ⫻ 3–6 mm, those of sterile fronds ca. 1⁄6–1⁄3 the sterile frond length, those of fertile fronds ca. 1⁄5–2⁄5 the fertile frond length, sparsely scaly; sterile blades chartaceous, deltate-

BOLBITIS HEMIOTIS

155

lanceolate, 35–110 ⫻ 15–50 cm, once-pinnate, apices Ⳳ conform, without buds; sterile pinnae to ca. 15 pairs, proximal pinnae stalked to ca. 10 mm, bases rounded to cuneate, apices acuminate, margins crenate to very shallowly lobed; veins areolate, areoles with 1–3 excurrent, usually free veinlets; fertile blades oncepinnate, 20–70 cm long; sporangia covering the abaxial surfaces of fertile blades.

Distribution. Terrestrial or epipetric in wet forests of lower middle elevation of the Atlantic slope; 150–1350 m. Mexico; Bel (Holst 5757, UC), CR. Selected Specimens Examined. Chis (Ghiesbreght 370, BM, GH, K). Oax (Mickel 5903, ENCB, MEXU, US; 7192, ENCB, MEXU, UC, US; Hallberg 1597, 1616, US). Ver (Wendt et al. 3853, CHAPA, MEXU, NY, UC; Ventura A. 15149, IEB, MEXU).

Distribution. Hemiepiphytic in lowland rain forests; 100–1000 m. Mexico; Guat, Bel, Hond, Nic, CR, Pan; Col, Ec. Selected Specimens Examined. Chis (Calzada 2952, XAL; Purpus 7246, MO, NY, US; Ton 5972, MEXU). Oax (Herna´ndez G. 1187, 1789, 2066, 2551, CHAPA, NY; Torres C. 11181, MEXU). Ver (Bourgeau 2008, NY, US).

The hemiepiphytic habit and strongly denticulate rhizome scales are unique among Bolbitis species in Mexico.

3. BOLBITIS HASTATA Bolbitis hastata (E. Fourn.) Hennipman, Amer. Fern J. 65: 30. 1975. Figs. 58H, J, K. Acrostichum hastatum Liebm., Mexic. Bregn. 172 (reprint 20). 1849, hom. illeg., non Thunb., 1784. Gymnopteris hastata E. Fourn., Mexic. Pl. 1: 70. 1872. Leptochilus liebmannii Maxon, Proc. Biol. Soc. Wash. 43: 86. 1930, nom. superfl. Bolbitis liebmannii (Maxon) C. Chr., Index Filic., Suppl. 3: 49. 1934. Type. Mexico. Veracruz: Arroyo de Isapa, Barranca de Jovo, near Sta. Marı´a Tlapacoyo, Liebmann s.n. [Pl. Mex. 2446] (C!; B, frag. NY!, K, US!).

Rhizomes creeping, 3–5 mm diam.; rhizome scales dark brown, lustrous, 0.5–1.5 mm long, entire; fronds subdistant, 35–95 cm long; stipes 15–45 cm ⫻ 2–4 mm, those of sterile fronds ca. 1⁄3– 1 ⁄2 the sterile frond length, those of fertile fronds to ca. 3⁄4 the fertile frond length, with scattered scales; sterile blades chartaceous, deltate, mostly 20–50 ⫻ 16–25 cm, pinnatifid, cut 3⁄4–9⁄10 the way to the rachises, with 3–5(–8) pairs of lobes, the confluent apical lobe 1⁄2 the blade length, acuminate, the blade and lobe tips often with a small bud on the adaxial surface 5–10 mm from tips; sterile lobes entire to irregularly undulate, proximal pair of lobes decurrent, acuminate at the tips; veins areolate, areoles with mostly recurrent, branched, free veinlets; fertile blades pinnatifid, 9–20 ⫻ 6–12 cm; sporangia covering the abaxial surfaces of fertile blades.

The pinnatifid fronds with free, included veinlets, entire rhizome scales, and subterminal laminar buds distinguish B. hastata from other species of Bolbitis in Mexico.

4. BOLBITIS HEMIOTIS Bolbitis hemiotis (Maxon) Ching in Christensen, Index Filic., Suppl. 3: 48. 1934. Figs. 58C–E. Leptochilus hemiotis Maxon, Amer. Fern J. 14: 101. 1925. Type. Trinidad. Fendler 101 (US!; isotypes B, BM, G, K, MICH!, NY!–4 sheets, S, UC!, US!).

Rhizomes creeping, to 10 mm diam.; rhizome scales dark brown, 5–10 ⫻ 1–3 mm, entire; fronds subdistant, 40–115 cm long; stipes 15–65 cm ⫻ 3–8 mm, those of sterile fronds ca. 1⁄2 the sterile frond length, those of fertile fronds ca. 2⁄3 the fertile frond length, sparsely scaly; sterile blades firmly chartaceous, ovate, 25–60 ⫻ 15–45 cm, pinnate, with conform terminal pinna, proliferous buds in axils of the distal pinnae; sterile pinnae (1– )2–3(–4) pairs, 20–30 ⫻ 6–15 cm, entire to undulate, proximal pinnae sessile to stalked to 10 mm, broadly elliptic or unequally rhomboid, at bases broadly cuneate acroscopically, cordate to rounded basiscopically, acute at tips; veins irregularly and finely areolate, areoles with mostly recurrent, branched, free veinlets; fertile blades once-pinnate, 20–35 cm long; sporangia covering the abaxial surfaces of fertile blades.

156

BOLBITIS

Distribution. Terrestrial in wet lowland and lower montane forests; 100–1000 m. Mexico; Bel, CR, Pan; Col (Forero et al. 6649, MO), Ven, Trin, Ec. Specimens Examined. Oax (Herna´ndez G. 1746, CHAPA, MEXU). Ver (Cedillo T. 3314, MEXU; Lira 103, MEXU, XAL; Nee & Taylor 29956, MO, UC; Wendt 3847a, CHAPA, NY, UC).

Distribution. Wet lowland forests; 200–600 m. Mexico; Guat, Bel, Hond, Nic, CR, Pan; Gr Ant; Col, Ven, Ec. Also cited from Peru by Hennipman & Moran (in Davidse et al. 1995: 249). Selected Specimens Examined. Chis (Breedlove 57295, CAS; Martı´nez S. 13468, MEXU; Martı´nez S. 25349, 25890, MEXU). Oax (Herna´ndez G. 1005, 2086, CHAPA, NY, 1567, 2071, CHAPA). Ver (Martı´nez C. 2189, CHAPA; van Rooden 757, NY).

The pinnate fronds with a conform terminal pinna, 3–9 alternate pinnae with strongly asymmetric bases, included free veinlets, and axillary proliferous buds distinguish the species from all others in the New World.

The lack of axillary buds and pinnae with more or less symmetric bases distinguish this species from B. hemiotis.

5. BOLBITIS PERGAMENTACEA

6. BOLBITIS PORTORICENSIS

Bolbitis pergamentacea (Maxon) Ching in Christensen, Index Filic., Suppl. 3: 49. 1934. Figs. 58A, B.

Bolbitis portoricensis (Spreng.) Hennipman, Amer. Fern J. 65: 30. 1975. Figs. 56A–C.

Leptochilus pergamentaceus Maxon, J. Wash. Acad. Sci. 14: 144. 1924. Type. Jamaica. Green River Valley, Underwood 1426 (US!–2 sheets).

Acrostichum portoricense Spreng., Nova Acta Phys.-Med. Acad. Caes. Leop.Carol. Nat. Cur. 10: 226. 1821. Gymnopteris portoricensis (Spreng.) Fe´e, Me´m. Foug. 2: 85. 1845. Type. Puerto Rico. Bertero s.n. (G; isotypes P, TO). Acrostichum cladorrhizans Spreng., Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 10: 225. 1821. Leptochilus cladorrhizans (Spreng.) Maxon, Pteridophyt. Porto Rico 460. 1926. Bolbitis cladorrhizans (Spreng.) Ching in Christensen, Index Filic., Suppl. 3: 47. 1934. Type. Puerto Rico. Bertero s.n. (G; isotypes P, TO). Acrostichum irregulare Liebm., Mexic. Bregn. 173. (reprint 21) 1849. Type. Mexico. [Veracruz:] Barranca de Sta. Marı´a Tlatetla el Fortı´n, Liebmann s.n. [Pl. Mex. 2688, Fl. Mex. 666, 667] (C–folio herb!; isotype B).

Rhizomes long-creeping, to 10 mm diam.; rhizome scales brown, ca. 6 ⫻ 4 mm, entire; fronds subdistant, 50–140 cm long; stipes 20–80 cm ⫻ 3–10 mm, those of sterile fronds ca. 1⁄2 the sterile frond length, those of fertile fronds nearly 2⁄3 the fertile frond length, with scattered scales; sterile blades firmly chartaceous, ovate to elliptic, 25–60 ⫻ 20–50 cm, pinnate, each blade with a conform terminal pinna, not proliferous; sterile pinnae (0–)2–6 pairs, proximal pinnae stalked, somewhat unequally elliptic, with cuneate bases and acuminate apices, margins entire to obscurely undulate; veins finely areolate, areoles with excurrent and recurrent, often branched, free veinlets; fertile blades once-pinnate, 12–45 cm long, with 1–3 pairs of lateral pinnae and a slightly larger terminal pinna; sporangia covering the abaxial surfaces of fertile blades; 2n⫽82 (Jam).

Rhizomes short-creeping, 8–15(–20) mm diam.; rhizome scales reddish brown, to 10 ⫻ 2 mm, entire; fronds clumped, 40–140 cm long, the fertile ca. 2⁄3 as long as the sterile; stipes 4–60 cm ⫻ 2–6 mm, those of sterile fronds less than 1⁄2 the sterile frond length, those of fertile fronds ca. 2⁄3 the fertile frond length, sparsely scaly; sterile blades herbaceous to chartaceous, 15–80 ⫻ 7–50 cm, pinnate-pinnatifid, tapering to pinnatifid, apices acuminate, each apex often long-extended, with proliferous buds along its length; sterile pinnae 3–6 free pairs plus 1–4 partly adnate pairs, mostly 14–25 ⫻ 2–4(–5) cm, proximal pinnae stalked to 5 mm, unequally deltate, basiscopic side more developed, other pairs Ⳳ equilateral, shallowly lobed ca. 1⁄3 the way to the costae or less, lobes often toothed; veins areolate, areoles with mostly recurrent, often 1-forked, free veinlets; fertile blades oncepinnate, pinna pairs 5–10, mostly entire or with undulate mar-

BOLBITIS UMBROSA

gins except for proximal pair more deeply lobed and basiscopically exaggerated; sporangia covering the abaxial surfaces of fertile blades or sometimes along main veins only.

157

with an apical pinna similar to lateral pinnae, not flagellate or proliferous; sterile pinnae 10–20 ⫻ 2–3 cm, to ca. 12(–30) pairs, bases broadly cuneate, apices acuminate, margins crenate-serrate (sometimes doubly so), the serrations ca. 5 mm apart; veins areolate, the subcostal areoles with 0–1 excurrent, free veinlets; fertile blades once pinnate, with 8–12 pairs of short-stalked pinnae ca. 3 ⫻ 0.6–1 cm, pinnae narrowly elliptic, entire, rounded at tips; sporangia covering the abaxial surfaces of fertile blades.

Distribution. Terrestrial or epipetric, especially near streams, in wet lowland and montane forests; 50–1400 m. Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Ec. Selected Specimens Examined. Chis (Croat & Hannon 63344, UC). Gro (Vela´squez 357, ENCB, MEXU, US). Jal (Mexia 1233, UC). Mich (Hinton 16305, LL, UC, US). Nay (McVaugh 18983, IEB, MEXU, MICH). Oax (Mexia 2186a, UC). Pue (Herna´ndez X. & Sharp X-157, MEXU). Tab (Matuda 3379, MEXU, NY, US). Ver (Smith 2106, UC, US).

Distribution. Wet lowland forests, especially stream banks; 550 m. Mexico; Nic, CR, Pan; Col, Ven, Sur, Ec, Peru, Braz, Bol, Parag, nw Arg. Specimens Examined. Chis (Breedlove 32817, DS, MEXU; Ghiesbreght 409, GH, cited by Hennipman, 1977, but not verified; Mu¨nch 19, DS, US).

This is the most common Bolbitis in Mexico; the flagelliform apices are characteristic of most sterile fronds. Bolbitis portoricensis is most easily confused with B. aliena and B. umbrosa (two other species with pinnatifid blade apices), both of which differ in lacking free included veinlets in the areoles. Bolbitis hastata differs in its pinnatifid blades. The sole collection from “Puebla” is undoubtedly mislabeled (“slopes of Ixtaccihuatl above Huejotzingo, 13,700 ft”).

Bolbitis serratifolia differs from congeners in Mexico by the pinnate blades with narrowly elliptic, entire pinnae; from B. hemiotis and B. pergamentacea it differs by the lack of recurrent veinlets in the areoles. It usually has more pairs of lateral pinnae than any other species of Bolbitis in Mexico, excepting B. bernoullii, which differs in its hemiepiphytic habit and denticulate rhizome scales.

7. BOLBITIS SERRATIFOLIA

8. BOLBITIS UMBROSA

Bolbitis serratifolia (Mertens ex Kaulf.) Schott, Gen. Fil., t. 14. 1835. Figs. 57D, E.

Bolbitis umbrosa (Liebm.) Ching in Christensen, Index Filic., Suppl. 3: 50. 1934. Figs. 57F–H.

Acrostichum serratifolium Mertens ex Kaulf., Enum. Filic. 66. 1824. Leptochilus serratifolius (Mert. ex Kaulf.) C. Chr., Bot. Tidsskr. 26: 289, f. 3. 1904. Type. Brazil. Rio de Janeiro, s. coll. s.n., Herb. Mertens (not found by Hennipman, 1977; isotype C). Leptochilus mexicanus Christ, Bull. Herb. Boissier, se´r. 2, 7: 414. 1907. Bolbitis mexicana (Christ) C. Chr., Index Filic., Suppl. 3: 49. 1934. Type. Mexico. Chiapas: San Cristo´bal, Mu¨nch 19 (presumably P, not found; isotypes DS!, US!).

Acrostichum umbrosum Liebm., Mexic. Bregn. 174 (reprint 22) 1849. Leptochilus umbrosus (Liebm.) C. Chr., Index Filic. 388. 1906. Syntypes. Mexico. Veracruz: Consoquitla pr. Mirador, Oct 1841, Liebmann s.n. [Pl. Mex. 2690, Fl. Mex. 665 (folio)] (C!); Hac. de Mirador, Barranca de S. Francisco, Nov 1841, Liebmann s.n. [Pl. Mex. 2689, Fl. Mex. 663, 664 (folios)]. We here designate Fl. Mex. 663 (photo US!) as lectotype; isolectotypes B-frag., NY!, BM, K!, UC! Hennipman (1977: 283) chose as lectotype a specimen at BM, but the true lectotype should be selected from Liebmann’s herbarium at C, which we now do.

Rhizomes long-creeping, not climbing, to 15 mm diam.; rhizome scales dark brown, to 6 ⫻ 2 mm, entire; fronds subdistant, mostly 50–100 cm long; stipes 20–70 cm ⫻ 3–6 mm, those of sterile fronds less than 1⁄2 the sterile frond length, those of fertile fronds ca. 2⁄3 the fertile frond length, with appressed scales; sterile blades firmly chartaceous, mostly 25–50 ⫻ 20–25 cm, pinnate,

Rhizomes short-creeping, ca. 7 mm diam.; rhizome scales brownish, to 5 ⫻ 2 mm, entire; fronds subdistant, 50–115 cm long, the fertile shorter than the sterile; stipes 20–65 cm ⫻ 2–5 mm, those of the sterile fronds less than 1⁄2 the sterile frond length, those of fertile fronds 1⁄2–3⁄4 the fertile frond length; sterile

158

BOMMERIA

blades thinly chartaceous, 30–70 cm, pinnate-pinnatifid, apices pinnatifid, not flagellate or proliferous; sterile pinnae 12–21 ⫻ 3.7–7(–10) cm, 3–5 free pairs ⫹ 4–10 additional pairs connected by wings along rachises, proximal pinnae inequilateral with basiscopic segments longer and more deeply lobed than acroscopic segments, distal pinnae Ⳳ equilateral and strongly decurrent; sterile segments subentire to shallowly lobed or pinnatifid, with a small tooth at the base of each sinus between segments; veins areolate, each areole usually lacking included free veins; fertile blades pinnate-pinnatifid, 10–35 ⫻ 7–16 cm; sporangia Ⳳ covering the abaxial surfaces of fertile blades but with a sterile zone along the main veins.

There are five species of Bommeria, all in the New World (southwestern United States to Costa Rica), mostly in dry, rocky regions. All five occur in Mexico. Bommeria is related, but not very closely, to Hemionitis as shown in its netted veins, gymnogrammoid sori, and pedate fronds. References Gastony, G. J. & C. H. Haufler. 1976. Chromosome numbers and apomixis in the fern genus Bommeria (Gymnogrammaceae) Biotropica 8: 1–11; Haufler, C. H. 1979. A biosystematic revision of Bommeria. J. Arnold Arbor. 60: 445–476; Maxon, W. R. 1913. Studies of tropical American ferns No. 4. Contr. U.S. Natl. Herb. 17: 133–179; Ranker, T. A. & C. H. Haufler. 1990. A new combination in Bommeria (Adiantaceae). Amer. Fern J. 80(1): 1–3.

Key to the Mexican Species of Bommeria

Distribution. Terrestrial in wet montane forests; 450–1000 m. Mexico; Hond (Hawkins 1000, UC). Specimens Examined. Chis (Breedlove 21621, 22483, DS, NY; 30866, DS, MEXU). Ver (Purpus 4355, UC, US; Purpus 4354, 4371, 11142, 12082, 16723, US).

Bolbitis umbrosa is easily confused with B. aliena, which has similar venation, without included veinlets in the areoles. The former differs in having more numerous pinna pairs, to 5 pairs of completely free pinnae, more deeply pinnatifid (even bipinnatifid) proximal pinnae, and small teeth at the bases of sinuses.

20. BOMMERIA Bommeria E. Fourn. in Baillon, Dict. Bot. 1: 448. 1876 [1877]. Lectotype (chosen by Maxon, 1913: 169): Bommeria ehrenbergiana (Klotzsch) Underw. [⬅ Gymnogramma ehrenbergiana Klotzsch]. Terrestrial or epipetric; rhizomes short- to long-creeping, scaly; fronds small, erect; stipes brown, hairy to glabrous; blades usually pentagonal to deltate, pinnatifid to tripinnatifid at bases, herbaceous to coriaceous, abundantly hairy on both sides, sometimes also with scales abaxially; veins free or anastomosing; sporangia along veins in distal third to two-thirds of their lengths; indusia and paraphyses absent, margins plane or inrolled (in B. ehrenbergiana); spores tetrahedral-globose; x ⫽ 30.

1. Blades palmate, 3–5(–7)-lobed, subglabrous, hairs scattered, inconspicuous, 0.5 mm, the lobes 2–5.5 cm wide; veins netted; sori running along vein length except near midveins. .......................................................... 2. B. elegans. 1. Blades pinnatifid to pinnate, with abundant hairs 0.5–1 mm, the lobes or segments 0.3–1 cm wide; veins free or netted; sori along the distal 1⁄4–1⁄3 of veins. 2. Blades with hairs and scales along costae. 3. Blades pinnate-pinnatifid above basal pinna pair; rhizomes long-creeping; veins free ........................ 3. B. hispida. 3. Blades pinnatifid above basal pinna pair; rhizomes compact; veins netted. .................... 5. B. subpaleacea. 2. Blades with hairs and only rare linear scales 2–8 cells wide along costae. 4. Veins netted. ................................................... 1. B. ehrenbergiana. 4. Veins free. .................................................................. 4. B. pedata.

1. BOMMERIA EHRENBERGIANA Bommeria ehrenbergiana (Klotzsch) Underw. in Baillon, Dict. Bot. 1: 448. 1876 [1877]. Figs. 59B, C. Gymnogramma ehrenbergiana Klotzsch, Linnaea. 20: 411. 1847. Stenogramma ehrenbergiana (Klotzsch) E. Fourn., Mexic. Pl. 1: 71. 1872. Type. “Ad thermas mexicanas, prope Grande in regno Mexico,” Ehrenberg 662 (B!; isotype UC!). Gymnogramma podophylla Hook., Sp. Fil. 5: 152. 1864. Hemionitis podophylla (Hook.) J. Sm., Hist. Fil. 150. 1875. Bommeria podophylla (Hook.) E. Fourn., Bull. Soc. Bot. France 27: 328. 1880. Type. Mexico. Veracruz: Between Vera Cruz and Orizaba, Mu¨ller 719 (K!; isotype BM!, frag. NY!).

Rhizomes short-creeping, horizontal; rhizome scales linearlanceolate, orange-tan, ca. 5 mm; fronds 12–40 cm, approximate; stipes 1⁄2–2⁄3 the frond length, terete at bases to grooved distally, castaneous to atropurpureous, glabrous except for acicular hairs in grooves; blades pedate, deltate, bipinnatifid, 5–16 cm; costae castaneous to atropurpureous; adaxial surfaces with abundant unicellular, acicular hairs 0.3–0.5 mm; abaxial surfaces with rare scales 2–8 cells wide and acicular hairs 0.3–1 mm; veins netted; sori along the distal 1⁄4–1⁄2 of veins; spores 64 per sporangium.

BOMMERIA HISPIDA

159

Distribution. Terrestrial and epipetric in periodically moist deciduous woods; 1550–1920 m. Mexico.

Distribution. Terrestrial, on moist banks in mesic woods; 750– 2000 m. Mexico.

Selected Specimens Examined. Gro (Lorea 2658, IEB, FCME). Hgo (Gastony & Haufler 1086, GH, MO, UC, US). Jal (Cochrane & Santana M. 13025, WIS). Me´x (Matuda 28799, US). NL (Mueller & Mueller 662, F, GH, MICH). Oax (Calzada 21303, MEXU). Pue (Arse`ne 2147, MICH, UC). Qro (Rzedowski 53516, IEB). SLP (Poust 98, NY). Tam (Leyva B. s.n., 19 Oct 1985, UNL). Ver (Mu¨ller 719, NY).

Selected Specimens Examined. Jal (Pringle 2581, GH, NY, UC; Rose & Painter 6928, US). Me´x (Hinton 4740, US). Mor (Pringle 7946, DS, MICH, MO, NY, TEX, 11782, CAS, ENCB, MO, TEX, UC, US; Rose & Rose 11083, NY, US). Nay (McVaugh 16340, CAS, MEXU, MICH, NY, US). Oax (Mickel 6250, ENCB, NY, UC; Ranker & Yatskievych 820, UC). Zac (Correa 50, MICH).

This species resembles B. subpaleacea in having netted veins but is distinct in having only linear abaxial costal scales (vs. conspicuous, abundant, lanceolate to ovate scales) and sori along distal 1⁄4–1⁄2 of veins (vs. 1⁄2–3⁄4 of veins).

Bommeria elegans is distinct by its large size, deeply lobed, subglabrous blades, and lack of laminar buds. It has traditionally been placed in Hemionitis, but Ranker and Haufler (1990) transferred this species to Bommeria based on details of spores, rhizome scales, and gametophyte symmetry. Molecular evidence (Gastony & Rollo, 1995) confirms this transfer.

2. BOMMERIA ELEGANS Bommeria elegans (Davenp.) Ranker & Haufler, Amer. Fern J. 80: 2. 1990. Fig. 59A. Hemionitis elegans Davenp., Gard. & Forest 190: 484. 1891. Type. Mexico. Jalisco: shaded banks and ledges of the barranca near Guadalajara, 24 Sep 1889, Pringle 2585 (GH!; isotypes NY!, UC!, US!–2 sheets). Hemionitis hederifolia J. Sm. in Seemann, Bot. Voy. Herald 339. 1854. Type. Mexico. Leibold s.n. (J. Smith Herb. 1843) (BM!; isotype US! photo ex BM).

Rhizomes horizontal, compact, 3–5 mm diam.; rhizome scales bicolorous, central portion dark brown, margins light brown, with marginal cilia 0.3 mm long, scales 3 ⫻ 0.8–1 mm, linearlanceolate, becoming ovate, 5 ⫻ 2 mm, concolorous at bases of stipes; fronds 26–45 ⫻ 9–22 cm; stipes 3⁄5–3⁄4 the frond length, 1.5–2.7 mm diam., dark reddish brown with scales and hairs 1 mm long at bases, glabrous otherwise, terete; blades deeply palmately lobed into 3–5(–7) lobes, the lobes entire, narrowly deltate or lanceolate, acuminate, margins setose, not recurved; abaxial surfaces with stiff, one-celled, acicular hairs 0.5 mm, most abundant among sporangia and near the blade margins, scattered, long, soft hairs 1.5–2 mm grading into narrow scales 3 mm long at frond bases; adaxial surfaces nearly glabrous, with some hairs 0.5 mm long on midveins and near the blade margins; veins netted, major veins dark near their bases; sori along all veins except near lobe midveins; spores tan; 2n⫽60 (Oax).

3. BOMMERIA HISPIDA Bommeria hispida (Mett. ex Kuhn) Underw., Bull. Torrey Bot. Club 29: 633. 1902. Figs. 59D–F. Gymnogramma hispida Mett. ex Kuhn, Linnaea 36: 72. 1869. Gymnopteris hispida (Mett.) Underw., Native Ferns, ed. 6, 84. 1900. Type. U.S.A. Texas: Pass of the Limpia, Wright 819 (B; isotypes GH!, MO!, NY!, US!, YU!). Bommeria schaffneri E. Fourn., Bull. Soc. Bot. France 27: 327. 1880. Type. Mexico. In montibus Escabrillos prope San Luis Potosı´, 1877, Schaffner 6 (P; GH photo, US photo!). Gymnogramma ehrenbergiana Klotzsch var. muralis Pringle ex Davenp., Bot. Gaz. 19: 392. 1894. Type. Mexico. Jalisco: on faces of cliffs near Tequila, Pringle 4420 (GH, not found; isotypes BM!, BR!, F, GH!, K!, MEXU!, MSC, TEX!, UC!, US!, VT).

Rhizomes long-creeping; rhizome scales linear-lanceolate, orange-tan, 1.5–2 mm; fronds (2–)7–10(–40) mm apart; stipes 4– 20 cm long, 3⁄4–4⁄5 the frond length, light grayish brown, terete to flattened adaxially, scaly and hairy at bases, mostly sparsely hairy, densely so near blade bases; blades 2–6 ⫻ 2.5–8 cm, pentagonal, pedately divided, bipinnatifid, the basal pinna pair deeply pinnatifid with greatly exaggerated basiscopic pinnatifid pinnules, the distal pinnae pinnatifid, rachises dilated laterally between the pinnules, or some forming entire rounded foliaceous lobes between pinnae; blade surfaces densely hispid adaxially, with stiff, unicellular, acicular hairs 1 mm long, the abaxial

160

BOMMERIA

surfaces bearing similar (but mostly curled) hairs in addition to tan lanceolate scales 0.2–0.5 mm wide on the rachises and midveins; sori along the distal half of veins, the pinnule margins often slightly incurved; veins free, obscure; spores tan, 64 per sporangium; 2n⫽60 (Jal, USA).

Distribution. Terrestrial on shady rock bases, oak forests; pineoak forests, matorral, eroded volcanic rocks, base of sandstone cliffs; 1500–2300(–3000) m. Sw USA (Ariz, N Mex, Tex); Mexico. Selected Specimens Examined. Ags (McVaugh 18309, MICH, NY). BCS (Carter et al. 2035, DS, K, UC, US). Chih (Correll & Johnston 21596, TEX, US). Coah (Wynd & Mueller 575, K, NY, US). DF (Schaffner 32, NY, US). Dgo (Pringle 10151, NY, UC, US). Hgo (Hitchcock & Stanford 7250, US). Jal (Pringle 4420, BM, BR, K, MEXU, UC, US). Me´x (Hinton 1431, GH). Mich (King & Soderstrom 4699, NY). SLP (Parry & Palmer 1006, F, GH, MO, NY p.p., US). Son (Wiggins 7403, DS, US). Zac (McVaugh 17756, IEB, NY).

This species is distinct by its small size, long acicular hairs, and free veins.

4. BOMMERIA PEDATA Bommeria pedata (Sw.) E. Fourn., Bull. Soc. Bot. France 27: 327. 1880. Figs. 60A–D. Hemionitis pedata Sw., Syn. Fil. 20, 209, t. 1, f. 3. 1806. Gymnogramma pedata (Sw.) Kaulf., Enum. Filic. 69. 1824. Gymnopteris pedata (Sw.) C. Chr., Index Filic. 341. 1905. Type. s. coll. s.n.; according to Haufler (1979) perhaps collected by Ne´e in Mexico and sent by Cavanilles to Swartz (presumably at S).

Rhizomes short-creeping; rhizome scales linear-lanceolate to ovate-lanceolate, concolorous to bicolorous with age, light brown with dark center and narrow light brown margins, 3–4 ⫻ 0.8–1 mm with cilia 0.3 mm long; fronds 1 mm apart; stipes 1 ⁄2–2⁄3 the frond length, castaneous to atropurpureous, terete to sulcate at apices, glabrous except for scattered scales and hairs 0.5–1 mm long at bases; blades pentagonal, pedately divided, basal pinna pairs pinnatifid, each with a greatly exaggerated basiscopic pinnatifid pinnule, remainder of blades pinnatifid or with one pair of suprabasal pinnatifid pinnae; lobes entire,

rounded, the margins slightly recurved; abaxial surfaces densely hispid, with stiff, unicellular, acicular hairs 0.3–0.8 mm long and scales 0.1–0.5 mm wide (2–8 cells wide at bases); adaxial surfaces with sparse, unicellular, acicular hairs 0.2–0.5 mm long; veins free, endings enlarged; sori along the distal 2⁄3–3⁄4 of veins; spores tan, brown or gray, with reticulate markings; n⫽2n⫽90, apogamous (Mor).

Distribution. Terrestrial and epipetric on rocky banks, on steep slopes in open mesic and xeric woods, pine-oak forests, matorral, igneous slopes; (50–)900–1800(–2550) m. Mexico; Guat, Hond, Salv, Nic, CR. Selected Specimens Examined. BCS (Breedlove 45619, CAS). Chih (Knobloch 916, US). Chis (Breedlove 51173, CAS, ENCB, NY). Col (Jones 540, US). DF (Orcutt 4331, US). Dgo (Garcı´a 796, US). Gro (Hinton 9209, NY, US). Gto (Galva´n & Galva´n 2370, 2408A, ENCB, IEB). Jal (Pringle 1861, BM, NY, US). Me´x (Rzedowski 29601, CAS, ENCB). Mich (King & Soderstrom 4677, NY, UC). Mor (Orcutt 4390, DS, US). Nay (McVaugh 16424, MEXU, MICH, NY, US). Oax (Mickel 6244, NY, UC). Pue (Nicola´s s.n., 27 Oct 1909, NY p.p.). Sin (Keil & Canne 8823, NY, US). SLP (Schaffner s.n., NY). Son (Pennell 19673, US). Ver (Seaton 41, F, NY, US). Zac (McVaugh 17652, MICH). Unverified, Doubtful, or Mistaken Reports. Ags (Garcı´a 3278, HUAA, and three other collections, cited by Siqueiros-Delgado & Gonza´lez-Adame, 2004, but not verified).

Bommeria hispida is similar in its free veins but has longcreeping rhizomes (fronds 7–10 mm distant), broad abaxial blade scales, and dimorphic hairs abaxially (acicular and curled).

5. BOMMERIA SUBPALEACEA Bommeria subpaleacea Maxon, Contr. U.S. Natl. Herb. 17: 169. 1913. Type. Mexico. Puebla: Purpus 4025 (US!; isotypes BM!, DS!, F, GH!, MSC, NY!). Figs. 60E–G. Bommeria knoblochii Maxon, Amer. Fern J. 32: 59 1942. Type. Mexico. Chihuahua: Mojarachic, Knobloch 6044 (US!; isotypes BM, DS, GH!, MSC).

Rhizomes short-creeping, compact; rhizome scales linearlanceolate to ovate-lanceolate, 1.5–2 ⫻ 0.5–0.8 mm, with shortfimbriate margins, slightly to strongly bicolorous, centers dark brown, borders wide, of clear cells with brown walls; fronds 1 mm apart; stipes castaneous to atropurpureous, terete to sulcate

BOTRYCHIUM

distally, 2⁄3–3⁄4 the frond length, with acicular hairs 0.3 mm long, and scattered ovate, tan scales 2 ⫻ 1–1.5 mm at bases; blades pentagonal, 2–11 ⫻ 2–4.5 cm wide, pedately divided, the basal pinna pair pinnatifid and greatly exaggerated basiscopically, remainder of blades pinnatifid; segments entire; abaxial surfaces with rare, small, tan, linear-lanceolate to ovate scales 1–1.5 ⫻ 1.5 mm; 10 or more cells wide at bases, also with dense acicular hairs 0.4–1 mm long; adaxial surfaces densely hispid with acicular hairs 0.2–0.5 mm long; sori covering distal 2⁄3–3⁄4 of veins; veins netted, but conspicuous; spores tan to brown, cristate.

Distribution. Terrestrial and epipetric in xeric, lightly shaded regions, rare throughout range; 1000–2000 m. Mexico. Selected Specimens Examined. Chih (Knobloch 6044, US, 8000, MSC, US). Gto (Dı´az Barriga 5847, MEXU, UAMIZ; Kenoyer 2381, US). Hgo (Sa´nchez-Mejorada 195, US). DF (Schaffner 7, NY). Mor (Matuda 32509, MEXU). Oax (Mickel 4556, NY). Pue (Purpus 4025, BM, DS, F, GH, MSC, NY, UC, US). SLP (Parry & Palmer 1006, NY p.p., YU). Unverified, Doubtful, or Mistaken Reports. Sin (reported by Vega A. et al., 1995, but no specimen cited; from range, more likely to be B. hispida).

Bommeria subpaleacea is distinct by its linear-lanceolate to ovate blade scales, netted veins, and the sporangia on more than 1⁄2 the vein length from costae to margins. Specimens from Oaxaca and Puebla (including the type) have ovate rather than linear-lanceolate blade scales characteristic of other specimens.

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21. BOTRYCHIUM Botrychium Sw., J. Bot. (Schrader) 1800(2): 8, 110. 1801. Lectotype (chosen by J. Smith, Hist. Fil. 369. 1875): Botrychium lunaria (L.) Sw. [⬅ Osmunda lunaria L.]. Sceptridium Lyon, Bot. Gaz. 40: 457. 1905. Lectotype (chosen by Clausen, 1938: 24): Sceptridium obliquum (Muhl. ex Willd.) Lyon [⬅ Botrychium obliquum Muhl. ex Willd.] ⬅ Botrychium dissectum Spreng. f. obliquum (Muhl. ex Willd.) Fernald. Botrychium sect. Osmundopteris Milde, Verh. K. K. Zool.-Bot. Ges.-Wien 19: 96. 1896. Botrychium subg. Osmundopteris (Milde) R. T. Clausen, Mem. Torrey Bot. Club 19(2): 93. 1938. Osmundopteris (Milde) Small, Ferns S.E. States 377, 482. 1938. Type: Botrychium virginianum (L.) Sw. [⬅ Osmunda virginiana L.]. Botrypus Michx., Fl. Bor.-Amer. 2: 274. 1803. Type: Botrypus virginianus (L.) Mich. [⬅ Osmunda virginiana L.] ⬅ Botrychium virginianum (L.) Sw.

Terrestrial; stem subterranean, erect, fleshy; roots thick, fleshy; fronds mostly one (rarely two), with two distinct parts: a spreading pinnately to ternately divided sterile blade (sometimes called trophophore), with free veins, and an erect fertile portion (sometimes called sporophore), 2–3 times divided, each sporophore stalked and terminated by a panicle-like sporangial cluster, usually exceeding the sterile blade in length, arising from base of the sterile blade or lower on stipe, sometimes at or below ground level; sporangia large, globose, opening by a horizontal slit; spores tetrahedral, one thousand to several thousand per sporangium; gametophytes subterranean, whitish or brownish, mycorrhizal, non-green; x⫽45. Botrychium comprises about 60 species, largely of temperate regions, usually growing in pastures, grassy meadows, and open second-growth woodlands, often in areas of disturbance. There is considerable variation in frond size and dissection, and this variation is further confounded by the tendency to form “genus communities” with two or three species present in a population. Twenty new species have been recently described in North America, showing that much work is needed for a full understanding of the genus. Highland pastures and grassy road shoulders should be searched for more populations of these unusual plants. For studies of variation, all fronds should be taken, leaving the underground stems unharmed to produce new fronds the following season. References Clausen, R. T. 1938. A monograph of the Ophioglossaceae. Mem. Torrey Bot. Club 19(2): 1–177; Hauk, W. D. 1995 [1996]. A molecular assessment of relationships among cryptic species of Botrychium subgenus Botrychium (Ophioglossaceae). Amer. Fern J. 85: 375–394.

Key to the Mexican Species of Botrychium 1. Vegetative blades narrowly oblong, less than 3 ⫻ 1 cm; buds glabrous; fertile stalks always developed (subg. Botrychium). 2. Distal margins of pinnae entire to very shallowly crenulate, pinna bases to 90⬚ angle, anterior margins straight or slightly convex; vegetative blades lacking supernumery sporangia (NL). .................................................................................................... 2. B. minganense. 2. Distal margin of pinna coarsely jagged, pinna bases at 50–60⬚ angle, anterior margins concave; vegetative blades commonly bearing supernumery sporangia (Dgo, Me´x). ........................................................................................................................................ 5. B. tolucaense. 1. Vegetative blades deltate, more than 5 cm wide; buds hairy; fertile stalks commonly absent. 3. Fertile portion attached at base of sterile blades; leaf sheaths open; texture thin (subg. Osmundopteris). .............................. 6. B. virginianum.

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BOTRYCHIUM 3. Fertile portion attached at some distance below the blades, generally at or below ground level; leaf sheaths closed; texture fleshy, thick-herbaceous (subg. Sceptridium). 4. Terminal segments mostly elongate and pointed, the apices cut only shallowly or not at all, pinnules show strong reductions from base to tip and segments grade into one another distally. ......................................................................... 1. B. decompositum. 4. Terminal segments ovate, acute to rounded at apices, mostly nearly equal in size; blades cut symmetrically to pinna apices. 5. Plants robust, mostly 4-pinnate, the segments mostly overlapping; Socorro Island. .......................................................... 4. B. socorrense. 5. Plants not robust, mostly to 3(–4)-pinnate, the segments mostly approximate to remote; mainland Mexico. .............. 3. B. schaffneri.

1. BOTRYCHIUM DECOMPOSITUM Botrychium decompositum M. Martens & Galeotti, Me´m. Foug. Mexique 51, pl. 1. 1842. Figs. 61F–H, 62A, 63A–D. Sceptridium decompositum (M. Martens & Galeotti) Lyon, Bot. Gaz. 40: 457. 1905. Botrychium dissectum Spreng. subsp. decompositum (M. Martens & Galeotti) R. T. Clausen, Bull. Torrey Bot. Club 19(2): 58. 1938. Type. Mexico. Veracruz: Orizaba, Galeotti 6452 (BR!, photos GH!, US!).

Buds hairy; leaf sheaths closed; fronds 15–50 cm long; sterile blades 5–15 ⫻ 7–22 cm, deltate to pentagonal, ternately compound; pinnae other than the basal pair pinnate to bipinnatepinnatifid, tips elongate; segments acute, ovate, often divided to main vein, terminal segments ovate to lanceolate, usually pointed, apices cut only shallowly or not at all, irregularly serrulate, usually considerably larger than other segments; leaf tissue fleshy or thick-herbaceous, glabrous or with few scattered hairs; sporophores arising from the common stalk at or just below the ground surface and far exceeding the sterile blades in height, commonly absent.

enable us to describe them with confidence at this time. There are three major variants that seem to run together. Botrychium decompositum in the strict sense has very coarse segmentation with blunt-tipped pinnules and pinnae. The degree of margin toothing/denticulation is quite variable (Figs. 63A, B; e.g., Pringle 15706, F, GH, UC, from Hidalgo). A second form has pinnules and segments strongly ascending and narrowly lanceolate, remote, pointed. Juvenile plants have cutting like that of the adults. It grows in sunny, grassy roadsides at the edge of forests. Where populations are more shaded, the plants become less vigorous and less frequent (Fig. 63D; e.g., Mickel 9627, 9687, NY, from Oaxaca). The third variant has pinnules and segments only moderately ascending, lanceolate, approximate, and sharply pointed (shortacuminate). Juvenile plants have long, undivided pinna tips (Fig. 61F; e.g., Wagner 80805, US, from Hidalgo). It is found in moist pine-oak forests, especially in disturbed areas such as along overgrown lumber roads (Fig. 61G; e.g., Mickel 9681, NY, from Oaxaca). The type specimen of B. decompositum, however, has somewhat pointed segment apices, and the variants do not seem to have unique morphologies. More collections and additional study are needed to discern species limits in this group.

2. BOTRYCHIUM MINGANENSE Botrychium minganense Vict., Proc. & Trans. Roy. Soc. Canada, ser. 3, 21: 331. 1927. Type. Canada. Que´bec: Arch’pel de Mingan, ˆIle Niapisca, Marie-Victorin & Rolland-Germain 18122 (isotype US). Figs. 61C–E.

Distribution. Grassy areas or open forests; 900–2800 m. Mexico; Guat, Hond, CR, Pan. Selected Specimens Examined. Chis (Breedlove 31940, DS, F, NY). Hgo (Pringle 8922, UC). Me´x (Hinton 1850, MEXU). Mich (Dı´az Barriga 2265, IEB). NL (Rodrı´guez L. 48, ASU). Oax (Mickel 7253, NY). Pue (Sharp 441189, US). Qro (Guzman 214, UAMIZ). Tam (Reznicek 8423, CHAPA, MICH). Ver (Taylor & Nee 296, NY).

The name “B. decompositum” has been used for a complex related to B. dissectum Spreng. of United States, Canada, and Greater Antilles. There are several variations that might, with further study, prove to be distinct species. Unfortunately, the distinguishing characters are few and their variation too great to

Buds glabrous; fronds ca. 6 cm long, glabrous; sterile blades 1.5 ⫻ 0.8 cm, pinnate; pinnae flabellate, distal margin entire or shallowly crenulate, base to 90⬚ angle, anterior margin straight to slightly convex; leaf tissue herbaceous; sporophores 2 cm long, arising from the common stalk just below the base of the sterile blades and exceeding the sterile blades in height; 2n⫽180 (USA).

BOTRYCHIUM SOCORRENSE

Distribution. Grassy meadows in pine forests; 3700 m. Mostly Canada and Alaska, northernmost United States and western mountains to Arizona; Mexico. Specimen Examined. NL (Schneider 966, F)

This is a small plant, differing from B. tolucaense in its smaller size and the lack of sporangia on the vegetative blades.

3. BOTRYCHIUM SCHAFFNERI Botrychium schaffneri Underw., Bull. Torrey Bot. Club 30: 51. 1903. Figs. 64A–D. Sceptridium schaffneri (Underw.) Lyon, Bot. Gaz. 40: 458. 1905. Type. Mexico. San Luis Potosı´: In mountains near San Luis Potosı´, Oct 1879 [& 1881], Schaffner s.n. (NY!; isotype K!). Botrychium pusillum Underw., Bull. Torrey Bot. Club 30: 50. 1903. Sceptridium pusillum (Underw.) Lyon, Bot. Gaz. 40: 458. 1905. Botrychium schaffneri Underw. var. pusillum (Underw.) R. T. Clausen, Mem. Torrey Bot. Club 19(2): 42. 1938. Type. Mexico. Me´xico: Sierra de las Cruces, Pringle 5192 (NY!; isotype GH!).

Buds hairy; fronds (4–)10–40 cm long, sterile blades 8–12 ⫻ 12–16 cm, deltate, ternately compound to tripinnate-pinnatifid, mostly 3(–4)-pinnate; segments ovate, oblong or roundish, small, to 6 ⫻ 5 mm, margins serrulate to entire, all segments roughly the same size, terminal segment of each pinna not distinctly larger; leaf tissue fleshy, thick-herbaceous, glabrous or with a few hairs; sporophores inserted at or just below the ground surface, exceeding the sterile blades in height, commonly absent; 2n⫽ca. 90 (Me´x).

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Distribution. In pine-oak woods. 1000–3050 m. Mexico; Guat, Hond, Salv, CR; Col, Ven, Ec, Peru, Bol, Arg. Selected Specimens Examined. Ags (Siqueiros D. 4315, IEB). Chih (Jones s.n., 26 Sep 1903, DS, F, MO). Chis (Breedlove 12920, DS, MICH). Col (Guizar Nolazco 1182, MEXU). DF (Pringle s.n., 19 Sep 1903, NMC). Dgo (McVaugh 21730, MICH). Gro (Rzedowski 26025, MICH). Hgo (Pringle 8922, CAS, GH, MICH, PH, UC). Jal (McVaugh 19629, MICH, NY). Me´x (Hinton et al. 8301, GH, MO, NY, US). Mich (Garcı´a Lo´pez & Dı´az B. 3895, IEB). Mor (Mendoza 55, UAMIZ). Nay (Breedlove 44529, DS). NL (Conteras Ocanas s.n., Nov 1985, UNL). Oax (Mickel 3944, NY). Pue (Purpus 5687, UC). Qro (Dı´az Barriga 6908, IEB). SLP (Schaffner 25, NY). Son (Van Devender 95-390, ARIZ). Ver (Mu¨ller s.n., in 1855, NY).

Botrychium schaffneri is distinguishable from B. decompositum in having smaller, ovate to oblanceolate segments of relatively uniform size and shape. There is considerable variation in plant size and compactness.

4. BOTRYCHIUM SOCORRENSE Botrychium socorrense W. H. Wagner in G. A. Levin & Moran, Mem. San Diego Soc. Nat. Hist. 16: 20, f. 9B. 1989. Type. Mexico. Socorro Island: slope ca. 1 km NW of summit of Cerro Evermann, Levin 2044 (SD!; isotypes BM!, MEXU!, MICH!, MO!, UC!, US!). Figs. 64E, F. Buds hairy; leaf sheaths closed; fronds 30–60 cm long; sterile blades 13–20 ⫻ 17–20 cm, deltate to pentagonal, ternately compound, 4-pinnate, cut symmetrically to pinna apices; segments acute to rounded at apices, mostly overlapping, mostly nearly equal in size, ovate, often divided to main vein, irregularly serrulate; leaf tissue fleshy, thick-herbaceous, glabrous or with few scattered hairs; sporophores arising from the common stalk at or just below the ground surface and far exceeding the sterile blades in height, commonly absent.

164

BOTRYCHIUM Specimens Examined. Dgo (Gordon s.n., 23 August 1963, GH). Me´x (only the type collection).

Distribution. Oak or pine-oak forests, in sparse grass and under brush, north and west slopes of Cerro Evermann; 900– 1000 m. Mexico. Specimens Examined. Rev (Guizar Nolazco 1182, MEXU; Levin 1814, MICH, SD, 2115, SD; Moran 5854, 5854A, SD).

Botrychium socorrense closely resembles specimens of B. schaffneri, the former differing in its extremely robust habit (40–60 cm tall vs. mostly 15–35 cm) and strongly overlapping blade divisions. It is also close to B. multifidum (S. G. Gmel.) Rupr. of California (sometimes called B. silaifolium C. Presl), resembling it in the large, floppy frond divisions.

5. BOTRYCHIUM TOLUCAENSE Botrychium tolucaense W. H. Wagner & Mickel, sp. nov. Type. Mexico. Me´xico: volcano of Toluca, in a wet canyon, 11,500 ft, Pringle 5193 (holotype GH!, isotype VT). Figs. 61A, B. A B. minganensi grandioribus frondibus, sporangiis in laminis vegetativis ferentibus abstans.

Buds glabrous; fronds ca. 12 cm long; sterile blades 2.4 ⫻ 1 cm, narrowly oblong, pinnate; pinnae flabellate, entire or lobed, bearing sporangia on their margins; leaf tissue herbaceous; fertile stalks 1.8 cm long, each arising from the common stalk shortly below the vegetative blade and exceeding the sterile blade in height.

Distribution. Grassy areas or open forests; 3500 m. Mexico.

It is distinct from B. minganense by the supernumery sporangia, and the larger plant size. Confusion about the identity of this plant is attested to by its previous identifications, namely Botrychium lunaria var. minganense and B. matricariifolium. The three excellent plants at VT are clearly distinct from any previously described species of Botrychium. It is no surprise that the elevation of collection was 11,500 ft. In 1991, Wagner visited the volcano in the hope of rediscovering this species. Although there were a number of seemingly appropriate habitats, he failed to find it. However, it is entirely possible that the season was wrong, or that droughts had kept the leaves from appearing. This species should be sought again at Volca´n Toluca, as well as in similar sites elsewhere. Botrychium tolucaense resembles B. ⫻ watertonense W. H. Wagner of western Alberta, Canada, in the presence of sporangia along the pinna margins on the otherwise sterile blade.

6. BOTRYCHIUM VIRGINIANUM Botrychium virginianum (L.) Sw., J. Bot. (Schrader) 1800(2): 111. 1801.Fig. 62B. Osmunda virginiana L., Sp. Pl. 2: 1064. 1753. Type. “America.” Kalm s.n. (LINN 1244.3). Osmunda cicutaria Savigny in Lamark, Encycl. 4: 650. 1797 [1798]. Botrychium cicutarium (Savigny) Sw., Syn. Fil. 171. 1806. Type. Plumier, Traite´ Foug. Ame´r. 136, pl. 159, 1705, based on a plant from Hispaniola collected by Plumier. Botrychium virginianum (L.) Sw. var. mexicanum Hook. & Grev., Bot. Misc. 3: 223. 1833 (as B. virginicum β mexicanum). Syntypes. Mexico. Veracruz: Jalapa, Chamisso s.n., “Rigla,” Capt. Veitch s.n. (K?). Botrychium brachystachys Kunze, Linnaea 18: 305. 1844. Type. Mexico. Roemer 35 (LZ, destroyed). Botrychium virginianum (L.) Sw. var. meridionale Butters, Rhodora 19: 213. 1917. Botrychium β virginianum subsp. meridionale (Butters) R. T. Clausen, Mem. Torrey Bot. Club 19(2): 103. 1938. Type. Mexico. Chiapas: Ghiesbreght 252 (GH!; isotype YU!).

Buds hairy; leaf sheaths open; fronds 15–75 cm long, sparsely hairy to glabrous; sterile blades 7–21 ⫻ 10–24 cm, deltate, ternately compound, to tripinnate-pinnatifid, pinnae above the basal pair mostly bipinnate-pinnatifid; ultimate segments 4–12 mm long, oblong to ovate, dentate or acutely incised at apex; leaf tissue thin, herbaceous, glabrous or with a few hairs; sporophores 5–15 cm long, attached at the base of sterile blades, equaling to exceeding the length of sterile blades, commonly absent; 2n⫽90 (Michigan).

CAMPYLONEURUM

Distribution. Terrestrial in commonly disturbed, moist, shady forests; 1380–2900 m. Can, USA; Mexico; Guat, Hond, Salv, CR, Nic, Pan; Gr Ant; Col, Ven, Ec, Peru, Braz, Bol; Europe, Asia. Selected Specimens Examined. Chih (LeSueur 1131, MO, TEX). Chis (Breedlove 34639, NY). Dgo (Ownbey & Ownbey 2000, US). DF (Reicke s.n., Oct 1913, MEXU). Gro (Anderson & Anderson 4932, NY). Gto (Dı´az Barriga 4003, ENCB). Hgo (Mickel 644, NY). Jal (McVaugh 14311, MICH, US). Me´x (Rzedowski 30909, MEXU). Mich (McVaugh 17830, NY). Mor (Lyonnet 896, GH, NY, US). Nay (McVaugh 18869, MICH). NL (Hinton et al. 22306, TEX). Oax (Mickel 3799, NY). Pue (Arse`ne 3524, MEXU). Qro (Barrie 1098, MEXU, TEX). Sin (Breedlove 44953, CAS). SLP (s. coll. 145, MEXU). Son (Fishbein 722, MEXU). Tam (Hinton et al. 24847, TEX). Tlax (Reeves 5929, ASC). Ver (Pringle 7753, GH, MO).

Botrychium cicutarium was reported by Clausen (1938) to have its fertile part equal to or shorter than the sterile and persistent leaves, but this character appears unreliable. Some of the Mexican material seems to have coarser cutting and more conspicuously pilose midribs than material of this species from the United States, but delimitation of B. cicutarium as a distinct taxon does not seem possible. A strange specimen from Hidalgo (Sa´nchez M. 130, US) has very large fronds (blades 26 ⫻ 36 cm, common stalks to 17 mm wide (pressed flat), roots 2–5 mm diam., segments to 13 ⫻ 9 cm, dissection bipinnate-pinnatifid to tripinnate. Most likely this represents the most extreme of the wide variation seen in B. virginianum.

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Epiphytic, occasionally epipetric or terrestrial; rhizomes shortto long-creeping; rhizome scales concolorous, brownish, often clathrate, surface glabrous, margins entire; fronds clumped to distant, monomorphic, sessile to long-stipitate, narrowly linear to elliptic, attenuate at apices and bases, articulate on phyllopodia; margins entire, modified; blades thin to usually thick, simple (in ours, rarely pinnate in a few extraterritorial species), glabrous (or sparingly hairy in a few extraterritorial spp.); venation areolate with 1 to usually several or many rows of areoles formed by anastomosing cross-veins from main lateral veins, the crossveins producing 2 (or sometimes more) excurrent, simple, free veins, a third excurrent vein sometimes completely bisecting the areole; sori round, exindusiate, non-paraphysate, at ends of free included veins, usually in 2 (to ca. 4) rows between main lateral veins; sporangia glabrous; spores bilateral, verrucate with spherical deposits; x⫽37. Campyloneurum is now usually segregated from Polypodium s.l. and includes approximately 50 neotropical species. It is characterized by the monomorphic, simple leaves (two species in South America are 1-pinnate), non-paraphysate sori borne on the free included vein tips, and usually clathrate, glabrous rhizome scales. There have been recent attempts at revision (Lellinger, 1988; Leo´n, 1992), but the genus is still inadequately understood. We depart in some respects from the circumscriptions adopted in both of those treatments, in particular with the species C. angustifolium and C. serpentinum. Several species, including C. phyllitidis, C. repens, and C. xalapense in Mexico, have been reported as having nectaries on the abaxial side of the blades (Mickel & Beitel, 1988; Leo´n, 1992). These occur at the juncture of the midribs and the main lateral veins, and apparently exude sugars. Whether there is regular ant association in Campyloneurum, as reported in some polypodiaceous ferns (Koptur et al., 1982), is unknown. Campyloneurum, including the two 1-pinnate species mentioned above (these excluded by Lellinger, 1988), is monophyletic and clearly allied to a large clade of other New World Polypodiaceae (Haufler, unpubl. data; Schneider et al., 2003), most especially to Niphidium and then to Microgramma. Nearly all species in this clade are simple-bladed. Species of Campyloneurum occupy an unusually wide variety of habitats, from sea level to above 4000 meters in paramo vegetation; species can be terrestrial, epipetric, or epiphytic. References

2 2 . C A M PYL O N E UR U M Campyloneurum C. Presl, Suppl. Tent. Pterid. 189. 1836. Lectotype (chosen by J. Sm., Hist. Fil. 95. 1875): Campyloneurum repens (Aubl.) C. Presl [⬅ Polypodium repens Aubl.].

Lellinger, D. B. 1988. Some new species of Campyloneurum and a provisional key to the genus. Amer. Fern J. 78: 14–35; Leo´n, B. 1992. A taxonomic revision of the fern genus Campyloneurum (Polypodiaceae). Unpubl. Ph.D. thesis, Aarhus University, Denmark; Koptur, S., A. R. Smith & I. Baker. 1982. Nectaries in some neotropical species of Polypodium (Polypodiaceae): preliminary observations and analyses. Biotropica 14: 108–113.

Key to the Mexican Species of Campyloneurum 1. Main lateral veins obscure or barely visible on the adaxial blade surfaces, if barely visible then main lateral veins and cross-veins similar in coloration; fronds 0.4–6 cm wide; sori 1–6(–8)-seriate between midribs and blade margins. 2. Main lateral veins barely visible; rhizomes not or only slightly pruinose; rhizome scales 2–3 ⫻ 0.8–1 mm, with toothed margins; sori 4–6(–8)-seriate between midribs and blade margins.

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3. Blades oblanceolate or obovate-lanceolate; 300–750 m. .................................................................................................................. 3. C. costatum. 3. Blades linear-lanceolate; (300–)750–2200(–2700 m). ...................................................................................................................... 9. C. xalapense. 2. Main lateral veins obscure; rhizomes strongly pruinose; rhizome scales 3–6 ⫻ 1–2.5 mm, entire; sori 1–4-seriate between midribs and blade margins. 4. Blades (1–)1.7–3.5 cm wide; sori 2–4-seriate between midribs and blade margins; rhizome scales 3–6 ⫻ 1.5–2.5 mm; (1150–) 1750–2700(–3000) m. ...................................................................................................................................................................... 1. C. amphostenon. 4. Blades 0.4–1.2(–1.6) cm wide; sori 1–2-seriate between midribs and blade margins; rhizome scales 3–4 ⫻ 1–1.5 mm; (150–) 900–2200 m. 5. Cells of rhizome scales oblong, often at least twice or more as long as wide; scales each with apex acuminate; blades usually straight. .........................................................................................................................................................................2. C. angustifolium. 5. Cells of rhizome scales roundish to squarish; scales each with apex rounded to shortly acute; blades often curved. ... 4. C. ensifolium. 1. Main lateral veins clearly visible on adaxial blade surfaces; fronds 2–8(–12) cm wide; sori 6–15-seriate between midribs and blade margins. 6. Rhizomes long-creeping, 3 mm or less diam., greenish; rhizome scales 0.4–1 mm wide; fronds sessile or extremely short-stiped; 100–1000 m elevation. 7. Blades greater than 4–6(–8) cm wide; rhizome scales brown, bicolorous, with narrow, lighter colored margins. ..................... 6. C. repens. 7. Blades 2–4(–5.5) cm wide; rhizome scales dark brown, concolorous. ...................................................................................... 7. C. serpentinum. 6. Rhizomes short-creeping, 4 mm or more in diam., black or brown; rhizome scales 1–1.5 mm wide; fronds sessile or long-stiped; 0–2300 m elevation. 8. Fronds sessile or extremely short-stiped, much less than 1⁄3 of frond length; blade tips acute; 0–900(–1400) m elevation. ................................................................................................................................................................................................................ 5. C. phyllitidis.

8. Fronds long-stiped, 1⁄3–1⁄2 of frond length; blade tips caudate; 1000–2300 m elevation. ............................................................ 8. C. tenuipes.

1. CAMPYLONEURUM AMPHOSTENON Campyloneurum amphostenon (Kunze ex Klotzsch) Fe´e, Me´m. Foug. 5: 258. 1852. Figs. 65C, D, 66G. Polypodium amphostenon Kunze ex Klotzsch, Linnaea 20: 399. 1847. Polypodium angustifolium Sw. var. amphostenon (Kunze ex Klotzsch) Baker in Martius, Fl. Bras. 1(2): 530. 1870. Campyloneurum angustifolium (Sw.) Fe´e var. amphostenon (Kunze ex Klotzsch) Farw., Amer. Midl. Naturalist 12: 296. 1931. Type. Venezuela. Me´rida, Moritz 120b (holotype B; isotypes BM, K!, frag. NY!).

Rhizomes moderately long-creeping (⬎5 mm between stipe bases), pruinose, (2–)3–5 mm diam.; rhizome scales dense, tan to brownish, ovate-lanceolate, 3–6 ⫻ (1–)1.5–2.5 mm, clathrate, the cells elongate, at least twice as long as wide; fronds 30–70(–100) cm long, usually somewhat distant (often 5–10 mm apart); stipes mostly 3–10 cm long; 1⁄10–1⁄5 the length of the blades; blades subcoriaceous, narrowly elliptic, (1–)1.7–3.5 cm wide, bases subabruptly tapering, apices acuminate; lime dots absent adaxially; areoles 2–4(–5) rows between midribs and margins, excurrent veinlets 1 or 2 per primary areole; main lateral veins immersed, faintly if at all discernible, secondary veins irregularly anastomosing; sori 2–4-seriate, in Ⳳ irregular rows between midribs and blade margins, one or two sori per primary areole.

Distribution. Epiphytic and on moist rocks in montane forests, pine-fir forests, pine-oak-fir forests, evergreen cloud forests; (1150–)1750–2700(–3000) m. Mexico; Guat, Hond, Salv, CR, Pan; Cuba, Jam, Hisp; Col, Ven, Ec, Peru, Bol. The specimen cited by Leo´n (1992) from nw Argentina (Fabris et al. 5135, UC) is probably C. lorentzii (Hieron.) Ching. Selected Specimens Examined. Chis (Breedlove 26260, DS, NY). DF (Lyonnet 1634, MEXU). Gro (Rzedowski & McVaugh 258, ENCB, MEXU, NY). Hgo (Sa´nchez M. 155, US). Jal (McVaugh 10084, MEXU, MICH, NY, US). Me´x (Purpus 1841, NY, US). Mor (Lyonnet 471200020, MEXU). Oax (Mickel 3829, NY, UC). Pue (Rzedowski 29998, ENCB, NY). SLP (Schaffner 76, NY). Ver (Ventura A. 7921, ENCB, F, NY).

This species is closely related to C. angustifolium, with which it has often been combined. Campyloneurum amphostenon differs from that in having broader blades (and hence more rows of sori between the midribs and blade margins), longer stipes, and longer and wider, lighter colored rhizome scales. We have seen one mixed collection of the two species, Dorantes et al. 5120 (NY), from Edo. Veracruz, Las Vigas, 2200 m, and on this gath-

CAMPYLONEURUM ANGUSTIFOLIUM

ering, the differences in rhizome scales and soriation can readily be seen. Schaffner 76 (NY) is represented by two sheets, one labeled “Valle de Me´xico” collected in 1875, the other labeled “San Luı´s Potosı´” with no collection date. Leo´n (1992) recognized C. amphostenon var. irregulare (Lellinger) B. Leo´n as also occurring in Mexico, and cited a single collection, from Veracruz: Nee & Hansen 18701 (F). We have not seen this specimen, but doubt its distinctness from var. amphostenon. Var. irregulare (type from Ecuador) was said (by Leo´n) to be distinguished by the irregular arrangement of the cells in most of the stem scales; it was ascribed a range of s. Mexico (Ver), Guatemala, Costa Rica, Panama, Ecuador, Peru, and Bolivia by Leo´n. Leo´n (1992; in Davidse et al., 1995) cited Mexico in the range of C. densifolium (Hieron.) Lellinger, a species said to occur from Mexico to Bolivia and the Greater Antilles. Campyloneurum densifolium was distinguished by Leo´n by light brown, ovatelanceolate (vs. dark brown and lanceolate) subappressed (vs. spreading) rhizome scales generally two times longer than wide (vs. three or more times longer than wide in C. amphostenon); in addition, the cells of the scales were said to be almost isodiametric in C. densifolium (vs. longer than wide in C. amphostenon). The distribution of C. densifolium in Mexico was given by Leo´n as Durango (Ownbey & Ownbey 1935, GH, NY, US) and San Luı´s Potosı´ (Pringle 3821, BM, F, GH, MO, NY); we have seen both of these gatherings in various herbaria and regard the former as C. ensifolium and the latter as C. angustifolium. There are specimens from southern Mexico, especially Chiapas, that possess some of the characteristics of C. densifolium, but we are not convinced that the species can be distinguished insofar as Mexican specimens are concerned.

2. CAMPYLONEURUM ANGUSTIFOLIUM Campyloneurum angustifolium (Sw.) Fe´e, Me´m. Foug. 5: 257. 1852. Figs. 65A, B, 66F. Polypodium angustifolium Sw., Prodr. 130. 1788. Type. Jamaica. Swartz s.n. (S, not found; isotypes BM, LD, S).

Rhizomes short-creeping, pruinose, 3–5 mm diam.; rhizome scales dense at the rhizome apices, dark brown, lanceolate, 3–6 ⫻ 0.7–1(–1.5) mm, clathrate, the cells elongate, at least twice as long as wide; fronds 15–70 cm long, clumped; stipes absent or extremely short, 2 cm long, usually not clearly demarcated; blades subcoriaceous, linear, 0.5–1.2(–1.6) cm wide, bases gradually tapering, apices long-acuminate; lime dots absent adaxially; areoles 1 or 2(–3) rows between midribs and margins, excurrent veinlet 1 per areole; main lateral veins immersed and not discernible, secondary veins irregularly anastomosing; sori mostly in single or irregular double rows running the length of the blades, one Ⳳ regular or two irregular rows on each side of the midrib; 2n⫽74 (Peru), 148 (Jam, CR).

167

Distribution. Epiphytic and epipetric in wet montane forest, pine forests, pine-oak-Liquidambar forests, evergreen cloud forests; (150–)900–2200 m. USA (Fla); Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Gr & L Ant; Col, Ven, Ec, Peru, Braz, Bol, nw Arg. Selected Specimens Examined. Chis (Croat & Hannon 65085, UC). Gro (Mexia 9097a, UC). Hgo (Moore 3211, UC, US). Jal (McVaugh 21447, IEB, MEXU). NL (Hinton 24585, MEXU). Oax (Mickel 6887, NY, UC, US). Pue (Sa´nchez M. 599, US). Qro (Rzedowski 42563, IEB). SLP (Pringle 3821, BR, DS, K, MEXU, NY, UC, US). Tab (Cowan 3423, NY). Tam (Hinton 25115, MEXU, US). Ver (Purpus 7238, UC). Unverified, Doubtful, or Mistaken Reports. DF (reported by Mickel & Beitel, 1988, and Mickel, 1992, but not verified). Me´x (reported by Mickel & Beitel, 1988, and Mickel, 1992, but not verified; also reported by Tejero-Dı´ez & Arreguı´n-Sanchez, 2004, but specimens are C. amphostenon). Son (Van Devender 95-902, ARIZ, MO, UCR, USON, reported by Reina Guerrero in Madron˜o 44:209. 1997, but not verified; specimens could be C. ensifolium).

Smith (1981), Stolze (1981), Mickel and Beitel (1988), Leo´n (in Tryon & Stolze, 1993; in Davidse et al., 1995), and many others, in floristic accounts, have chosen to synonymize C. ensifolium under C. angustifolium, or have ignored it altogether. We now believe that it is subtly distinguishable, and occupies a more-or-less discrete range. However, further study, especially cytological and isozymic, is needed. The marked variation in cell size and shape of rhizome scales of C. angustifolium s.l. suggests to us that hybridization and polyploidy may be responsible for some of the differences seen. Even with the removal of C. ensifolium, there is an element within the C. angustifolium complex in Mexico that appears to be substantively different. This entity, confined to southern Mexico and probably northern Central America, has the rhizome scales darker brown, narrower (1 mm wide), with the cell lumina very narrow (lateral walls wider than the lumina). This variant almost always occurs at lower elevations than C. angustifolium, usually below 1000 m (vs. above 1000 m for C. angustifolium s.str.; it is also much less common in Mexico than is C. angustifolium. We regard it as probably distinct specifically, but are unable to find an existing name for it; pending further study, we include it in a broad concept of C. angustifolium. Because of the narrow rhizome scales with small cell lumina, the species approaches what has been called C. angus-

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tipaleatum (Alston) Lellinger, known from Colombia to Bolivia. Specimens of this narrow-scaled form include the following, three of which were cited as C. angustifolium by Leo´n (marked by an asterisk*): Chis (*Palacios-Rios 2884, UC). Oax (Mickel 5792, UC; *Nee 32158, UC; Wendt et al. 4007, UC). Tab (Cowan 3423, UC). Ver (Calzada 10263, UC; Copeland herb. 154, MEXU, UC; *Paxson et al. 17M680, UC). Judging by the scales, C. angustifolium is probably more closely related to C. amphostenon, which has similar rhizome scales, than it is to the narrow-scaled variant with very narrow cell lumina mentioned above. A few specimens seem somewhat intermediate and difficult to place in one species or the other, e.g., Camp 2740 (UC), from Oaxaca. One collection listed above, from San Luis Potosı´, was cited under C. densifolium by Leo´n (1992); we discuss C. densifolium under C. amphostenon.

Guat, Bel, Hond, Nic, CR, Pan; Gr Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Braz. Specimens Examined. Chis (Breedlove 33188, 33190, DS, MEXU; Palacios-Rios 2802, UC; Reyes-Garcı´a & Sousa S. 2257, MEXU). Oax (Mickel 1449, US). Ver (Vera-Santos 3010, US).

This species has features of both C. xalapense and C. serpentinum, and differs from the former in having shorter petioles, oblanceolate blades, and growing at generally lower elevations. It is a much less common species than is C. xalapense. From C. serpentinum (and C. repens) it differs in its shorter-creeping rhizomes and much less obvious secondary venation, and thinner blade texture. Campyloneurum aphanophlebium (Kunze) T. Moore, which bears short hairs on the petioles and abaxially at the bases of the blades, apparently does not occur in Mexico, although it is known from Belize, Honduras, and further south (Leo´n, 1992, and in Davidse et al., 1995).

3. CAMPYLONEURUM COSTATUM 4. CAMPYLONEURUM ENSIFOLIUM Campyloneurum costatum (Kunze) C. Presl, Suppl. Tent. Pterid. 190. 1836. Figs. 66C, D. Polypodium costatum Kunze, Linnaea 9: 38. 1834. Type. Cuba. Limonar, Poeppig s.n. (LZ destroyed; isotypes B, BM!, C, K!, L–photos C!, P!).

Rhizomes short-creeping, not pruinose, 3–6 mm diam.; rhizome scales dense, brown, lustrous, narrowly lanceolate, 3–5 ⫻ 0.8–1(–1.5) mm, clathrate, the cells elongate, at least twice as long as wide; fronds 20–60 cm long, crowded to somewhat distant; stipes 1⁄10–1⁄5 the length of the blades; blades chartaceous to subcoriaceous, oblanceolate or obovate-lanceolate, 3–6 cm wide, bases attenuate or narrowly cuneate, apices generally caudate; lime dots absent adaxially; areoles 5–10 rows between midribs and margins, excurrent veinlets 3 or 4 per primary areole, the middle one completely bisecting the areole; main lateral veins inconspicuous or faintly visible on both sides, secondary veins regularly anastomosing; sori 5–10-seriate between the midribs and blade margins, in two very regular rows between main lateral veins, 2 sori per primary areole; 2n⫽74 (CR, Jam).

Distribution. Epiphytic or on limestone rocks in lowland and lower montane rain forests; 300–750 m. USA (s Fla); Mexico;

Campyloneurum ensifolium (Willd.) J. Sm., Cat. Cult. Ferns 12. 1857. Fig. 66E. Polypodium ensifolium Willd., Sp. Pl., ed. 4, 5(1): 152. 1810. Polypodium angustifolium Sw. var. ensifolium (Willd.) Hicken, Revista Mus. La Plata, Secc. Bot. 5: 271. 1908. Campyloneurum angustifolium (Sw.) Fe´e var. ensifolium (Willd.) Farw., Amer. Midl. Naturalist 2: 296. 1931. Type. Said to be from Peru, Ne´e s.n. (B-Willd. 19610, fiche UC!; photos BM, USM), but Lellinger (1988: 34) and Leo´n (1992) speculated that Ne´e probably collected it in Mexico.

Rhizomes short-creeping, usually not pruinose, 2–4(–5) mm diam.; rhizome scales dense, tan to brownish, ovate-lanceolate, 3–4 ⫻ 1–1.5 mm, clathrate, the cells roundish to squarish or at least isodiametric; fronds 15–60 cm long, clumped (1–4 mm apart); stipes absent or extremely short, not clearly demarcated; blades coriaceous or thick-chartaceous, linear, 0.4–1 cm wide, gradually tapering at bases to the rhizomes or nearly so, longacuminate at apices, often curved, margins often strongly revolute and cartilaginous; lime dots absent adaxially; areoles 1 or 2 rows between midribs and margins, excurrent veinlet one per primary areole; main lateral veins immersed and not discernible, secondary veins irregularly anastomosing; sori mostly in single rows running the length of the blades, one row on each side of the midrib, rarely an incomplete, irregular second row present outside of the first row.

CAMPYLONEURUM PHYLLITIDIS

169

mm, subclathrate, the cells elongate, at least twice as long as wide; fronds (30–)50–120 cm long, clumped (2–5 mm apart); stipes absent or extremely short (to narrowly winged to 15 cm), often not clearly demarcated; blades rigidly chartaceous to subcoriaceous, often lustrous, oblanceolate, 5–8(–12) cm wide, gradually tapering nearly to rhizomes at bases, apices acute to acuminate; lime dots absent adaxially; areoles (6–)8–16 rows between midribs and margins, excurrent veinlets 3 or 4 per primary areole, a middle one usually completely bisecting the areole; main lateral veins prominent, secondary veins regularly anastomosing; sori 6–15-seriate between the midribs and blade margins, in two very regular rows between main lateral veins, 2 sori per primary areole; 2n⫽74 (Fla, Jam, Galapagos), 148 (Jam). Distribution. Epiphytic or occasionally on mossy boulders, in lowland and montane forests; 200–1800 m. Mexico; Guat, Salv (Jaurequi 87, UC). Leo´n (1992) also cited specimens from Honduras and Nicaragua, but these have not been verified. Selected Specimens Examined. Chih (Spellenberg 12030, MEXU, UC). Chis (Croat 78450, UC; Go´mez Lo´pez 278, UC; Purpus 7238, UC). Col (Espero´n Mun˜oz 18298, UC; Lott 2953, UC). Dgo (Ownbey & Ownbey 1935, NY, UC, US). Jal (Dı´az-Luna 5598, 8933, UC). Nay (Norris 13199, UC). Oax (Mickel 6887, UC). Unverified, Doubtful, or Mistaken Reports. Mich (Hinton et al. 15685, MO, US, and 15543, MO, cited by Leo´n, 1992, but not verified). Sin (Breedlove 35732, MO, cited by Leo´n, 1992, but not verified). Ver (Barnett et al. 85a, MO; Matuda 208, MO; Nee & Hansen 18570, F; Nee & Taylor 26818, F; Nee 29080, F, all cited by Leo´n, 1992, but not verified).

This species is closely related to C. angustifolium and is often regarded as a variety of that, but Lellinger (1988) and Leo´n (1992) clarified the species’ differences, namely that C. ensifolium has rhizome scales with cells rounded to squarish or at least isodiametric (vs. elongate, 2–3 times as long as wide), scales generally acute at the tips and curling at the edges, and blades often narrower and usually curved (vs. straight). From C. amphostenon, C. ensifolium differs by the narrower, curved, coriaceous and revolute-margined blades, fewer rows of vein areoles and sori between midribs and blade margins, narrower rhizome scales, and occurrence at lower elevations. Campyloneurum ensifolium seems to occur predominantly along the Pacific Coastal states in Mexico to El Salvador (and perhaps further south). It is certainly much more common in Mexico than C. angustifolium in states northwest of Guerrero.

5. CAMPYLONEURUM PHYLLITIDIS Campyloneurum phyllitidis (L.) C. Presl, Suppl. Tent. Pterid. 190. 1836. Figs. 65L–N. Polypodium phyllitidis L., Sp. Pl. 2: 1083. 1753. Lectotype (chosen by Proctor, 1977: 341). Plumier, Descr. Pl. Ame´r., pl. 38, 1693, representing a plant from “Antilles.”

Rhizomes short-creeping, not pruinose, 5–8 mm diam.; rhizome scales dense, brown, deltate-lanceolate, 4–8 ⫻ 1.5–2(–3)

Distribution. Epiphytic or epipetric in wet montane forests, tropical subdeciduous forests, evergreen rain forests; 0–900(– 1400) m. USA (Fla); Mexico; Guat, Bel, Hond, CR, Pan; Bah, Gr & L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol. Selected Specimens Examined. Camp (Martı´nez S. 28001, XAL). Chis (Breedlove 31569, DS, NY). Gro (Lorea 113, IEB). Hgo (Moore 3038, UC, US). Jal (Alcocer R. 11111, UC). Mich (Hinton 15969, DS, NY, UC). Oax (Herna´ndez G. 284, ENCB, IEB, MEXU). Pue (Campos V. 342, IEB, MEXU). Qro (Dı´az Barriga & Zamudio 4632, ENCB, IEB, MEXU). QR (Ucan & Flores 939, MEXU). SLP (King 4284, NY, UC). Tab (Cowan et al. 3540, NY, UC). Ver (Mickel 7227, NY, UC). Yuc (Schott 756, MO).

Specimens determined as Camplyoneurum latum T. Moore or C. brevifolium (Lodd. ex Link) Link in various herbaria (e.g., Mickel 5846, NY, UC) are referable here. Both Lellinger (1989) and Leo´n (1992, and in Davidse et al., 1995) considered C. phyllitidis and C. latum [⫽ C. brevifolium] to occur in Mexico. The distinctness of C. brevifolium from C. phyllitidis remains in doubt, and even if the two are not the same elsewhere in the range, Mexican specimens so identified as C. latum are not distinguishable from C. phyllitidis. Leo´n (1992) cited one of the specimens listed above (Alcocer R. 11111 from Jalisco) as C. brevifolium; we have also seen the three Mickel collections she cited as C. brevifolium from Oaxaca, and all are inseparable from C. phyllitidis, in our opinion. The venation in Mexican material of this species is variable, with the primary areoles being often subdivided, not only by

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6. CAMPYLONEURUM REPENS

the broader, weakly bicolored rhizome scales. Campyloneurum fasciale was treated by Leo´n (1992; in Davidse et al., 1995) as the accepted name for what is here called C. serpentinum, but the type of Polypodium fasciale, from eastern Venezuela, seems indistinguishable from the many specimens of C. repens seen from that same area. More study of this group is needed.

Campyloneurum repens (Aubl.) C. Presl, Suppl. Tent. Pterid. 190. 1836. Figs. 66A, B.

7. CAMPYLONEURUM SERPENTINUM

excurrent veinlets but also by some recurrent and/or forking veinlets. The fronds are broad and the margins undulate to a greater degree than in other parts of the range of this species.

Polypodium repens Aubl., Hist. Pl. Guiane 2: 962. 1775, non Sw., 1788. Lectotype (chosen by Proctor, 1985: 340). Plumier, Traite´ Foug. Ame´r., pl. 134, 1705, based on material from Morne Rouge, Martinique. Polypodium fasciale Willd., Sp. Pl., ed. 4, 5(1): 156. 1810. Campyloneurum fasciale (Willd.) C. Presl, Suppl. Tent. Pterid. 190. 1836. Type. Venezuela [Edo. Monagas], Caripe, Humboldt 426 (B-Willd. 19632, fiche UC!, isotype P).

Rhizomes slender, creeping, mostly 2–3 mm diam., green, nearly naked at maturity; rhizome scales sparse, narrowly lanceolate-deltate with long-attenuate tips, 2–4 ⫻ 0.5–1 mm, weakly clathrate, the cells elongate, at least twice as long as wide, margins lighter colored; fronds 20–50 cm long, distant (7–15 mm apart); stipes absent or very short, 1–10 cm long, 1⁄20–1⁄5 the length of the blades; blades chartaceous, narrowly elliptic to narrowly oblanceolate, 4–6(–8) cm wide, narrowly cuneate and decurrent at bases, apices acute to long-acuminate; lime dots absent adaxially; areoles (4–)6–12 rows between midribs and margins, 1.5–3 times broader than long, excurrent veinlets usually 2 per primary areole, free, not partitioning the areoles; main lateral veins prominent, secondary veins regularly anastomosing; sori 6– 12-seriate between midribs and blade margins, in two regular rows between main lateral veins, two per primary areole.

Distribution. Epiphytic in wet montane forests, tropical rain forests, tropical evergreen forests; 130–600 m. Mexico; Guat, Bel, Hond, Nic, CR, Pan; Cuba, Jam, PR, L Ant; Col, Ven, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol. Specimens Examined. Chis (Breedlove 34063, DS, NY, 57985, CAS; Martı´nez S. 23395, 24977, 26130, MEXU).

Campyloneurum repens is hardly distinguishable from C. serpentinum of Central and South America, but several specimens more nearly resemble C. repens in the broader fronds, more numerous series of sori between midribs and blade margins, and

Campyloneurum serpentinum (Christ) Ching, Sunyatsenia 5: 263. 1940. Figs. 65J, K. Polypodium serpentinum Christ, Bull. Herb. Boissier, se´r. 2, 6: 51. 1906. Type. Costa Rica. Navarro, Werckle´ s.n. (P, photo BM).

Rhizomes greenish, long-creeping, 1–2.5 mm diam., twining (the plants vine-like), not pruinose; rhizome scales sparse, dark brown, concolorous, linear-lanceolate to subulate, 2–3 ⫻ 0.4–1 mm, clathrate, the cells elongate, at least twice as long as wide, lacking differentiated margins; fronds 18–40 cm long, distant (to 20 mm apart); stipes 1–2(–3) cm long, less than 1⁄8 the blade length; blades herbaceous to chartaceous, narrowly elliptic, 2– 4(–5.5) cm wide, tapering gradually on both sides of the middle, proximally nearly to the rhizome, bases attenuate, apices acuminate; lime dots absent adaxially; areoles 5–8 rows between midribs and margins, 1.5–2 times broader than long, excurrent veinlets usually 2 per primary areole, free, not partitioning the areoles; main lateral veins prominent, straight to slightly flexuous, secondary veins regularly anastomosing; sori 5–8-seriate between midribs and blade margins, in two regular rows between main lateral veins, two per primary areole.

Distribution. Epiphytic in lowland rain forests, seasonal evergreen forests, pine-oak forests; 250–1000 m. Mexico; Bel, CR, Pan; Col, Ven, Braz. Leo´n (in Davidse et al., 1995) included French Guiana, Ecuador, Peru, and Bolivia in the range, but her circumscription of the species, which included C. fasciale, differs somewhat from the one adopted here (see C. repens for discussion). Selected Specimens Examined. Chis (Breedlove 32816, DS, MEXU; Dressler 1617, MEXU, NY, US; Matuda 207, MEXU, US; Purpus 6751, NY, UC, US). Gro (Lorea 2534, FCME, NY). Oax (Cruden 1113, UC;

CAMPYLONEURUM XALAPENSE

171

Herna´ndez G. 1391, 1465, CAS, CHAPA, MEXU, NY). Tab (Cowan 2967, CHAPA, ENCB, MEXU, NY). Ver (Conzatti & Gonza´lez 590, MEXU, US; Dorantes et al. 2453, UC; Mu¨ller 2227, K, NY).

and blade margins, in two regular rows between main lateral veins, each sorus in its own areole; 2n⫽74 (Oax, Chis).

Leo´n (1992, and in Davidse et al., 1995) treated this taxon under the name C. fasciale, which is here regarded as a synonym of C. repens (see above). Campyloneurum serpentinum is indeed very closely related to C. repens and differs from that by having narrower blades that are gradually reduced at both distal and proximal ends; also, the rhizome scales of C. serpentinum are undifferentiated at the margins, contrasting with differentiated margins in C. repens. Leo´n (1992, and in Davidse et al., 1995) further utilized cell shape in the scales, as a distinguishing feature: the central cells in scales of C. repens were said to be 1–2 times longer than broad (vs. 3–4 times longer than broad in C. serpentinum); we are unable to apply this difference to specimens we have studied, and in fact the character is often difficult to assess because there are so few scales on the rhizome in C. serpentinum. Some specimens here regarded as C. serpentinum were cited as C. repens by Leo´n, e.g., Cruden 1113 (UC), Mickel 6374 (NY, UC), from Oaxaca; Mickel and Beitel (1988) cited Herna´ndez 1391 and 1465, as well as Mickel 6374 from Oaxaca under C. repens, but Herna´ndez 1465 was cited as C. serpentinum by Leo´n. A few collections from Chiapas seem somewhat intermediate between the two species, and are difficult to place. Specimens cited as C. occultum (Christ) L. D. Go´mez [⫽ C. aphanophlebium (Kunze) T. Moore] by Smith (1981) for Chiapas are C. serpentinum vel aff., differing primarily in the more obscure main lateral veins and also the quite obscure connecting veins; it is quite possible that these specimens (Breedlove 26527, 35385, DS; also Breedlove 48384, CAS) represent an undescribed species or distinct variety. Campyloneurum serpentinum appears to be much more common in southern Mexico (21 collections seen) than is C. repens (three collections seen), which is known only from Chiapas.

Distribution. Terrestrial or epipetric, rarely epiphytic in wet montane rain forests, evergreen cloud forests, pine-oak-Ostrya forests, Liquidambar forests, woods along streams; 1000–2300 m. Mexico; Guat, Hond.

8. CAMPYLONEURUM TENUIPES

Selected Specimens Examined. Chis (Breedlove 22308, DS, NY; Miller et al. 2697, UC). Col (Sanders 11708, NY, UC). Gro (Hinton 14059, NY, UC; Lorea 2890, IEB, UC). Oax (Mickel 4828, 5097, 6050, NY, UC). Pue (Ventura A. 22364, IEB, NY). Qro (Carranza 1049, IEB). Ver (Conzatti 590, MEXU). Unverified, Doubtful, or Mistaken Reports. Mich (Hinton 11177, US, cited by Leo´n, 1992, but not verified).

Campyloneurum tenuipes is similar to C. xalapense, but has longer stipes, larger rhizome scales with scales more decidedly clathrate and larger cell lumina, conspicuous lateral veins, and 6–10-seriate sori.

9. CAMPYLONEURUM XALAPENSE Campyloneurum xalapense Fe´e, Me´m. Foug. 5: 258. 1852. Fig. 65H.

Polypodium tenuipes (Maxon) C. Chr., Index Filic., Suppl. 1: 63. 1913. Type. Guatemala. Alta Verapaz: Near Coba´n, Tu¨rckheim II 1952 (US!; isotype US!).

Polypodium xalapense (Fe´e) Christ, Bull. Soc. Roy. Bot. Belgique. 35(1): 231. 1896. Type. Mexico. [Veracruz:] “Xalapaˆ Mexicanorum,” Galeotti 6273 (P!; isotypes BR!, K!, frag. NY!). Campyloneurum caudatum Fe´e, Me´m. Foug. 8: 96. 1857. Polypodium weatherbyanum F. Seym., Phytologia 31: 171. 1975, non P. caudatum Raddi, 1819. Type. Mexico. [Veracruz:] “Cordoba et Huatusco,” Schaffner 176 (P?; isotype K!).

Rhizomes short-creeping, 4–7 mm diam., brown; rhizome scales dense, brown, linear-lanceolate, each with a nearly hairlike tip and peltate base, 5–8 mm long, 1–1.5 mm wide, clathrate; fronds 30–80 cm long, clumped (2–10 mm apart); stipes 8–18 cm mm, 1⁄3 to equalling the blade length; blades chartaceous, narrowly elliptic to linear-oblong, 4–9 cm wide, bases cuneate, apices abruptly narrowed and caudate; lime dots absent adaxially; areoles 6–10 rows between midribs and margins, excurrent veinlets 2 or 3 per areole, if 3, the middle one sometimes completely bisecting the areole; main lateral veins prominent, secondary veins regularly anastomosing; sori 6–10-seriate between midribs

Rhizomes short-creeping, 4–7(–10) mm diam., blackish or slightly pruinose; rhizome scales dark brown, 2–4 mm long, 0.8–1 mm wide, obscurely clathrate, lanceate with expanded bases, acuminate at tips, margins entire to faintly erose; fronds (20–) 30–60 cm long, clumped; stipes 2–12(–25) cm, ca. 1⁄10 the blade length; blades narrowly elliptic or linear-oblong, often parallelsided for much of their length, 3–5.5 cm wide, bases cuneate and then abruptly curving to meet the stipes, apices acuminate or often abruptly narrowed and long-caudate; lime dots often present adaxially; areoles 4–7 rows between midribs and margins, excurrent veinlets 2–4 per primary areole; main lateral veins im-

Campyloneurum tenuipes Maxon, Contr. U.S. Natl. Herb. 13: 7. 1909. Figs. 65E–G.

172

CERADENIA

mersed to faintly discernible, secondary veins Ⳳ regularly anastomosing; sori 4–6-seriate in two Ⳳ regular rows between midribs and blade margins; 2n⫽74 (Oax), 148 (CR).

Distribution. Epiphytic, occasionally epipetric in montane forests, pine-oak forests, hardwood forests, limestone cliffs; (300–) 750–2200(–2700 m). Mexico; Guat, Bel, Hond, Salv, Nic, CR. Selected Specimens Examined. Chis (Davidse et al. 29866, UC). Gro (Martı´nez S. 982, MEXU). Hgo (Moore 3019, UC, US). Nay (McVaugh 18988, MEXU, NY). Oax (Mickel 7167, NY, UC). Pue (Campos V. 405, IEB, MEXU). Qro (Rubio 2566, IEB, UC). SLP (Rzedowski 10944, NY). Tab (Cowan 2967, NY). Ver (Pringle 6082, NY, UC; Purpus 2163, NY, UC). Unverified, Doubtful, or Mistaken Reports. Jal (McVaugh 20436, MICH, cited by Mickel, 1992, but not verified).

Campyloneurum xalapense can be distinguished from congeners in Mexico by the presence of lime dots on the adaxial surface of the blades, by the more or less parallel-sided blades that are reduced proximally and then abruptly meeting the stipes, by the caudate blade tips, and by the stramineous, raised, angular midribs abaxially. It seems most closely related to C. tenuipes, from which it also differs by having shorter stipes and nonclathrate rhizome scales with smaller, less transparent cell lumina. Campyloneurum xalapense is somewhat intermediate between C. tenuipes and the C. ensifolium-amphostenon complex; one wonders if allopolyploidy is involved. Excluded Species Campyloneurum aphanophlebium (Kunze) T. Moore [syn.: C. occultum (Christ) L. D. Go´mez]. See comments under C. serpentinum.

2 3 . C E R A DE N I A Ceradenia L. E. Bishop, Amer. Fern J. 78: 2. 1988. Type: Ceradenia curvata (Sw.) L. E. Bishop [⬅ Polypodium curvatum Sw.]. Most commonly epiphytic, at times epipetric or terrestrial at high elevation; rhizomes short-creeping or erect, dorsiventral or

radially symmetrical; rhizome scales orangish to castaneous or atropurpureous, non-clathrate, usually setulose or with marginal glands (rarely, scales absent: C. jungermannioides); fronds monomorphic, determinate (ours) or indeterminate; stipes often articulate, bearing setae and, abaxially, usually branched hairs; blades rarely simple, commonly pinnatifid, pinnatisect, or perpinnate; costae not readily visible abaxially; veins usually obscure, pinnate, simple or often 1-forked from the costae; hydathodes absent, vein tips not visible adaxially; indument of reddish brown to atropurpureous, stiff setae often 1–3 mm long on stipes, rachises, costae, and sometimes laminae, also of shorter branched or unbranched hairs, these often septate and sometimes bearing whitish globose or oblong glands, usually 2–5 glands per hair; sori round or slightly oblong, superficial or slightly impressed; indusia absent; soral paraphyses present, in the form of hairs bearing whitish to occasionally tan or yellowish wax-like globose or pyriform glands that function to protect the developing sporangia (and obscure them in young sori); sporangia glabrous; spores tetrahedral-globose; x⫽37. Ceradenia is widespread in the wet Neotropics and comprises about 55 species; a single species (C. jungermannioides) extends to the Azores, and another (C. pruinosa) is probably conspecific with the African species Ctenopteris punctata F. Ballard. The genus is most closely related to Enterosora (which see) and probably also to Zygophlebia, with about eight, mostly neotropical species. Recent molecular work on Grammitidaceae by Ranker et al. (2004) indicates that Ceradenia is very likely monophyletic. Soral paraphyses, whitish glands, and anhydathodous blades distinguish Ceradenia from other Mexican genera of Grammitidaceae. The glands are usually globose or pear-shaped and are borne terminally and laterally on short uniseriate hairs; the glands may completely hide the young sporangia and can be mistaken for spores when sporangia mature. Specimen-drying treatments, especially if an alcohol-based preservative is used before drying, may obscure or apparently destroy the glands. Some species in subg. Ceradenia (e.g., C. pruinosa) also have branched gland-bearing hairs on the laminar tissue, on the abaxial surface or on both surfaces. Often there are three (2–5) globose or oblong glands per hair. In Mexico and usually wherever they grow, species of Ceradenia are infrequent or rare, often very local, and usually found only in relatively undisturbed cloud forests with constant high humidity. There are relatively few collections of all four Mexican species. Reference Bishop, L. E. 1988. Ceradenia, a new genus of Grammitidaceae. Amer. Fern J. 78: 1–5.

Key to the Mexican Species of Ceradenia 1. Blades simple, entire; rhizome scales absent. .. 1. C. jungermannioides. 1. Blades pinnatifid or pinnatisect; rhizome scales present. 2. Stipes absent or, if present, less than 1⁄4 the blade length. ......................................................................... 4. C. pruinosa. 2. Stipes equalling or often exceeding blade length.

CERADENIA OIDIOPHORA 3. Rhizome scales atropurpureous; blades abaxially lacking white glands, except in sori. ........... 2. C. margaritata. 3. Rhizome scales orangish; blades abaxially with numerous whitish, oblong glands. ................. 3. C. oidiophora.

1. CERADENIA JUNGERMANNIOIDES Ceradenia jungermannioides (Klotzsch) L. E. Bishop, Amer. Fern J. 78: 4. 1988. Figs. 67C, D. Polypodium jungermannioides Klotzsch, Linnaea 20: 373. 1847. Grammitis jungermannioides (Klotzsch) Ching, Bull. Fan Mem. Inst. Biol. 10: 240. 1941. Type. Venezuela. Me´rida: Moritz 312 (B; isotype BM!).

Rhizomes ascending, radially symmetric, caudices 3–5 mm diam.; rhizome scales absent; fronds tightly clustered, to 7(–10) cm long; stipes dark brown, 3–5 ⫻ 0.2–0.3 mm, with reddish brown setae 1–2 mm long; blades herbaceous, narrowly ligulate, entire to repand at margin, 2.5–7(–10) cm ⫻ 2–4 mm, bases attenuate, apices obtuse; segments absent; veins simple or 1-forked from midrib; indument abaxially (on and between veins) of scattered, reddish brown setae 1–2 mm long and also of sparse, gland-bearing hairs 0.1–0.2 mm; indument adaxially of scattered, reddish brown setae 1–2 mm long; sori medial to supramedial, in a single row on each side of blade midrib.

173

Grammitis margaritata A. R. Sm., Proc. Calif. Acad. Sci., ser. 4, 40: 221, f. 5D–E. 1975. Type. Mexico. Chiapas: 3–4 km W of El Porvenir, Breedlove 31801 (DS!).

Rhizomes short-creeping, dorsiventral, caudices 6–10 mm diam.; rhizome scales castaneous, to ca. 5 ⫻ 0.5 mm, margins bearing whitish setulae 0.1–0.3 mm long; fronds closely spaced, 15–25 cm long; stipes dark red-brown, to ca. 14(–20) cm ⫻ 1 mm, setose to glabrescent, setae dark red-brown, to 3.5 mm long; blades subcoriaceous, lanceolate to narrowly deltate, pinnatisect, to 12 ⫻ 2.5 cm, bases truncate, apices acuminate; segments ca. 20 pairs, to 15 ⫻ 4.5 mm, bases dilate and adnate, margins sparingly setose, also with a few branched hairs, apices rounded or subacute; veins to ca. 7 pairs per segment; indument abaxially absent on costae, veins, and laminar tissue, or of scattered branched hairs to 0.3 mm mainly on costae and veins; indument adaxially absent, except for setae along rachis; sori medial to supramedial, to 7 pairs per segment, in single rows on each side of costae; 2n⫽74 (Chis).

Distribution. Epipetric on mossy rock ledges in cloud forests; 2800 m. Mexico; Guat, Hond, Salv. In Mexico known only from the type.

Distribution. Epiphytic in wet montane forests; 1000–1700 m. Mexico; Guat, Hond, Salv, CR, Pan; Jam, Hisp; Col, Ven, Braz; Azores.

Ceradenia margaritata is most closely related to Antillean and Central American species in subg. Filicipecten L. E. Bishop, e.g., C. brunneoviridis (Baker ex Jenman) L. E. Bishop and C. kookenamae (Jenman) L. E. Bishop.

Specimens Examined. Chis (Breedlove 32442, 32645, DS). Oax (Hallberg 1524, NY, UC).

3. CERADENIA OIDIOPHORA

This is the only simple-bladed Ceradenia, and the only one that lacks rhizome scales. It superficially resembles, but is not closely related to, Enterosora parietina, which lacks the peculiar soral paraphyses, has golden brown rhizome scales 0.5–0.8 mm long, and has laminar setae that tend to be clustered around and through the sori.

Ceradenia oidiophora (Mickel & Beitel) A. R. Sm., Novon 5: 20. 1995. Figs. 67J, K.

2. CERADENIA MARGARITATA Ceradenia margaritata (A. R. Sm.) L. E. Bishop, Amer. Fern J. 78: 5. 1988. Figs. 67E–H.

Grammitis oidiophora Mickel & Beitel, Mem. New York Bot. Gard. 46: 202. 1988. Type. Mexico. Oaxaca: Distrito Ixtla´n, trail from San Pedro Nolasco N to the Llano Verde, Mickel 3822 (NY!; isotype UC!).

Rhizomes short-creeping, dorsiventral, caudices ca. 4–6 mm diam.; rhizome scales orange, 4–5 ⫻ 0.3–0.5 mm, margins bearing whitish glands 0.1 mm long; fronds closely spaced, 10–25 cm long; stipes light brown, to 10 cm ⫻ 1 mm, with dense, reddish setae 0.8–1 mm long and short, pale, Ⳳ appressed hairs; blades subcoriaceous, lanceolate to narrowly deltate, deeply pinnatifid,

174

CERATOPTERIS

8–20 ⫻ 1.5–4 cm, bases truncate, apices acuminate; segments 12– 30 pairs, to 25 ⫻ 3 mm, bases dilate and decurrent, barely adnate, margins glabrous, apices obtuse to subacute; veins to ca. 12 pairs per segment; indument abaxially of reddish brown setae ca. 1 mm mostly on rachises and costae, setae sparse on veins, laminae also with numerous branched hairs 0.1–0.3 mm, these bearing whitish, globose, oblong, or obpyriform glands; indument adaxially of scattered setae on blade and a few minute gland-bearing hairs like those abaxially; sori medial, 5–12 pairs per segment, in single rows on each side of costae.

Distribution. Epiphytic on trunks and epipetric in exposed rock crevices in cloud forests; 2150–2550 m. Mexico.

orange-brown setae 0.8–1 mm; blades thick-herbaceous, pinnatifid, linear-oblanceolate, 4–9 cm ⫻ 8–10 mm, tapered at both ends; segments 8–12 pairs, 5–7 ⫻ 2–3 mm, bases joined, margins setose, apices obtuse; veins to 4 pairs per segment; indument abaxially, on and between veins, of red-brown setulae 0.3–0.5 mm long and gland-bearing hairs 0.1–0.2 mm, glands whitish, globose; indument adaxially of red-brown setulae 0.3–0.5 mm long; sori medial, to 4 pairs per segment, in single rows on each side of costae.

Distribution. Epiphytic in cloud forests along continental divide; 1200–1350 m. Mexico; Nic, Pan; s Ven, s Braz; Liberia? Specimen Examined. Oax (Hallberg 1724, NY).

Specimens Examined. Oax (Mickel 4927, 7045, NY; Smith 472, UC). Unverified, Doubtful, or Mistaken Reports. Gro (Lorea 4355, FCME, cited by Lorea-Herna´ndez & Vela´zquez M., 1998, but not verified).

Ceradenia oidiophora is readily distinguished from all other Mexican species of the genus by its orange rhizome scales and numerous white, oblong glands on the abaxial laminae. It has characteristics of both subg. Filicipecten (well developed stipes, dorsiventral rhizomes) and subg. Ceradenia (abaxial lamina bearing numerous whitish glands).

4. CERADENIA PRUINOSA Ceradenia pruinosa (Maxon) L. E. Bishop, Amer. Fern J. 78: 5. 1988. Figs. 67A, B. Polypodium pruinatum Baker in Hooker & Baker, Syn. Fil., ed. 2, 508. 1874, hom. illeg., non Sw., 1801. Polypodium pruinosum Maxon, Proc. Biol. Soc. Wash. 52: 117. 1939. Ctenopteris pruinosa (Maxon) Copel., Philipp. J. Sci. 84: 470. 1955 [1956]. Grammitis pruinosa (Maxon) C. V. Morton, Contr. U.S. Natl. Herb. 38: 262. 1973. Type. Nicaragua. Chontales: Tate 44 (K, frag. NY!, US!). Grammitis kirkbridei Mickel, Amer. Fern J. 74: 117. 1984. Type. Panama. Darie´n: Tres Bocas on Rı´o Coasi, Kirkbride & Duke 1381 (NY!; isotypes MO!, UC!).

Rhizomes ascending, radially symmetric, caudices ca. 3–5 mm diam.; rhizome scales orange-brown, 1.5–2 ⫻ 0.3 mm, margins setulose; fronds tightly clustered, 4–10 cm long; stipes stramineous to light brown, to 1 cm ⫻ 0.2–0.5 mm, bearing dense,

Mickel and Beitel (1988) compared this species to Grammitis flexuosa Kunth [⬅ Eriosorus flexuosus (Kunth) Copel.], but this is presumably an error for Polypodium flexuosum Maxon [⬅ Grammitis maxoniana Lellinger], which is now Terpsichore flexuosa (Maxon) A. R. Sm., known only from Cuba and Haiti. Ceradenia pruinosa is similar in frond dissection and rhizome scales to Terpsichore flexuosa, but the latter has vein tips visible adaxially and setose sporangia; it also lacks Ceradenia-type glands in the sori and on the blades. The nearest relationship of Ceradenia pruinosa is not known, but the species is certainly a member of subg. Ceradenia, comprising ca. 25 species. Ctenopteris punctata F. Ballard (Kew Bull. 1955: 468), from Liberia, is almost exactly the same as Ceradenia pruinosa except that the rhizome scales are larger, to 6 mm long. This represents an extremely unusual disjunct distribution or at least a very closely related species-pair. The report of this species from Guatemala by Mickel and Beitel (1988) has not been confirmed.

24. CERATOPTERIS Ceratopteris Brongn., Bull. Sci. Soc. Philom. Paris, se´r. 3, 8: 186. 1821. Type: Ceratopteris thalictroides (L.) Brongn. [⬅ Acrostichum thalictroides L.]. Parkeria Hook., Exot. Fl. 2: t. 147. 1825. Type: Parkeria pteridoides Hook. [⬅ Ceratopteris pteridoides (Hook.) Hieron.].

CERATOPTERIS PTERIDOIDES

Rooting in mud or free-floating; rhizomes poorly developed, short-creeping, sparsely and obscurely scaly; fronds clustered, the fertile and sterile strongly dimorphic; sterile blades simple or merely lobed to pinnatifid, or 1–3-pinnate, glabrous, succulent to herbaceous; veins areolate, lacking included veinlets; vegetative buds often present in sinuses of pinnae; fertile blades 2–4-pinnate, erect, taller than the sterile, the blade divisions linear, 1–3 mm wide, the margins inrolled over the sporangia; sori marginal, paraphyses absent; sporangia sessile or nearly so, at the ends of veins; annuli vestigial or well developed with to ca. 70 indurate cells; spores tetrahedral-globose, with Ⳳ parallel ridges, lacking equatorial flanges; x⫽39. There are three or four species of Ceratopteris in tropical regions, usually in standing or slow-moving water or rooted in mud. The genus has been widely utilized in studies of reproductive biology and fern genetics because of its relatively quick growth and ease of culture. Affinities of Ceratopteris, often placed in the monotypic family Parkeriaceae, are clearly with the Pteridaceae s.l. Tryon and Tryon (1982) and Kramer (in Kubitzki, 1990), respectively, included Ceratopteris as a tribe Ceratopterideae, or subfamily Ceratopteroideae, within the Pteridaceae. All recent molecular data support inclusion of Ceratopteris with the Pteridaceae s.l. (Gastony & Rollo, 1995, 1998; Hasebe et al., 1995; Pryer et al., 1995), or the breakup of the Pteridaceae into ca. 5 smaller monophyletic families. In all of these phylogenetic topologies, Ceratopteris is the sister group to Acrostichum. However, Ceratopteris is aberrant in the family in the chromosome base number (x⫽39), in occupying aquatic habitats, in dimorphic fronds, in spore morphology, and in the ill-defined annulus with the large number of weakly indurate cells. References Hickok, L. G. 1977. Cytological relationships between three diploid species of the fern genus Ceratopteris. Canad. J. Bot. 55: 1660–1667; Hickok, L. G. & E. J. Klekowski, Jr. 1974. Inchoate speciation in Ceratopteris: an analysis of the synthesized hybrid C. richardii ⫻ C. pteridoides. Evolution 28: 439–446; Lloyd, R. M. 1974. Systematics of the genus Ceratopteris Brongnart (Parkeriaceae) II. Taxonomy. Brittonia 26: 139–160.

Key to the Mexican Species of Ceratopteris 1. Plants generally floating; stipes often inflated or enlarged distally; proximal pinnae of sterile blades (or proximal lobes of simple blades) opposite; buds or plantlets generally developed in sinuses of pinnae of sterile blades on actively growing fronds; annuli vestigial or of 0–10(–40) poorly developed cells. .................... 1. C. pteridoides. 1. Plants generally rooted in mud; stipes nearly the same width through their length; proximal pinnae of sterile blades alternate; buds generally developed only on senescent blades; annuli with 15–70 clearly indurate cells. .............................................................................. 2. C. thalictroides.

1. CERATOPTERIS PTERIDOIDES Ceratopteris pteridoides (Hook.) Hieron., Bot. Jahrb. Syst. 34: 561. 1905. Figs. 68A–D.

175

Parkeria pteridoides Hook., Exot. Fl. 2: t. 147. 1825. Lectotype (chosen by Lloyd, 1974: 156). Guyana. Essequibo, Parker s.n.. (K–photo GH; isotype GH).

Plants floating, occasionally rooted in mud; stipes 4–17 mm wide, often inflated or enlarged distally; sterile fronds ca. 8–30 ⫻ 5–25 cm; sterile blades mostly simple and palmately 3-lobed or pinnatifid and 5-lobed (occasionally pinnate-pinnatifid at the base), the proximal pinnae (or lobes) opposite or nearly so; buds often present in blade sinuses of actively growing fronds; fertile fronds ca. 15–50 ⫻ 8–35 cm, pinnae 4–9 pairs, ascending, proximal pairs often opposite or subopposite; sporangia with annuli absent or vestigial, the indurate cells 0–12(–40); spores 32 per sporangium; 2n⫽78 (Salv).

Distribution. Floating in marshes, ponds, or in slow-moving rivers or streams; 0–50 m. Se USA; Mexico; Guat, Hond (Saunders 507, NY), Salv, CR, Pan; Cuba, Hisp; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol, Parag, n Arg. Also cited from Nicaragua by Moran (in Davidse et al., 1995: 121), but specimen (Grijalva 1851, MO) not found. Selected Specimens Examined. Chis (Breedlove 56074, CAS, ENCB, LL, MEXU, NY). Gro (Lozada P. 926, FCME; Vela´zquez 740, ENCB). Nay (Te´llez 10363, MEXU). Tab (Cowan 2804, CAS, CHAPA, ENCB, MEXU, NY, 2882, CHAPA, ENCB, NY; Haynes 5500, NY; Zamudio 1057, MEXU). Ver (Lot 1625, ENCB, IEB, MEXU; Palacios-Rios 3232, XAL).

Lloyd (1974) recognized four species in the genus, three of which were cited from both Guatemala and the United States. Although he cited only one specimen of Ceratopteris from Mexico (Liebmann s.n., US!) and this was identified as a hybrid between C. pteridoides and C. richardii Brongn., we refer all but one recent Mexican collection to C. pteridoides. Ceratopteris pteridoides has 32 spores per sporangium, stipe width 4–17 mm, sterile blades often simple (palmately or pinnately lobed), sometimes pinnate, and proximal pinnae or segments of sterile blades opposite. Ceratopteris richardii Brongn. (Dict. Class. Hist. Nat. 3: 351. 1823) reputedly has 16 spores per sporangium, stipe width 0.8–6.5(–12) mm, sterile blades pinnate to bipinnate, and proximal pinnae of sterile blades inserted mostly alternately. Tryon and Tryon (1982) treated C. richardii as a synonym of C. thalictroides. Hickok and Klekowski (1974) and Hickok (1977) found that synthesized hybrids between C.

176

CHEILANTHES

pteridoides and C. richardii, both diploids, resulted in 40% viable spores, suggesting that those two species are not completely reproductively isolated. The affinities of C. richardii are still uncertain; Lloyd (1974) thought it intermediate morphologically between C. thalictroides and C. cornuta (P. Beauv.) Lepr., an African species. Should C. richardii prove to be synonymous with C. pteridoides, then the former name has priority.

2. CERATOPTERIS THALICTROIDES Ceratopteris thalictroides (L.) Brongn., Bull. Sci. Soc. Philom. Paris, se´r. 3, 8: 186. 1821. Figs. 68E–G. Acrostichum thalictroides L., Sp. Pl. 2: 1070. 1753. Lectotype (designated by Pichi Sermolli, Webbia 12: 650. 1957). Sri Lanka. Hermann s.n. (BM-Herb. Hermann 3: 42, specimen 377).

Plants generally on mud, sometimes in shallow water; stipes 0.7–8 mm wide, about the same width throughout their length; sterile fronds ca. 6–60 ⫻ 5–20 cm; sterile blades (1–)2–3-pinnate, the proximal pinnae alternate; buds absent or usually dormant on actively growing blades, generally developed only on senescent blades; fertile fronds usually ca. 15–60 ⫻ 10–35 cm, pinnae 4–6 pairs, ascending, proximal pairs subopposite; sporangia with annuli of 15–70 clearly indurate cells; spores 32 per sporangium; 2n⫽156 (Hawaii, Japan).

Distribution. In slow-moving rivers or streams; 10 m. USA (Calif, Fla, La, Tex); Mexico; Guat, Salv, Nic, CR, Pan; Ven, Sur, Ec, Braz; Africa, Asia, Australia, islands of the Pacific. Specimen Examined. Gro (Fonseca 1643, FCME). Unverified, Doubtful, or Mistaken Reports. Oax (Pe´rez-Garcı´a 1033, MEXU, cited by Tejero-Dı´ez & Mickel, 2004, but not verified). Tab (reported by Magan˜a, 1992, probably based on a misidentification of C. pteridoides).

Citation of this species from Guerrero by Lorea-Herna´ndez & Vela´zquez M. (1998) was based on a specimen they identified as C. pteridoides; they did not cite the Fonseca collection cited above, which was collected in 1989. Lellinger (1989) and Moran (in Davidse et al., 1995) cited Mexico (Tabasco) in the range of C. thalictroides, but no specimens have been seen from there; the citation by Moran was based on Cowan (1983), but these specimens are all C. pteridoides. Another possible record of C. thal-

ictroides from Mexico is a specimen from Tamaulipas, “presa Vicente Guerrero, en El Campo recreativo Morris, 2 km de V. casas, Mpio., Villa de Casas,” 250 m, Martı´nez 929 (MO), but the specimen is poor, sterile, and of uncertain identity. Stolze’s (1981: 125) illustration of “C. thalictroides” is similar to the Chiapas collection identified here as C. pteridoides, and it seems possible the specimen on which the illustration was based is misidentified. Sterile fronds of C. thalictroides are generally more finely divided with much narrower segments than those of C. pteridoides, and the stipes are not inflated.

25. CHEILANTHES Cheilanthes Sw., Syn. Fil. 126. 1806, nom. cons. Type: Cheilanthes micropteris Sw. Terrestrial or epipetric; rhizomes compact to long-creeping, occasionally ascending, scaly; roots wiry; fronds monomorphic, small to medium-sized, 1- to 5-times pinnate, hairy, scaly, farinose, or glabrous abaxially; veins free; sori near the blade margins, usually with the margins inrolled and differentiated; spores tetrahedral-globose; x⫽29, 30. Cheilanthes has approximately 150 species, mostly of warm, dry, rocky regions, with nearly half the species in dry parts of Mexico and the southwestern United States. The apogamous condition in many species is generally suggested by the presence of 32 spores per sporangium rather than the customary 64, and some species have more than one cytotype. Blade indument is highly diverse in the genus. Scales show every transition to hairs in the C. lendigera complex. A few species have hairs with a glandular tip. Some species have hairs ascending to spreading directly from their bases, but many species have a unique type of hair with the basal 2–4 cells very small, generally starting basiscopically and then having larger cells that grow acroscopically (Fig. 87N). Cheilanthes has undergone major changes in circumscription recently. For many years Notholaena was separated from Cheilanthes on the basis of the sori running along the veins near their tips and the blade margins not differentiated, but these characters were shown to be artificial and unreliable as generic characters. Mickel (1979, 1992) and Mickel and Beitel (1988) combined the two under Cheilanthes. Cheilanthes is sometimes confused with Pellaea, and several species traditionally maintained in the latter genus are here treated in Cheilanthes and Argyrochosma. There are several natural species groupings still within Cheilanthes, but at what level they should be recognized is still uncertain. Molecular studies by Gastony and Rollo (1998) indicate that Cheilanthes, as defined here, is polyphyletic and might better be divided into several genera. Some species should perhaps be treated in other existing genera, e.g., C. aurea, C. aurantiaca, and C. leucopoda may be better placed in Notholaena, C. farinosa in the principally Asian genus Aleuritopteris. In other cases, genera recognized as distinct in this treatment are perhaps better in Cheilanthes, e.g., species treated herein as Mildella.

CHEILANTHES References Gastony, G. J. & D. R. Rollo. 1998. Cheilanthoid ferns (Pteridaceae: Cheilanthoideae) in the southwestern United States and adjacent Mexico–a molecular phylogenetic reassessment of generic lines. Aliso 17: 131–144; Gastony, G. J. & M. D. Windham. 1989. Species concepts in pteridophytes: the treatment and definition of agamosporous species. Amer. Fern J. 79: 65–77; Mickel, J. T. 1979. The fern genus Cheilanthes in continental United States. Phytologia 41: 431–437; Mickel, J. T. 1987. A new fern from western Mexico and its bearing on the

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taxonomy of the cheilanthoid ferns. Amer. Fern J. 77: 109–114; Reeves, T. 1979. A monograph of the fern genus Cheilanthes subgenus Physapteris (Adiantaceae). Unpublished Ph.D. dissertation, Arizona State University; Saiki, Y. 1984a. Note on ferns: 2. Asiatic species of the Aleuritopteris farinosa complex. J. Phytogeogr. Taxon. 32: 1–13; Saiki, Y. 1984b. Note on ferns: 3. African and neotropical species of the Aleuritopteris farinosa group. J. Phytogeogr. Taxon. 32: 81–90; Saiki, Y. 1984c. Note on ferns: 4. Classification of the Aleuritopteris farinosa group. J. Phytogeogr. Taxon. 32: 91–98; Tryon, R. M. 1956. A revision of the American species of Notholaena. Contr. Gray Herb. 179: 1–106.

Key to the Mexican Species of Cheilanthes 1. Blades farinaceous abaxially. 2. Farina white; blades bipinnate-pinnatifid, glabrous except for farina; rhizome scales concolorous, dark reddish brown, or bicolorous with black center and brown margins. ................................................................................................................................ 23. C. farinosa. 2. Farina yellow-orange; blades pinnate-pinnatifid to tripinnate-pinnatifid, with hairs in addition to farina; rhizome scales concolorous, tan, or bicolorous with age in C. aurantiaca. 3. Blades pinnate-pinnatifid to bipinnate, lanceolate to elliptic; hairs on blade margins, adaxial laminar surfaces, and adaxial rachises (absent on abaxial surfaces); basiscopic segments not exaggerated; rhizome scales bicolorous with age. .............. 7. C. aurantiaca. 3. Blades 2–3-pinnate-pinnatifid, broadly deltate to pentagonal, basal basiscopic segments exaggerated; abaxial blade surfaces with long white hairs along axes, major veins, blade margins, and adaxial surfaces; rhizome scales concolorous, tan. ............ 8. C. aurea. 1. Blades lacking farina abaxially. 4. Stipes stramineous, green, or gray to dull gray-brown. 5. Blades broadly pentagonal; fronds densely patent-hairy, hairs patent; rhizome scales orange-tan. ........................................ 32. C. leucopoda. 5. Blades narrowly oblong or lanceolate, or if deltate, then glabrous adaxially and sparsely and laxly hairy abaxially; rhizome scales concolorous light brown or bicolorous. 6. Blades tripinnate-pinnatifid to quadripinnate-pinnatifid, glabrous, broadly deltate; stipes stramineous; rhizome scales concolorous, tan, entire; margins of indusia with whitened cells; spores gray-brown; base of root enlarged as storage organ (2–4 mm long, 1–2 mm wide). ..................................................................................................................................... 9. C. bolborrhiza. 6. Blades pinnate to tripinnate-pinnatifid, linear-lanceolate to ovate-lanceolate; stipes gray to grayish brown, green, or stramineous; rhizome scales bicolorous with thick reddish brown to black center and pale erose-denticulate margin (breaking into 1-mm hairs); adaxial laminar surface with hairs (0.1 mm long) especially near the margin, hairs (0.5–1 mm long) or narrow scales on pinnae undersurface; modified false indusia entire, extending the length of the pinnules; spores tan; base of roots unmodified. 7. Blades pinnate to bipinnate-pinnatifid, narrowly oblong; stipes dull gray to gray-brown. ......................................... 35. C. lozanoi. 7. Blades 2–3-pinnate-pinnatifid, broadly deltate to ovate-lanceolate; stipes green to stramineous. ............................. 52. C. skinneri. 4. Stipes castaneous (usually lustrous), atropurpureous or black. 8. Blades pinnate to pinnate-pinnatifid. 9. Pinnae abaxially with erose scales or serrulate scales and hairs; stipes scaly; rhizome scales 6–12 ⫻ 0.1–0.5 mm, with occasional teeth; laminar margins strongly differentiated into 0.5 mm wide, thin, false indusia; pinnae nonarticulate. ...................................................................................................................................................................................... 11. C. brachypus. 9. Pinnae abaxially with long, matted hairs only; stipes with 1–2 mm long white hairs only; rhizome scales 3 ⫻ 0.5 mm; laminar margins not differentiated; pinnae articulate. 10. Abaxial blade hairs tangled; stipe hairs straight, slightly curved downward at base then up, slender, not jointed; rhizome scales strongly bicolorous, lustrous; plants to 75 cm; widespread. ............................................................. 10. C. bonariensis. 10. Abaxial blade hairs straight; stipe and rachis hairs jointed, straight; rhizome scales concolorous pale brown, dull; plants small, 10–15 cm; Sonora. ............................................................................................................................... 59. C. yatskievychiana. 8. Blades bipinnate to quadripinnate-pinnatifid. 11. Blade axes glabrous, lacking multicellular hairs and/or scales; vein endings pronounced and conspicuous on adaxial blade surfaces. 12. Stipes terete, or grooved only distally; rachises and rachillae grooved with pale shoulders, at least distally; blades pentagonal. 13. Segments mostly 3–6 mm long, terminal ones 5–8(–12) mm. 14. Indusia long-decurrent, broad, covering the abaxial segment surfaces. ........................................................ 21. C. decurrens. 14. Indusia not decurrent or only slightly so, narrower, abaxial segment surfaces visible. 15. Rhizome scales concolorous black; Tam, Gto, NL, Ver, Hgo, SLP, Qro. ............................................. 50. C. purpusii. 15. Rhizome scales bicolorous black with brown margins; Gro, Oax, Chis. .......................................... 16. C. complanata. 13. Segments mostly 7–20 mm long, terminal ones 12–30 mm. 16. Segments mostly 10–15 mm long, terminal ones 20–25 mm; blades quadripinnate; indusia decurrent; NL. .............................................................................................................................................................................. 5. C. apiacea. 16. Segments mostly 7–20 mm long, terminal ones 12–20(–30) mm; blades 2–3-pinnate, or 4-pinnate, not NL.

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CHEILANTHES 17. Indusia not decurrent; pinnae 4–6 pairs; segments mostly 8–20 mm long (terminal ones to 30 mm); blades 2–3-pinnate; veins ending flush with non-farinose surfaces; NL. ........................................ 27. C. hintoniorum. 17. Indusia decurrent; pinnae 8–10 pairs; segments mostly 7–12 mm long (terminal ones to 20 mm); blades 3–4-pinnate; veins ending in pits, the pits often farinose; Jal, Gro, Tam, Oax. .................... 47. C. pellaeopsis. 12. Stipes grooved; rachises grooved, but shoulders not distinctly paler; blades lanceolate, ovate, or deltate. 18. Spores black; rhizome scales dark castaneous to black, concolorous; indusia 0.1–0.8 mm wide with suberose to erose, non-decurrent margins, indusial surfaces white-papillate or not; veins prominent abaxially. 19. Blades 2–3-pinnate, lanceolate or ovate to deltate (length:width ratio 1.3–4:1), 3–9 cm wide; fronds 12–32 cm long. 20. Blades deltate (length:width ratio 1.3–2:1); pinnae (5–)7–10 pairs, basal pinnae largest; terminal segments of pinnae linear, 1.5–4 cm ⫻ 1–1.5(–2) mm (length:width ratio 15–40:1); indusia strongly erose, 0.1–0.3 mm. ..................................................................................................................................... 4. C. angustifolia. 20. Blades lanceolate (length:width ratio 2–4:1); pinnae 10–20 pairs, basal pinnae slightly reduced; terminal segments of pinnae ovate, 0.5–1.5 cm ⫻ 2–4 mm (length:width ratio 2.5–7:1); indusia suberose, 0.5–0.8 mm wide. ........................................................................................................................... 19. C. cuneata. 19. Blades tripinnate-pinnatifid to quadripinnate, strongly deltate (length:width ratio 0.8–1:1), 12–20 cm wide; fronds 30–52 cm long. 21. Terminal pinna segments conform, lanceolate, 0.7–1.5 cm ⫻ 1.5–2.5 mm (length:width ratio 4.5–6:1); basal pinnae 15–20 cm long; basal lobe of basal basiscopic pinnule of basal pinnae with 2–3 lobes. .......................................................................................................................................................... 20. C. decomposita. 21. Terminal pinna segments pinnatifid, ovate, 0.4–0.8 cm ⫻ 2–3 mm (length:width ratio 1.8–3:1); basal pinnae 10–14 cm long; basal lobe of basal basiscopic pinnule of basal pinnae with 4–7 lobes. .......................................................................................................................................................... 13. C. chaerophylla. 18. Spores brown or tan; rhizome scales castaneous concolorous (in C. membranacea) or bicolorous with black center and narrow light brown margins; indusia 0.3–1 mm wide with entire to long-fimbriate margins, longdecurrent or not, indusial surfaces often white-pebbly to long-papillate (C. pyramidalis); veins not prominent abaxially. 22. Stipes 1.5–3.5 mm diam., tightly clustered; rhizome scales castaneous concolorous; rachises flexuous; indusia long-decurrent. ................................................................................................................................. 39. C. membranacea. 22. Stipes 1–1.5 mm diam., approximate, not tightly clustered; rhizome scales bicolorous, black-centered with brown to tan margins; rachises straight; indusia decurrent or not. 23. Indusial margins papillate, not at all decurrent. ........................................................................................ 6. C. arizonica. 23. Indusial margins short- to long-fimbriate, generally short- to long-decurrent. 24. Recurved adaxial blade and indusial surfaces with white papillae (spicules) 0.3 mm long; indusia slightly decurrent to decurrent to segment base (rarely to next segment), fimbriate; 850– 2750 m. ............................................................................................................................................... 51. C. pyramidalis. 24. Indusia long decurrent onto subtending axis to the next segment; indusial surfaces glabrous; (2200–)2700–3500 m. 25. Recurved adaxial blade surfaces glabrous, spicules absent from segments. ........................ 36. C. marginata. 25. Recurved adaxial blade surfaces spiculate. .................................................................................53. C. spiculata. 11. Blade axes with scales and/or hairs; vein endings inconspicuous adaxially. 26. Rachises grooved. 27. Rhizomes compact; fronds approximate, mostly 20–30 cm long. 28. Rachises and stipes densely clothed with a mixture of hairs and linear scales; articulation lines absent from stipes; Oax. .....................................................................................................................................................41. C. mickelii. 28. Rachises and stipes glabrous except for a few scales at bases of pinnae; older fronds with articulation lines on stipes proximally; widespread in Mexico. .................................................................................................. 3. C. allosuroides. 27. Rhizomes creeping; fronds 1–3 mm distant, mostly 10–15 cm long (some longer). 29. Fronds nearly glabrous, with scattered to sparse scales and hairs on abaxial rachises and stipes. .............. 58. C. wrightii. 29. Fronds with conspicuous scales on axes. 30. Stipes and rachises with indument primarily of scales; stipes reddish brown. ...................................... 49. C. pringlei. 30. Stipe and rachises with indument predominantly hairs, scales very few and narrow; stipes purplish brown. ........................................................................................................................................................ 48. C. peninsularis. 26. Rachises and stipes terete. 31. Adaxial laminar surfaces with conspicuous pustulate spiculate hairs. .................................................................. 28. C. horridula. 31. Adaxial laminar surfaces lacking pustulate spiculate hairs, glabrous or with fine hairs. 32. Indusia fimbriate. ................................................................................................................................................. 26. C. fimbriata. 32. Indusia entire. 33. Costae with scales more than 5 cells wide, lanceolate to deltate. 34. Blade scales ciliate. 35. Blade scales ciliate only in basal half of scale.

CHEILANTHES

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36. Abaxial blade scales 2 mm or more long, whitish with castaneous base, exceeding the segment margins, the visible part of the abaxial scales entire; the laminar scales ciliate only at their bases. ................................................................................................................... 17. C. covillei. 36. Abaxial blade scales ca. 1 mm long, dark reddish brown, covering the surfaces but not exceeding the margins, largely the long-attenuate scale tips and lateral cilia; the laminar scales ciliate their entire length. ....................................................................................... 15. C. clevelandii. 35. Blade scales ciliate throughout the scale length. 37. Pinnae densely pubescent adaxially; abaxial costal scales with fine, curly cilia forming entangled masses that are visible adaxially; ultimate segments minute, 0.5–1 mm. .......................................................................................................................... 33. C. lindheimeri. 37. Pinnae glabrous or sparsely pubescent adaxially; abaxial costal scales with coarse cilia that are not strongly entangled; ultimate segments larger, at least some 1–3 mm. 38. Pinnae glabrous adaxially; costal scales often ciliate only in proximal 1⁄2; rhizome scales usually brown, concolorous brown to dark brown to weakly bicolorous, loosely appressed and deciduous on older portions of stem. ................................. 57. C. wootonii. 38. Pinnae sparsely pubescent adaxially; costal scales usually ciliate their entire length; rhizome scales dark brown to blackish, bicolorous, usually strongly appressed and persistent. .................................................................................................................... 60. C. yavapensis. 34. Blade scales entire to erose-denticulate. 39. Segments glabrous adaxially; rhizomes short- to long-creeping. ............................................. 25. C. fendleri. 39. Segments densely to sparsely pubescent adaxially; rhizomes short-creeping. 40. Rachises with linear scales, pinna rachis scales lanceolate. ................................................ 22. C. eatonii. 40. Rachises with broad, ovate-lanceolate scales. 41. Scales not far exceeding the segment margins, not visible as viewed from above; adaxial surfaces glabrous or with scattered hairs. ................................................. 43. C. myriophylla. 41. Scales far exceeding the segment margins, conspicuous as viewed from above; adaxial surfaces moderately to densely hairy. ................................................................. 55. C. villosa 33. Costae with hairs only, or with linear scales 2–3(–5) cells wide. 42. Stipes and both blade surfaces with short, rigid, glandular hairs. 43. Blades broadly deltate to pentagonal, pedate; rhizome scales reddish black. ..................... 30. C. kaulfussii. 43. Blades narrowly deltate to narrowly oblong; rhizome scales reddish orange or orange-tan. 44. Blades not conspicuously pubescent, but mostly the stipes, laminar glands mostly sessile; rhizome scales reddish orange, flexuous. ................................................................. 56. C. viscida. 44. Blades conspicuously pubescent; rhizome scales tan to orange-tan, straight. 45. Blades narrowly oblong; fronds 28–40 cm; laminar hairs mostly ca. 2 mm; Tam. ............................................................................................................................. 38. C. maxoniana. 45. Blades narrowly deltate to lanceolate; fronds 9–30 cm; laminar hairs 0.2–2 mm; NL to Jal and Oax. ....................................................................................................... 34. C. longipila. 42. Stipes and blade surfaces without short, rigid glandular hairs. 46. False indusia strongly differentiated, forming a pouch on the bead-like segment, segment margins often puckered. 47. Indusia to ca. 0.1 mm wide. ..............................................................................................40. C. mexicana. 47. Indusia 0.2–0.5 mm wide. 48. Stipes and rachises with linear scales mixed with hairs; indusia glabrous at indusiallaminar interface; indusia 0.3–0.5 mm wide. .................................................... 37. C. marsupianthes. 48. Stipes and rachises with hairs only; with hairs at indusial-laminar interface; indusia 0.2–0.3 mm wide. ......................................................................................................... 31. C. lendigera. 46. Indusia weakly differentiated, not forming a pouch. 49. Blades only with straight-based hairs. 50. Segments nearly round, bead-like. ..................................................................................... 24. C. feei. 50. Segments oblong, elongate, or if ovate-roundish, 2–6 mm diam. and mostly adnate. 51. Segments long-villous to nearly glabrous abaxially, segment surfaces not concealed. ............................................................................................................. 12. C. brandegeei. 51. Segments densely hairy abaxially, concealing the blade surfaces. 52. Hairs matted on segments, very fine; rhizome scales black, linearlanceolate; blades bipinnate-pinnatifid to tripinnate; sporangia evident. ....................................................................................................................... 44. C. newberryi.

52. Hairs spreading, fluffy, the segments appearing like balls of cotton; rhizome scales orange-tan, each occasionally with a thin black central stripe; blades tripinnate to tripinnate-pinnatifid; sporangia hidden. ......... 46. C. parryi.

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CHEILANTHES 49. Rachis pubescence consisting of narrow scales and hairs mixed, the hairs reversing direction near their base. 53. Blades 3–4-pinnate; segments oblong. 54. Rachises and costae with white, appressed to ascending, linear scales. ..... 29. C. jamaicensis. 54. Rachises and costae with orange-tan spreading scales and hairs. 55. Terminal segments distinctly oblong, usually 3 mm long; abaxial segment surfaces not completely covered with hairs, lamina at least slightly visible. ....................................................................................................... 14. C. chipinquensis. 55. Terminal segments more or less round, 1–2 mm diam.; abaxial segment surfaces completely covered with hairs. ................................................... 54. C. tomentosa. 53. Blades 2–3-pinnate; segments toothed or auriculate, not oblong-entire. 56. Rachises lacking narrow scales, with hairs only. 57. Blades deltate, tripinnate-pinnatifid to quadripinnate. ............................... 1. C. aemula. 57. Blades lanceolate, bipinnate to bipinnate-pinnatifid. 58. Rhizomes short- to long-creeping; blade segments mostly rounded, chartaceous; blades mostly 2–3- pinnate. ....................................... 42. C. microphylla. 58. Rhizomes compact to short-creeping; blade segments mostly acute to acuminate, coriaceous; blades mostly bipinnate to bipinnatepinnatifid. ............................................................................................ 2. C. alabamensis. 56. Rachises with linear scales mixed with hairs. 59. Blades bipinnate, pinnules unlobed, or with only a single acroscopic lobe at base; adaxial pinna surfaces bluish to whitish green; rhizome scales entire, occasionally with rhizome hairs showing among the scales; laminae subcoriaceous. ........................................................................................ 45. C. notholaenoides. 59. Blades bipinnate-pinnatifid to tripinnate, pinnules with acroscopic and basiscopic lobes; adaxial pinna surfaces green; rhizome scales with longciliate margins; laminae chartaceous. ......................................................... 18. C. cucullans.

1. CHEILANTHES AEMULA Cheilanthes aemula Maxon, Contr. U.S. Natl. Herb. 10: 495. 1908. Type. Mexico. Tamaulipas: Victoria, 1907, Palmer 187 (US!; isotypes GH!, K!, MICH!, NY!). Figs. 72J, K. Rhizomes short-creeping, horizontal, 1–2 mm diam.; rhizome scales linear, 2–3 mm long, concolorous tan to orange-brown, often darker at bases, margins entire; fronds to 55 cm tall, approximate; stipes 1⁄3–1⁄2 the frond length, atropurpureous to black, terete, with lax hairs 1 mm long and hairs less than 0.1 mm long on adaxial surface of distal stipes and rachises to glabrescent; blades deltate, tripinnate-pinnatifid to quadripinnate, the pinnae slightly more developed basiscopically, the basal pair basiscopically exaggerated, texture thin; adaxial blade surfaces glabrous or with scattered, 0.5–0.8 mm long hairs, costae shorthairy in grooves; abaxial blade surfaces with scattered 0.5–0.8 mm long hairs on laminae; sporangia on vein ends, margins curved, modified into 0.3 mm wide false indusia; sporangia 64-spored; spores tan; 2n⫽58 (NL, USA).

Distribution. Dry, rocky slopes, all limestone; 130–1800 m. USA (Tex); Mexico. Selected Specimens Examined. Chih (Knobloch 449, MSC). Chis (Breedlove 21902, DS). Coah (Knobloch 2428, MSC). Hgo (Matuda 23567, MEXU). Mich (Escobedo 2447, NY). NL (Pringle 1988, NY). Oax (Hellwig 364, NY). Qro (Rzedowski 48083, IEB, MEXU). SLP (Fryxell & Anderson 3554, NY). Tam (Bartlett 10633, MICH). Ver (Purpus 2173, DS).

Cheilanthes aemula is distinguished by the combination of deltate, tripinnate-pinnatifid to quadripinnate fronds, pinnules with lobes on both acroscopic and basiscopic sides, stipes and rachises terete with dense adaxial catenate hairs and sparse, longer, abaxial hairs, and lower leaf surfaces with scattered 0.3– 0.5 mm long whitish hairs. The closely related C. microphylla and C. alabamensis have fronds that are bipinnate-pinnatifid to

CHEILANTHES ALLOSUROIDES

tripinnate and lanceolate, and both C. cucullans and C. notholaenoides have a dense covering of 1–2 mm long golden hairs and linear scales on the rachises, and lanceolate fronds that are bipinnate to almost tripinnate.

2. CHEILANTHES ALABAMENSIS Cheilanthes alabamensis (Buckley) Kunze, Linnaea 20: 4. 1847. Figs. 78J–L. Pteris alabamensis Buckley, Amer. J. Sci. Arts 45: 177. 1843. Pellaea alabamensis (Buckley) Baker in Hooker & Baker, Syn. Fil. 148. 1867. Allosorus alabamensis (Buckley) Kuntze, Revis. Gen. Pl. 2: 806. 1891. Type. U.S.A. Alabama: At foot of Muscle Shoals, banks of Tennessee River, on limestone rocks, Buckley s.n. (PH; possible type, NY– “limestone rocks, Tennessee river, Florence, Ala.”).

Rhizomes compact to short-creeping, horizontal, 1–2 mm diam.; rhizome scales linear, concolorous orange-brown, slender, 2–3.5 mm long, entire; fronds 13–30 cm long, approximate; stipes black, lustrous, terete, glabrate, short-pubescent adaxially on distal stipes and rachises with scattered, twisted, catenate hairs 0.5– 0.8 mm; blades lanceolate, bipinnate to 2-pinnate-pinnatifid, coriaceous; segments lanceolate, auriculate, acute to acuminate at apices; adaxial surfaces glabrous except for rare 0.1(–0.3) mm long white hairs; abaxial surfaces glabrous or with rare white hairs 0.1–0.3 mm long; sori at vein ends, laminar margins curved, only slightly modified into 0.3 mm wide false indusia; sporangia 32spored; spores tan; 2n⫽58 (NL).

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3. CHEILANTHES ALLOSUROIDES Cheilanthes allosuroides Mett., Abh. Senckenberg. Naturf. Ges. 3: 78. 1859. Figs. 72A, B. Pellaea allosuroides (Mett.) Hieron., Hedwigia 62: 18. 1920. Type. Mexico. Schmitz s.n. (B). Pellaea arsenei Christ, Notul. Syst. (Paris) 1: 233. 1910. Type. Mexico. Michoaca´n: Morelia, Arse`ne 2496 (P!; isotypes B!, US!). Pellaea pallida Hook. & Baker, Syn. Fil., ed. 2, 478. 1874. Type. Mexico. Glennie s.n. (K!).

Rhizomes compact, horizontal, 2–3 mm diam.; rhizome scales linear, concolorous, orange-brown, 3–4 mm long, entire; fronds to 30 cm tall, clumped; stipes ca. 1⁄3 the frond length, grooved, dark castaneous to purplish brown, lustrous, with sparse, 2 mm long, appressed hairs, older fronds with articulation lines proximally, abscised old fronds leaving stipe bases 1–1.5 cm long; rachises glabrous except for a few scattered or sparse linear tan scales at pinna bases; blades lanceolate, 2–3-pinnate, texture firm; pinnae 8–13 pairs, segments mostly elliptical to lanceolate, often auriculate; adaxial surfaces glabrous and minutely papillate; abaxial surfaces glabrous except for a few long hairs on costae; sori elongate along vein ends, segment margins curved, firm, somewhat modified, thin only at the minutely erose edges; spores tan.

Distribution. Dry, rocky slopes, on basaltic or other rocks (all acidic, apparently); 1000–2400 m. Mexico.

Distribution. Limestone outcrops; 450–2400 m. Se USA; Mexico. Selected Specimens Examined. Chih (Pringle 449, NY). Coah (Wynd & Mueller 377, NY). Gto (Rzedowski 40582, IEB). NL (Pringle 1989, NY). Oax (Camp 2232, NY). Pue (Purpus 4097, UC). Qro (Zamudio 6101, IEB). SLP (Palmer 268, NY). Son (Van Devender s.n., 6 Feb 1978, UC). Tam (von Rozynski 285, NY p.p.).

Cheilanthes alabamensis can be distinguished from the closely related C. microphylla (q.v.) by its lanceolate segments glabrous on both sides or with only rare 0.1 mm white hairs abaxially, vs. oblong segments with scattered hairs 0.1–0.5 mm on both sides in C. microphylla.

Selected Specimens Examined. Ags (Reeder et al. 3339, ARIZ). Chih (Pringle 1443, NY). DF (Pringle 11271, NY). Dgo (Shreeve 9187, ARIZ). Gto (Sohs 337, US). Hgo (Orcutt 6237, MEXU). Jal (Pringle 5399, NY). Me´x (Rose 11039, NY). Mich (Arse`ne s.n., 9 Aug 1909, MICH). Nay (Breedlove 45565, CAS). Oax (Mickel 4952, NY). Pue (Purpus 3148, NY). Qro (Rose 11179, NY). SLP (Parry & Palmer 979, NY). Sin (Breedlove 43871, CAS, MEXU). Son (Boutin & Kimnach 3677, NY). Ver (Botteri 38, BM). Zac (Reveal 2675, US).

The poorly differentiated false indusia, sori elongate along the veins, and scattered linear scales on the major axes resemble those of C. notholaenoides and C. cucullans, but the glabrous blade surfaces and grooved rachises with only sparse scales distinguish C. allosuroides. It is related to C. wrightii, but the sori are protected by long continuous indusia rather than only interrupted lobes in C. wrightii. Cheilanthes allosuroides is generally

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taller than C. wrightii (15–30 cm vs. 8–22 cm) and has scales limited to pinna bases (vs. scattered scales on stipes).

4. CHEILANTHES ANGUSTIFOLIA Cheilanthes angustifolia Kunth, Nov. Gen. Sp. 1: 21. 1815. Figs. 71G, H. Allosorus angustifolius (Kunth) C. Presl, Suppl. Tent. Pterid. 152. 1836. Onychium angustifolium (Kunth) Kunze, Farrnkra¨uter 2: 11. 1848. Pellaea angustifolia (Kunth) Baker in Hooker & Baker, Syn. Fil. 150. 1867. Type. Mexico. [Michoaca´n:] “Crescit locis subapricis in monte ignivomo Jorullo, alt. 630 hexapl.” Humboldt & Bonpland s.n. (B-Willd. 20116!; P-Humb.!). Allosorus angustifolius (Kunth) C. Presl var. minor Liebm., Mexic. Bregn. 219 (reprint 67). 1849. Lectotype (chosen by Smith, 1981: 71). Mexico. Oaxaca: Yavesia, Liebmann s.n. [Pl. Mex. 2349, Fl. Mex. 231] (C!). Cheilanthes venusta Fe´e var. aurata Fe´e, Me´m. Foug. 9: 12. 1857, nom. nud., based on Mexico. [Oaxaca:] San Pedro de Nolasco, Galeotti 6560 (BR!–6 sheets).

Rhizomes compact, horizontal, 2–3 mm diam.; rhizome scales linear-lanceolate, concolorous, castaneous to black, lustrous, 2.5– 3 mm long, entire; fronds 12–32 cm long, clumped; stipes 1⁄2–2⁄3 the frond length, castaneous to blackish, lustrous, grooved, glabrous or with sparse 0.3–0.5 mm hairs and linear, brown scales; blades deltate, 2–3-pinnate, basal pinnae usually largest, 5–12 ⫻ 3.5–9 cm (length:width ratio 1.3–2:1), glabrous on both sides; pinnae (5–)7–10 pairs (apical 4–5 pinna pairs entire, median 2– 6 pinna pairs pinnate, basal 0–1 pair bipinnate), terminal segments linear, 1.5–4 cm ⫻ 1–1.5(–2) mm (length:width ratio 15– 40:1); basal basiscopic pinnule 1.8–3.5 cm long, 0–3-lobed, terminal segment 1–1.5(–2) mm wide; adaxial and abaxial surfaces glabrous, the veins prominent abaxially; laminar margin curved, strongly modified in 0.1–0.3 mm wide false indusia, deeply erose, nearly discontinuous, surface smooth or slightly white-papillate; spores black.

Distribution. Slopes in dry to moist oak or pine forests, on gravelly bluffs and limestone rocks; 100–2450 m. Mexico; Guat, Hond, Salv, Nic, CR, Pan. Selected Specimens Examined. Chih (Tenorio C. 2031, MEXU). Chis (Ventura & Lo´pez 4023, NY). Col (Sanders 8499, UC). Dgo (Maysilles 8026, MICH). Gro (Anderson & Laskowski 4394, NY). Gto (Rzedowski 51267, IEB). Jal (McVaugh 17184, NY). Me´x (Manning & Man-

ning 531117, MEXU). Mich (Hinton 12190, NY). Mor (Gold 208, MEXU). Nay (Alava & Cook 1574, UC). Oax (Mickel 1321, NY). Pue (Guı´zar 1429, MEXU). Qro (Zamudio & Carranza 7133, IEB). Sin (Anderson 12567, NY). Son (Van Devender 93-1326, ARIZ). Tam (Diggs & Nee 2378, XAL). Ver (Va´zquez 2151, XAL). Unverified, Doubtful, or Mistaken Reports. Ags (De la Cerda & Garcı´a 971, HUAA, cited by Siqueiros-Delgado & Gonza´lez-Adame, 2004, but not verified).

The C. angustifolia complex, including also C. cuneata, C. decomposita, and C. chaerophylla, is in need of careful biosystematic study. The large, round, black spores found in all its members seem to indicate apogamy, and intermediate specimens exist between the species as characterized here. Our concepts of the four species have been carefully matched with the illustrations in Martens & Galeotti (1842) (for C. decomposita and C. chaerophylla), microfiche of the Willdenow herbarium (for C. angustifolia), and photo of type material at Berlin (for C. cuneata), as well as examination of the type specimens. Cheilanthes angustifolia and C. cuneata, as indicated in the key, have blades that are less divided (2–3-pinnate vs. 3–4-pinnate), more narrowly deltate (vs. broadly deltate), and fronds that are shorter than those of C. decomposita and C. chaerophylla. Cheilanthes angustifolia has long, narrow terminal pinna segments, a deltate blade, and deeply erose, narrow indusia (see key), whereas C. cuneata has wider, ovate, terminal pinna segments, ovate blades, more pinna pairs, and wider, suberose indusia. These two species seem to intergrade and may prove to be conspecific. Cheilanthes decomposita has long, narrow, conform terminal segments, and larger basal pinnae, whereas C. chaerophylla has shorter, ovate, pinnatifid terminal segments, and smaller basal pinnae. The status of the intermediate specimens is unclear, and the presence of globose spores and presumed apogamy in the whole group precludes using spore abortion as a test for hybridity. The intermediates between C. angustifolia and C. cuneata have fronds that are tripinnate (less divided and smaller than C. chaerophylla and C. decomposita), parallel-sided for the basal half (not ovate like C. cuneata and not deltate like C. angustifolia), conform terminal pinna segments that are 0.8–2.2 cm long, 1.5–3 mm wide (length:width ratio 5–14:1), and ascending pinnae. These intermediates (all at NY) are known from the states of Jalisco, Durango, Me´xico, Oaxaca, and Chiapas. The intermediates between C. cuneata and C. decomposita have blades that are tripinnate to tripinnate-pinnatifid, narrowly deltate, and have broad, conform terminal pinna segments. These are known from the states of Sinaloa, Nayarit, Veracruz, and Oaxaca. Intermediate specimens considered as hybrids by Mickel and Beitel (1988) are here merely designated as intermediates. There seem to be no sharp boundaries between the members of the complex.

5. CHEILANTHES APIACEA Cheilanthes apiacea Mickel, sp. nov. Type. Mexico. Nuevo Leo´n: Mpio. Zaragoza, ca. 3–7 km ESE of Zaragoza, ca. 3 air km E of Zaragoza, W to WSW of Cerro El Viejo peak on local

CHEILANTHES ARIZONICA

1-track roads, Nesom 7716 (holotype NY!; isotypes CAS!, TEX!). Figs. 76D–F. A C. hintoniorum laminis grandioribus et late pentagonis, segmentibus longioribus et indusiis longi-decurrentibus distincta. (Alluding to the frond dissection appearing like some members of the Apiaceae.)

Rhizomes compact, ascending, 2–3 mm diam.; rhizome scales linear-lanceolate, dark red-brown to black, lustrous, 4–5 mm long, entire; fronds to 38 cm long, clumped; stipes ca. 1⁄3 the frond length, dark reddish brown, lustrous, terete, the rachises and rachillae deeply grooved and green-winged, glabrous; blades pentagonal, quadripinnate; pinnae 8–10 pairs, the basal pair exaggerated basiscopically; segments mostly 10–15 mm long, ca. 1 mm wide, terminal ones mostly 20–25 mm long; adaxial surface glabrous, slightly pebbly; abaxial surface glabrous; laminar margins recurved and well differentiated into 0.2–0.3 mm wide longdecurrent entire false indusia covering the abaxial laminar surfaces; vein ends visible but not in deep pits; spores brown.

183

Cheilanthes pyramidalis Fe´e subsp. arizonica Maxon, Amer. Fern J. 8: 116. 1918. Cheilanthes pyramidalis Fe´e var. arizonica (Maxon) Broun, Index N. Amer. Ferns 51. 1938. Type. U.S.A. Arizona: Huachuca Mts., Ramsey Canyon, Goodding 1327 (US!; isotype NY!). Cheilanthes lerstenii Mickel & Beitel, Mem. New York Bot. Gard. 46: 117. 1988. Type. Mexico. Oaxaca: Distrito Sola de Vega, 2–9 km N of bridge at Juchatengo, 47–54 km S of Sola de Vega, Mickel 6238 (NY!).

Rhizomes compact, horizontal, 2–3 mm diam.; rhizome scales linear-lanceolate, lustrous, bicolorous black, with narrow brown margins; fronds to 30(–50) cm long, clumped; stipes 1⁄2–2⁄3 the frond length, castaneous to dark brown, grooved, glabrous; blades ovate to deltate, 3–4-pinnate-pinnatifid, 4–10 cm wide; pinnae 5–10 pairs, most equilateral, basal pair conspicuously larger than adjacent pair, inequilateral, proximal basiscopic pinnules exaggerated; ultimate segments elliptic to linear-oblong, not bead-like, the largest 4–10 mm long; adaxial surfaces glabrous; abaxial surfaces with scattered to rare, minute, 1-celled hairs, in northern part of range as reddish, scattered, sessile glands; laminar margin recurved and strongly differentiated, 0.2–0.5 mm wide, covering most of the abaxial surface with short, regular, marginal papillae (0.1 mm long), not decurrent; sori more or less continuous around segment margins; sporangia 32-spored; spores brown to grayish brown; n⫽2n⫽90, apogamous (USA).

Distribution. Oak woods, steep slopes, base of boulders; 1950– 2100 m. Mexico. Specimens Examined. NL (Hinton et al. 23441, ANSM, TEX).

From C. hintoniorum, C. apiacea differs in having larger fronds (25–38 ⫻ 9–18 cm), broadly pentagonal blades, more dissected blades which are 4-pinnate at bases (vs. 3-pinnate), and longer segments (mostly 10–15 mm long, the longest ones 15–20 mm long, the terminal ones mostly 20–25 mm long). The indusia of C. apiacea are long-decurrent and more differentiated than those of C. hintoniorum, their margins are pebbly but not distinctly papillate, and vein endings lack the white farina found in C. pellaeopsis. In constrast to C. hintoniorum, the rhizome scales are black to dark red-brown, lustrous, entire, 4–5 ⫻ 0.3– 0.5 mm at base, 10–12 cells wide (vs. black, bristle-like, 5–7 mm long, 3–4 cells wide at base), and the fronds have 8–10 pairs of pinnae (vs. 4–6 pairs).

6. CHEILANTHES ARIZONICA Cheilanthes arizonica (Maxon) Mickel, Phytologia 41: 433. 1979. Figs. 71A, B, 77C–E.

Distribution. Rocky slopes and ledges, shady ravines, commonly on igneous substrates; (800–)1750–2700 m. USA (Ariz); Mexico; Guat, Hond. Selected Specimens Examined. Chih (Knobloch 654, DS, MSC). Chis (Breedlove 29955, NY). Coah (Wendt & Alemcowicz 521, LL, TEX). Dgo (Palmer 150, NY). Hgo (Breedlove 59484, CAS). Jal (Koch & Fryxell 89200, NY). Me´x (Tejero-Dı´ez, 2839, IZTA, NY). Mich (Madrigal Sa´nchez 2687, IEB). Nay (Rose 2211, NY p.p.). NL (Muller 2845, LL, UC, US). Oax (Mickel 5263, NY). Qro (Carranza 2862, IEB). Sin (Breedlove 18570, NY). Son (Phillips 642, ARIZ, 674, ARIZ; White 4323, NY). Zac (Fitzner s.n., 31 July 1970, MSC). Unverified, Doubtful, or Mistaken Reports. Gro (Lorea 2742, FCME, cited by Lorea-Herna´ndez & Vela´squez M., 1998, but not verified).

The non-decurrent indusia with glabrous surfaces and regular, short-papillate margins distinguish this taxon from C. marginata and C. pyramidalis (decurrent to long-decurrent, long-papillate indusial margins), C. complanata (long- or nondecurrent, entire indusia), and C. membranacea (see key for char-

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acters). Cheilanthes lerstenii was described from Oaxaca, but the only character separating it is the presence of reddish glands in specimens from the southwestern United States and northernmost Mexico. Collections from further south have the same 1celled glandular hairs, though they are inconspicuous. The color and abundance of these glands seem to be clinal, so the two species are here treated as one.

Cheilanthes aurantiaca is distinct by the lanceolate to elliptic, pinnate-pinnatifid to bipinnate blade and the gold farina on the abaxial surfaces of the pinnae.

8. CHEILANTHES AUREA Cheilanthes aurea Baker in Hooker & Baker, Syn. Fil. 453. 1868. Type. Guatemala, 1862, Salvin & Godman s.n. (K!).

7. CHEILANTHES AURANTIACA Cheilanthes aurantiaca (Cav.) T. Moore, Index Fil. xxxviii. 1857. Figs. 75A–D. Pteris aurantiaca Cav., Descr. Pl. 266. 1801. Type. Mexico. Me´xico: “montes de Chalma,” Ne´e s.n. (MA). Pteris lutea Cav., Descr. Pl. 267. 1801. Cheilanthes lutea (Cav.) T. Moore, Index Fil. 246. 1861. Aleuritopteris lutea (Cav.) E. Fourn., Mexic. Pl. 1: 121. 1872. Type. Mexico. Me´xico: “montes de Chalma,” Ne´e s.n. (MA). Allosorus ochracea Hook. in Bentham, Pl. Hartw. 55. 1840. Cheilanthes ochracea Hook., Sp. Fil. 2: 114. 1858. Type. Mexico. [Michoca´n:] Morelia, Hartweg 418 (K!; isotype L, photos NY!, TEX!, US!).

Rhizomes compact, horizontal to ascending, 3–4 mm diam.; rhizome scales lanceolate, concolorous, tan, bicolorous with age, entire, 4–5 mm long; fronds to 36 cm long, clumped; stipes 1/9–1⁄3 the length of the fronds, terete, red-brown, lustrous, with scattered to sparse, tan, lanceolate scales, varying to hairs on adaxial rachial surfaces; blades lanceolate to elliptic, 4–8.5 cm wide, narrowed at bases, pinnate-pinnatifid to bipinnate; pinnae 7–14 pairs, linear-lanceolate; adaxial surfaces with scattered, white, jointed hairs ca. 1 mm long, more abundant along blade margins; abaxial surfaces with yellow-orange farina; sori marginal; blade margins recurved and differentiated, entire; sporangia 32-spored; spores light to dark brown.

Rhizomes compact, stout, ascending, 3–5 mm diam.; rhizome scales ovate-lanceolate to linear-lanceolate, concolorous tan, entire; fronds to 32 cm tall, clumped; stipes 1⁄3–1⁄2 the frond length, abscised old fronds leaving 1–1.5 cm long stipe bases, castaneous, grooved, sparsely scaly near base, sparsely yellow farinose throughout, var. palmeri with white jointed hairs 1.5–2 mm long; blades 2–3-pinnate-pinnatifid, broadly deltate to pentagonal (or ovate-lanceolate), 5–13 cm wide, the basiscopic pinnules of the basal pinna pair somewhat to greatly exaggerated; pinnae 6–12 pairs, segments rounded to oblong; adaxial surfaces generally glabrous or with scattered jointed white hairs 0.4–1 mm long, but sparse, long, white hairs along the axes, major veins, and blade margins; abaxial surfaces densely yellow farinose (but not continuous to scattered), with scattered long segmented white hairs on axes, major veins, and blade margins; sori at vein ends, laminar margins glabrous, recurved with firm, modified, 0.5 mm wide false indusia on tips of smaller segments, continuous on larger segments, entire, glabrous or strongly fimbriate-ciliate; sporangia 32-spored; spores black. Distribution. Terrestrial on banks and among rocks in dryish woods; 450–2550 m. Mexico; Guat, Salv. Key to the Mexican Varieties of Cheilanthes aurea 1. Indusial margins entire; blade hairs 0.5 mm; indusial hairs 0.1–0.2 mm; stipe hairs 1 mm; farina abundant; blades deltate to pentagonal; pinnae 6–8 pairs. ........... 8a. var. aurea. 1. Indusial margins fimbriate; blade hairs 1 mm; indusial hairs 0.5 mm; stipe hairs 2 mm; farina sparse; blades lanceolate; pinnae 9–12 pairs. ...................................... 8b. var. palmeri.

8a. Cheilanthes aurea var. aurea

Figs. 70C, D.

Var. aurea is distinct from var. palmeri in having indusial margins entire, blade hairs 0.5 mm, indusial hairs 0.1–0.2 mm, stipe hairs 1 mm, farina abundant, blades deltate to pentagonal, and pinnae 6–8 pairs; 2n⫽60 (Oax).

Distribution. On and among rocks, mostly in open oak or pine-oak forest; 700–2200 m. Mexico. Selected Specimens Examined. Dgo (Sa´nchez 739, MEXU). Gro (Matuda 37306, MEXU). Hgo (Matuda 32549, MEXU). Jal (McVaugh 20111, MICH). Me´x (Hinton 918, NY, 4350, MEXU). Mich (Gonza´lez Medrano 17024, MEXU). Mor (Pringle 10265, NY). Nay (Breedlove 44622, CAS, MEXU). Oax (de Avila 359, MEXU). Sin (Breedlove 62991, CAS).

CHEILANTHES BONARIENSIS

185

Selected Specimens Examined. Jal (McVaugh 13489, MICH; Pringle 2584, NY, 5412, CHAPA, MEXU, 11779, UC; Knobloch 2477, US). Nay (Feddema 791, MEXU; Mexia 854, UC p.p.; Rose & Painter 7410, US, 7587, US).

9. CHEILANTHES BOLBORRHIZA Cheilanthes bolborrhiza Mickel & Beitel, Mem. New York Bot. Gard. 46: 107. 1988. Type. Mexico. Oaxaca: Distrito Pochutla, 43 km N of Pochutla, 11 km N of Candelaria, Mickel 5207 (NY!). Figs. 69C–E.

Distribution. Roadside banks and wooded river banks; 450– 1050 m. Mexico; Guat, Salv. Selected Specimens Examined. Gro (Berkley et al. 17M757, F; Mickel 726, NY; both intermediate between the two varieties). Me´x (Hinton 6137, US). Mor (Sandoval 36, 42, UAMIZ). Oax (Mickel 845, 6228a, 6249, 6269, 6945, NY).

Cheilanthes aurea var. aurea is distinguished by its yellowfarinose abaxial blade surfaces and deltate, tripinnate to tripinnate-pinnatifid blades. It is close to var. palmeri in dissection, having yellow farina, broad rhizome scales, broad indusia, and large black spores, but var. aurea has fewer and shorter hairs on blade margins (0.5 vs. 1 mm), indusia (0.1–0.2 vs. 0.5 mm), and stipes (mostly 1 vs. 2 mm), abundant farina (vs. very sparse), lanceolate blades (vs. deltate), fewer pinna pairs (6–8 vs. 9–12), and blades 5–7 cm vs. 6–13 cm wide.

Rhizomes short, compact, horizontal, 3–5 mm diam.; rhizome scales linear-lanceolate, orange-tan, dull, concolorous, 4 mm long, entire; root bases swollen (2–4 ⫻ 1–2 mm wide), black, filled with starchy storage material; fronds to 90 cm long, clumped; stipes 1⁄3–1⁄2 the frond length, stramineous, castaneous only at very base, lustrous, glabrous, grooved; blades broadly deltate, 3– 4-pinnate-pinnatifid, basal pinnae opposite to subopposite, rachis grooves running into pinna grooves, costae and blade rachises with green stripes on upper surfaces, costae winged, proximal pinnae exaggerated basiscopically; ultimate segments elliptic to lanceolate, thin, glabrous; veins evident; sori marginal, indusia lateral on segments (not at tips or in sinuses), 0.5 mm wide, margins entire, surfaces with whitened, modified cells; spores gray-brown.

8b. Cheilanthes aurea var. palmeri (D. C. Eaton) Mickel, comb. et stat. nov. Figs. 75E–H. Cheilanthes palmeri D. C. Eaton, Proc. Amer. Acad. Arts 22: 464. 1887. Type. Mexico. [Jalisco:] Guadalajara, Palmer 223 (YU!; isotypes GH!, NY!, YU!-2 sheets).

Var. palmeri is distinct from var. aurea in having indusial margins fimbriate, blade hairs 1 mm, indusial hairs 0.5 mm, stipe hairs 2 mm, farina sparse, blades lanceolate, and pinnae 9–12 pairs. Distribution. Moist, tropical wooded stream banks, and steep slopes in pine-oak woods; 900–1900 m. Mexico; Salv. Specimens Examined. Jal (Breedlove 64102, CAS, UC; Ruelas 16221, UC). Me´x (Tejero-Dı´ez 2952, IZTA). Mor (Lyonnet 2575, IEB, MEXU; Riba 1667, IEB, 1913, MEXU; Sandoval 9, NY). Oax (only the type).

Cheilanthes bolborrhiza may be related to C. skinneri but differs in its more delicate appearance, thinner texture, concolorous rhizome scales, indusia Ⳳ continuous, gray-brown spores, and unique root storage organs. It may represent a distinct group within Cheilanthes.

10. CHEILANTHES BONARIENSIS Distribution. Shaded banks, among rocks in pine-oak forests and dry tropical deciduous forests; 1500-2550 m. Mexico.

Cheilanthes bonariensis (Willd.) Proctor, Bull. Inst. Jamaica, Sci. Ser. 5: 15. 1953. Figs. 74A–E.

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CHEILANTHES

Acrostichum bonariense Willd., Sp. Pl., ed. 4, 5(1): 114. 1810. Notholaena bonariensis (Willd.) C. Chr., Index Filic. 6. 1905. Type. [Argentina. Buenos Aires:] “Bonaria” (B-Willd. 195371!). Pteris aurea Poir. in Lamarck, Encycl. 5: 710. 1804. Notholaena aurea (Poir.) Desv., Me´m. Soc. Linn. Paris 6: 219. 1827. Type. Peru. Joseph de Jussieu s.n. (P-Juss., sheet 1333!, photo GH). Cheilanthes ferruginea Willd. ex Link, Enum. Hort. Berol. Alt. 2: 463. 1822. Notholaena ferruginea (Link) Hook., Sec. Cent. Ferns, t. 52. 1861, hom. illeg., non Desv., 1813. Type. “America meridionali.” (B; photo GH). Notholaena chiapensis Rovirosa, Pteridogr. Sur Me´xico 229, t. 48, f. 1–6. 1909. Type. Mexico. Chiapas: “Prope El Sacramento,” Rovirosa 1077 (PH, frag. US!; isotype GH!).

indument abaxially of pectinate scales rather than long, matted hairs. There is considerable variation in the number of pinna lobes (3–8 pairs per pinna) and degree of cutting, occasionally less than 1⁄4 to 3⁄4 of the way to the costa. This is the most widespread and abundant pteridophyte species in Mexico, being in all states except Baja California Norte, Tabasco, and those of the Yucatan Peninsula.

Rhizomes short-creeping, horizontal, ca. 3 mm diam.; rhizome scales linear-lanceolate, strongly bicolorous with lustrous dark red-brown to black central stripe, 3 mm long, entire; fronds to 75 cm tall, clumped; stipes 1/6–1⁄3 the frond length, terete, castaneous to black, clothed with straight, somewhat appressed, white to tan hairs, 2 mm long; blades linear-elliptic, pinnatepinnatifid; pinnae 15–44 pairs, cut 1⁄4–3⁄4 to costae, with 3–5(–8) pairs of lobes; adaxial surfaces with abundant to scattered, appressed, stiff, 1-celled hairs (1 mm long); abaxial surfaces densely clothed with white to rusty matted hairs, these obscuring the laminar surfaces; sori at ends of veins, laminar margins slightly recurved, not modified into false indusia; sporangia 32-spored; spores gray, dark brown to black; n⫽2n⫽90, apogamous (Ariz).

Cheilanthes brachypus (Kunze) Kunze, Linnaea 23: 307. 1850. Figs. 74L–Q.

11. CHEILANTHES BRACHYPUS

Cheilanthes squamosa Hook. & Grev. var. brachypus Kunze, Linnaea 18: 340. 1844. Notholaena brachypus (Kunze) J. Sm., Ferns Brit. For. 172. 1866. Type. Mexico. Leibold 52 (B!; isotypes BM, photo US!, K!, photo GH, P!).

Rhizomes compact, ascending, ca. 3 mm diam.; rhizome scales linear-lanceolate, 6–12 mm long, concolorous orange-tan to slightly bicolorous with a castaneous center and pale margins, lustrous, entire or with occasional teeth; fronds to 30 cm tall, clumped; stipes ca. 1/10 the frond length, only ca. 2–3 cm long, dark brown, terete, with linear scales 2–3 mm long, bases swollen, erose, dull, light brown with whitish margins, mixed with fine hairs 2 mm or more long; blades oblanceolate, longest pinnae 1.2–3.5 cm long just above middle of blade, narrowing proximally, pinnate-pinnatifid; pinnae 9–20 pairs, equilateral; adaxial surfaces dark green with scattered to dense tomentum of curly hairs (1 mm long); abaxial surfaces densely covered with lanceolate scales, these erose at base, overlying a dense rusty tomentum of fine hairs; laminar margins recurved and strongly modified into false indusia 0.5 mm wide; spores tan to black; 2n⫽60 (Mor).

Distribution. Dry rocky slopes; 600–3200 m. Sw USA; Mexico; Guat, Hond, Nic, CR, Pan; Gr Ant; Col, Ven, Ec, Peru, Braz, Bol, Chile, Parag, Arg, Uru. Selected Specimens Examined. Ags (Cowan 4027, UC). BCS (Brandegee 658, NY). Chih (Pringle 462, NY). Chis (Ghiesbreght 304, NY). Coah (Palmer 1399, NY). Col (Alava & Cook 1706, UC). DF (Orcutt 3616, NY). Dgo (Breedlove 59252, NY). Gro (Hinton 9481, NY). Gto (Galva´n 3474, NY). Hgo (Gonza´lez Quintero 3136, NY). Jal (McVaugh 16803, NY). Me´x (Rose 6800, NY). Mich (Feddema 35, NY). Mor (Orcutt 3831, NY). Nay (Garcı´a Necchea 16910, UC). NL (Hinton 19912, MEXU). Oax (Mickel 6806, NY). Pue (Nee 33011, NY). Qro (Argu¨elles 125, MEXU). SLP (Parry & Palmer 994, NY). Sin (Breedlove 16936, NY). Son (Torres C. 3852, NY). Tam (Bartlett 10111, NY). Tlax (Clausen s.n., 23 Aug 1955, NY). Ver (Bourgeau 3153, NY). Zac (Anderson 12614, NY).

Cheilanthes bonariensis superficially resembles Astrolepis sinuata in dissection and heavy indument, but the latter species has

Distribution. Rocky slopes and stream banks; 50–1950 m. Mexico; Guat, Hond, Salv, Nic, CR. Selected Specimens Examined. Ags (Siqueiros D. 4285, IEB). Chis (Breedlove 22536, NY). Col (Lott 3038, NY). Gro (Iltis & Solheim 3072, NY). Gto (Galva´n 2407, IEB). Jal (Pringle 1787, NY). Me´x (Tejero-Dı´ez 2410, IZTA). Mich (Hinton 15433, NY). Mor (Ramı´rez R. 66, XAL). Nay (Feddema 457, NY). Oax (Mickel 6233, NY). SLP (Parry & Palmer 998, NY). Sin (Keil 8767, NY). Son (Fishbein 1897, ARIZ). Ver (Purpus 7870, NY).

CHEILANTHES CHIPINQUENSIS

Although this species traditionally was placed in Notholaena, it has well differentiated false indusia. Cheilanthes brachypus superficially resembles Astrolepis sinuata, but differs from that species in the false indusia, sori limited to vein ends, and nonarticulate pinnae with long, unbranched hairs on both surfaces (although mixed with scales on the abaxial surfaces).

12. CHEILANTHES BRANDEGEEI Cheilanthes brandegeei D. C. Eaton, Bull. Torrey Bot. Club 17: 215, t. 104. 1890. Syntypes. Mexico. Baja California Norte: Magdalena Island and San Benito, Jan, Mar, & Apr 1889, Brandegee s.n. (YU!; isosyntypes NY!–2 sheets, P!). Figs. 75M–O. Rhizomes stout, compact, horizontal, 2–3 mm diam.; rhizome scales linear-lanceolate, concolorous orange-tan, 3–4 mm long, entire; fronds to 20 cm long, clumped; stipes ca. 1⁄3 the frond length, castaneous, lustrous, glabrous, terete; blades deltate or pentagonal, bipinnate-pinnatifid to tripinnate at base, 6–7 cm wide; pinnae 5–7 pairs, the proximal ones slightly inequilateral; segments ovate with rounded apices; adaxial surfaces with sparse hairs 0.2–0.3 mm long; abaxial surfaces with scattered 1-celled white hairs 0.3–0.5 mm long; texture thin; laminar margins curved, deeply lobed, weakly differentiated; spores brown.

Distribution. Among (basaltic) rocks on shaded north slopes, and among trees and shrubs; 30–1250 m. Mexico. Selected Specimens Examined. BCN (Montgomery 83-28, NY; Moran 23952, SD; Wiggins 7649, NY, P, 15080, ARIZ; Wiggins & Thomas 176, NY). BCS (Breedlove 62331, NY; Moran 18934, MICH, 19679, ARIZ, ASC, SD, TEX). Cedros Isl (Palmer 707, NY, P).

Cheilanthes brandegeei is one of the most distinct species of the genus with its broadly deltate blades, deeply lobed indusia, and scattered, long, white, 1-celled hairs. Molecular studies (Cranfill, unpublished data) suggest that this species might be more closely allied to Bommeria than to Cheilanthes.

187

Allosorus chaerophyllus M. Martens & Galeotti, Me´m. Foug. Mexique 47, pl. 11. 1842. Type. Mexico. Oaxaca: Juquila, Galeotti 6367 (BR?, not found; isotype P!).

Rhizomes stout, compact, horizontal; rhizome scales linearlanceolate, 2–3 mm, concolorous, dark castaneous to black, lustrous; fronds 30–52 cm long, clumped; stipes 14–34 cm long, 1⁄2–3⁄4 the frond length, castaneous, grooved, glabrous; blades 12–20 ⫻ 12–20 cm (length:width ratio 0.8–1.1:1), tripinnate-pinnatifid to quadripinnate, strongly deltate or pentagonal, basal pinnae largest, glabrous on both surfaces; pinna pairs 15–20 (distal 2–3 pairs entire, upper median 5–8 pairs pinnate, lower median 3–5 pairs bipinnate, basal 1–3 pairs tripinnate), terminal segments pinnatifid, ovate, 0.4–0.8 cm ⫻ 2–3 mm (length:width ratio 1.8–3:1); basal pinnae tripinnate, exaggerated basiscopically, 10–14 ⫻ 7–10 cm (length:width ratio 1.1– 1.8:1), (12–)14–16 pinnule pairs (distal 2–6 pairs entire, median 5–7 pairs pinnate, 1–5 basal pairs bipinnate); basal basiscopic pinnule 7–7.5 cm long, with 10–12 lobes, terminal segment 2–3 mm wide, basal lobe with 4–7 lobules; adaxial and abaxial surfaces glabrous, the veins prominent abaxially; sori at vein endings, laminar margins curved, strongly modified into 0.3–0.5 mm wide false indusia, margins suberose, surfaces slightly whitened; spores black.

Distribution. Exposed or shaded banks; (500–)900–2500 (–3200) m. Mexico; CR; Col. Selected Specimens Examined. Chis (Breedlove 22683, NY). Col (McVaugh 16108, NY). DF (Lyonnet 239, MEXU). Dgo (Maysilles 7003, MEXU). Gro (Hinton 9491, NY). Gto (Rzedowski 40579, IEB). Jal (Pringle 2026, NY). Me´x (Mickel 680, NY). Mich (Hinton 12609, NY). Mor (Ramı´rez R. 60, XAL). Nay (Avelar Mun˜oz 16911, UC). Oax (Mickel 1318, NY). Pue (Purpus 4176, UC). Sin (Bartholomew 2516, NY). Son (Perrill 5256, MEXU).

Cheilanthes chaerophylla is the most widespread member of the C. angustifolia complex, which includes also C. cuneata and C. decomposita. See discussion under C. angustifolia.

13. CHEILANTHES CHAEROPHYLLA

14. CHEILANTHES CHIPINQUENSIS

Cheilanthes chaerophylla (M. Martens & Galeotti) Kunze, Linnaea 23: 243, 307. 1850. Fig. 70E.

Cheilanthes chipinquensis Knobloch & Lellinger, Amer. Fern J. 59: 8. 1969. Type. Mexico. Nuevo Leo´n: Chipinque Mesa,

188

CHEILANTHES

Rhizomes short-creeping, horizontal, 2–3 mm diam.; rhizome scales linear-lanceolate, 3–4 mm, bicolorous with black center and orange margins, tips attenuate, entire; fronds 12–37 cm, clumped; stipes 1⁄3–1⁄2 the frond length, brown, terete, densely clothed with appressed or ascending tan hairs 2–3 mm and hairlike scales; blades lanceolate to narrowly lanceolate, bipinnate to tripinnate-pinnatifid, 3.5–7.5 cm wide, firm to subcoriaceous; pinnae 17–25 pairs; costae with hairs and hair-like scales as on stipes; terminal segments often unequally pinnately lobed at bases, subterminal ultimate segments ovate, ca. 1 mm diam., terminal oblong, 3 ⫻ 1 mm; adaxial blade surfaces with scattered crinkly white hairs; abaxial blade surfaces sparsely to densely hairy with crinkly white or tan hairs, these generally not totally obscuring the laminar surfaces; indusia slightly modified, continuous; sporangia 64-spored; spores tan to light brown; 2n⫽58, 60 (NL).

Rhizomes short-creeping, horizontal, 1–3 mm diam.; rhizome scales linear-lanceolate, lustrous dark brown concolorous or with paler margin, entire; fronds to 40 cm long, approximate to distant; stipes ca. 1⁄2 the frond length, dark brown, lustrous, terete, clothed with 1–2 mm long hairs and filiform scales to glabrescent with age; blades oblong-lanceolate to ovate, quadripinnate, 2–8 cm wide; rachises terete, with scattered scales and sparse hairs; pinnae 10–12 pairs (more in California), equilateral; segments bead-like, most about 1 mm long; adaxial surface glabrous; abaxial surfaces with costal scales ovate-lanceolate to broadly deltate, deeply cordate at base, with overlapping basal lobes, conspicuous, ca. 1 ⫻ 0.5–1 mm, imbricate, concealing ultimate segments but not exceeding their margins, ciliate (usually only on proximal 1⁄2), often irregularly coarsely toothed and somewhat flexuous, on the segments reduced ciliate scales, some appearing as branched hairs, apices attenuate; lamina margins recurved, weakly differentiated, entire; sporangia 64-spored; spores brown.

Distribution. Terrestrial or epipetric on limestone in pine-oak forests; 800–1200m. Mexico.

Distribution. Rocky slopes and ledges, usually on igneous substrates; 0–1600 m. USA (Calif); Mexico.

outside Monterrey, Knobloch 1996B (MSC!; isotypes F!, GH!, MEXU!, MICH!, UC!, US!). Figs. 82F–H.

Specimens Examined. NL (Aguirre s.n., 6 Dec 1976, ENCB; McPherson 883, CAS, MICH; Muller 2701, GH p.p.; Pennell 16951, US; Pringle 2776, UC; Reeves 6087, 6098, ASC; Rodrı´guez L. s.n., 28 Apr 1985, UNL).

Cheilanthes chipinquensis is closely allied to the widespread C. tomentosa but can be distinguished by the less dense indument and the terminal segments about three times the length of the lateral segments. The former has 64-spored sporangia, versus the 32-spored sporangia in C. tomentosa. Knobloch (pers. comm.) suggested that the diploid C. chipinquensis may be a parent of the triploid C. tomentosa. Reeves (1979) pointed out some intermediate specimens, but was unsure if they were hybrids. Cheilanthes chipinquensis is generally found growing together with C. tomentosa, and care is needed for their separation.

15. CHEILANTHES CLEVELANDII Cheilanthes clevelandii D. C. Eaton, Bull. Torrey Bot. Club 6: 33. 1875. Type. U.S.A. California: San Diego, Cleveland s.n. (YU!; isotypes GH!, P!, US!). Figs. 80J–O.

Selected Specimens Examined. BCN (Brandegee s.n., 30 Apr 1889, UC p.p.; Fischer et al. 2073, ENCB; Hevly 2073, ARIZ; Moran 16168, MEXU, 24597, SD; Orcutt s.n., 13 Apr 1882, NY; Porter 160, CAS, MEXU; Pringle s.n., April 1882, LL, US; Wiggins & Gillespie 4048, CAS, DS, MEXU, NY, US; Yatskievych 82-171, ARIZ).

Cheilanthes clevelandii appears hairy adaxially because the ciliate scales on abaxial surfaces extend beyond the segment margins. It is close to C. covillei, but their ranges rarely overlap. Reeves (1979) recognized two varieties, var. clevelandii being widespread and var. clokeyi T. Reeves & Cochrane (ined.), confined to California. Cheilanthes covillei appears scaly only, has glabrous blades, and has costal scales ciliate only on basal lobes.

16. CHEILANTHES COMPLANATA Cheilanthes complanata A. R. Sm., Amer. Fern J. 70: 19. 1980. Type. Mexico. Chiapas: Mpio. Motozintla de Mendoza, slope of Cerro Mozotal, Breedlove 41747 (DS!; isotype MEXU!). Rhizomes compact, horizontal to ascending, 2–3 mm diam.; rhizome scales linear-lanceolate, bicolorous, black with narrow

CHEILANTHES COVILLEI

189

brown margin, lustrous, 3–4 mm long, entire; fronds 30–42 cm long, clumped; stipes 1⁄2–2⁄3 the frond length, castaneous, terete or slightly grooved distally, glabrous; blades deltate, attenuate at apices, tripinnate to tripinnate-pinnatifid; pinnae ca. 15 pairs, equilateral except the basiscopically exaggerated basal pair; segments narrowly obovate to narrowly elliptic; adaxial surfaces pebbly, but without papillae; abaxial surfaces glabrous except for scattered, minute, orange, 1-celled hairs; laminar margins recurved, somewhat differentiated into 0.3 mm wide, smooth false indusia, entire, not or only slightly decurrent along next axis; spores gray-brown.

At least some of the specimens cited by Yatskievych and Moran (in Davidse et al., 1995) are here treated as C. decurrens, which see.

Distribution. Terrestrial or epipetric in cloud forests; 1960– 3000 m. Mexico.

Rhizomes compact, ascending, 2–3 mm diam.; rhizome scales linear-lanceolate, 3–4 mm long, bicolorous with a black central stripe with brown margins, lustrous, entire; fronds to 42 cm long, clumped; stipes 2⁄3 the frond length, dark reddish brown, lustrous, grooved, glabrous; blades deltate, attenuate at apices, tripinnate at base; pinnae ca. 15 pairs, equilateral except basal pair which is exaggerated basiscopically, segments oblong, obtuse to acute; adaxial surfaces glabrous; abaxial surfaces glabrous except for sparse orangish 1-celled hairs; laminar margins recurved, somewhat differentiated as false indusia, 0.3–0.5 mm wide, lobed, discontinuous at segment apices, not at all decurrent; spores gray-brown.

Cheilanthes complanata is distinct by its non-decurrent, entire indusia, castaneous stipes and rachises, bicolorous rhizome scales, and conspicuous swollen vein endings (generally visible abaxially). It is close to C. harrisii Maxon of Jamaica in its nondecurrent, entire, barely modified false indusia and deltateattenuate blades, but C. harrisii differs in its rhizome scales being somewhat flexuous with kinky hair-tips, frond axes darker (atropurpureous vs. castaneous), and blades more dissected (to quadripinnate vs. tripinnate to tripinnate-pinnatifid).

16b. Cheilanthes complanata var. interrupta Mickel, var. nov. Type. Mexico. Guerrero: 16 km sobre el camino de Puerto del Gallo a Atoyac de Alvarez, Espin M. 149 (holotype NY!; isotypes ENCB!, FCME!). Figs. 77J–L. A C. complanata var. complanata indusiis lobatis vel discretis distincta. (Alluding to the indusia being interrupted along the segment margins, rather than continuous as in the type variety.)

Key to the Mexican Varieties of Cheilanthes complanata 1. Indusia continuous, unlobed; Chis. ..................... 16b. var. complanata. 1. Indusia discontinuous, deeply lobed; Gro. ............ 16a. var. interrupta.

16a. Cheilanthes complanata var. complanata Figs. 77F–H. Var. complanata differs from var. interrupta in having continuous (vs. discontinuous) indusia.

Distribution. Slopes in laurel forests; 1960 m. Known only from the type collection. Var. interrupta resembles var. complanata in its narrowly deltate-attenuate blades, castaneous blade axes, and bicolorous rhizome scales, but differs in the indusia being lobed to discrete, rather than entire.

17. CHEILANTHES COVILLEI Distribution. Terrestrial or epipetric in evergreen cloud forests, rocky slopes in pine woods; 2750–3000 m. Mexico. Specimens Examined. Chis (Breedlove 40335, DS, 55870, CAS, ENCB, NY). Unverified, Doubtful, or Mistaken Reports. Ver (reported by Mickel & Beitel, 1988, but not verified).

Cheilanthes covillei Maxon, Proc. Biol. Soc. Wash. 31: 147. 1918. Type. U.S.A. California: Surprise Canyon, Panamint Mts., Coville & Funston 593 (US!). Figs. 81Q–V. Rhizomes short-creeping, horizontal, 2–4 mm diam.; rhizome scales lanceolate to linear-lanceolate, concolorous, dark brown to black, sclerotic, often weakly bicolorous with narrow, tan mar-

190

CHEILANTHES

gins, ca. 2 mm long, entire; fronds to 30 cm long, clumped; stipes dark reddish brown to atropurpureous, lustrous, terete, with scattered linear-lanceolate whitish scales to 3 mm long; blades lanceolate to ovate, 3–4-pinnate, 1.5–5 cm wide; pinnae ca. 10 pairs, equilateral; segments bead-like, 1–3 mm long; adaxial segment surfaces glabrous; abaxial segment surfaces glabrous or with a few scales near segment base, costal scales ovate-lanceolate, whitish with castaneous base, cordate with overlapping lobes, 2– 3 ⫻ 0.4–1.5 mm, strongly imbricate, ciliate only on basal lobes, usually concealing ultimate segments and exceeding them; laminar margins recurved and weakly differentiated, entire; sporangia 64-spored; spores brown; 2n⫽60 (USA).

pinnatifid to tripinnate, chartaceous, pinnule lobes obcuneate on both acroscopic and basiscopic sides of pinnules; rachises and costae with dense appressed filiform scales 1–2 mm long, 2 cells wide; adaxial surfaces dark green, glabrous; abaxial surfaces with scattered to dense, 0.3–0.5 mm long, whitish, catenate hairs and sparse orange-tan, filiform scales 0.5–1 mm long; laminar margins recurved, especially at segment tips, slightly modified into 0.5 mm wide entire false indusia; spores tan.

Distribution. Dry to moist rocky slopes, often near streams; 1050–2350 m. Mexico; Guat.

Distribution. Rocky slopes, cliffs, and ledges, usually on igneous substrates; 650–2200 m. USA (Ariz, Calif, Nev, Utah); Mexico. Selected Specimens Examined. BCN (Carter 2555, MEXU, UC; Gentry & Cech 8910, MEXU; Moran 7997, MEXU, US, 11394, MEXU, 10297, DS, 14607, US, 11543, UC, US, 20960, DS; Wiggins 9898, UC).

Selected Specimens Examined. Ags (McVaugh & Koelz 124, MICH). Chih (Knobloch 5933, US). Chis (Breedlove 14501, DS). Coah (Johnston et al. 11506B, LL, TEX). DF (Orcutt 4330, NY). Gro (Rose 9396, NY). Gto (Rzedowski 40582, ENCB). Hgo (Sa´nchez Mejorada 682, MEXU). Jal (McVaugh 17424, NY). Me´x (Rzedowski 18719, NY). Mich (McVaugh 18156, NY). Mor (Lyonnet 2675, CAS). NL (Rodrı´guez L. 147, ANSM). Oax (Camp 2488, NY). Pue (Purpus 3151, NY). Ver (Murrill & Murrill 40, NY). Unverified, Doubtful, or Mistaken Reports. Qro (Pin˜a 92, ENCB, cited by Dı´az-Barriga & Palacios-Rios, 1992, but not verified).

Cheilanthes covillei differs from C. clevelandii by having glabrous blades and costal scales ciliate only on the basal lobes. It is distinct from the much rarer in Mexico C. fendleri by having rigid, dark brown, strongly appressed rhizome scales. It hybridizes with C. parryi, C. newberryi, and C. fendleri in the southwestern United States to form rare, sterile diploids.

Cheilanthes cucullans resembles C. microphylla but has abundant hair-like scales on rachises and costae, long-creeping rhizomes, and long-ciliate rhizome scales. See key for comparison with C. notholaenoides, which also has dense hairs on its rachises.

18. CHEILANTHES CUCULLANS

19. CHEILANTHES CUNEATA

Cheilanthes cucullans Fe´e, Me´m. Foug. 7: 39, t. 25, f. 4. 1857. Type. Mexico. Valley of Mexico, Schaffner 82 (RB; isotype K!, frag. US!). Figs. 72G, H. Cheilanthes micromera Link, Hort. Berol. 2: 36. 1833. Cheilanthes microphylla (Sw.) Sw. β micromera (Link) T. Moore, Index Fil. 247. 1861. Type. Mexico. Cultivated at B (B, photo NY!; possible isotype BR!).

Rhizomes long-creeping, horizontal, 2 mm diam.; rhizome scales linear-lanceolate, ca. 2 mm long, concolorous orange-tan, often blackish at base or with age, margins often with cilia, 0.1– 0.3 mm long; fronds 14–35 cm long, distant; stipes 1⁄3–1⁄2 the frond length, castaneous to black, lustrous, terete, with filiform orangetan scales 2 cells wide; blades oblong-lanceolate, bipinnate-

Cheilanthes cuneata Link, Hort. Berol. 2: 40. 1833. Figs 70F, 71E, F. Pellaea cuneata (Link) J. Sm., Hist. Fil. 286. 1875. Cassebeera cuneata (Link) J. Sm., J. Bot. (Hook.) 4: 159. 1841. Type. Described from cultivated material (B, photo NY!; isotype, PH!).

Rhizomes compact, horizontal, 2–3 mm diam.; rhizome scales linear-lanceolate, 2–2.5 mm long, concolorous, brown to black, lustrous, entire; fronds 18–34 cm long, clumped; stipes 1⁄3–2⁄3 the frond length, castaneous, lustrous, grooved, glabrous; blades 8– 18 ⫻ 4–8 cm (length:width ratio 2–4:1), bipinnate-pinnatifid to tripinnate, lanceolate, proximal pinnae slightly reduced; pinna pairs 10–20 (distal 2–4 pairs entire, median 6–14 pairs pinnate),

CHEILANTHES DECURRENS

proximal 0–5 pairs bipinnate), terminal segments ovate, 0.5–1.5 cm ⫻ 2–4 mm (length:width ratio 2.5–7:1); proximal pinnae bipinnate, slightly exaggerated basiscopically, 3.2–6.5 ⫻ 1.6–3 cm (length:width ratio 1.7–2.5:1), 5–10 pinnule pairs (distal 3–4 pairs entire, proximal 2–6 pairs pinnate; basal basiscopic pinnules 1–1.7 cm long, 1–3 lobed, terminal segment 2–3 mm wide; adaxial and abaxial surfaces glabrous, the veins prominent abaxially; laminar margins recurved, modified into 0.5–0.8 mm wide false indusia, suberose, surfaces and margins with white papillae; spores black; 2n⫽180 (Oax).

191

siscopically, 15–20 ⫻ 8–10 cm (length:width ratio 1.6–2.2:1), with 14–18 pinnule pairs (distal 5–6 pairs entire, median 8–10 pairs pinnate, basal 2–3 pairs bipinnate); basal basiscopic pinnules 6–8 cm long, 11–13 lobes, terminal segments 1.5–2 mm wide, basal lobe with 2–3 lobules; laminar margins recurved, strongly modified into 0.3–0.5 mm wide false indusia, suberose to erose, surface whitened with papillae; spores black.

Distribution. Shaded banks, often near streams; 900–2300 m. Mexico. Distribution. Shaded, often rocky slopes; 1050–2800 m. Mexico. Specimens Examined. Chis (Breedlove 22683, DS). Gro (Lorea 3240, FCME). Jal (McVaugh 24347, MICH). Me´x (Huerta G. 16, IEB). Mich (Escobedo 1838, IEB). Mor (Lyonnet 864, UC). Oax (Mickel 4881, NY). Pue (Tlapa & Ubierna 1117, IEB). Unverified, Doubtful, or Mistaken Reports. Ver (Zola´ 851, MEXU, XAL, cited by Palacios-Rios, 1992, but not verified).

Cheilanthes cuneata is part of the C. angustifolia complex (including also C. chaerophylla and C. decomposita). See discussion of the complex under C. angustifolia.

20. CHEILANTHES DECOMPOSITA Cheilanthes decomposita (M. Martens & Galeotti) Fe´e, Me´m. Foug. 9: 11. 1857. Figs. 77M, N. Allosorus decompositus M. Martens & Galeotti, Me´m. Foug. Mexique 48, pl. 10, f. 2. 1842. Type. Mexico. Oaxaca: Juquila, Galeotti 6362 (BR!).

Rhizomes compact, horizontal, 2–3 mm diam.; rhizome scales linear-lanceolate, concolorous, dark reddish brown to black, lustrous, entire, 2.5–3 mm long; fronds 30–50 cm long, clumped; stipes 1⁄2–3⁄4 the frond length, castaneous, lustrous, grooved, glabrous; blades 12–20 ⫻ 12–20 cm (length:width ratio 0.8–1:1), tripinnate-pinnatifid to quadripinnate, strongly deltate, basal pinnae largest, glabrous on both sides; pinna pairs 14–20 (apical 2–5 pairs entire, upper median 4–8 pairs pinnate, lower median 4–7 pairs bipinnate, basal 1–2 pairs tripinnate), terminal segments conform, narrow, lanceolate, 0.7–1.5 cm ⫻ 1.5–2.5 mm (length:width ratio 4.5–6:1); basal pinnae pair exaggerated ba-

Selected Specimens Examined. DF (Sa´nchez 151, MEXU). Dgo (Sa´nchez 741, MEXU). Gro (Campos R. 1315, FCME). Jal (Pringle 2588, NY). Me´x (Tejero-Dı´ez 2578, IZTA). Mich (Arse`ne s.n., 9 Aug 1909, NY). Nay (McVaugh 16427, NY). Oax (Mickel 6064, NY). Sin (Bartholomew 2522, CAS; Gentry 6501, ARIZ). Ver (Mu¨ller 66, NY).

Cheilanthes decomposita is part of the C. angustifolia complex (including also C. cuneata and C. chaerophylla). See discussion of the complex and intermediates under C. angustifolia.

21. CHEILANTHES DECURRENS Cheilanthes decurrens Mickel, sp. nov. Type. Guatemala. Volca´n de Agua, 3300–3600 m, Maxon & Hay 3709 (holotype NY!). Figs. 77A, B, O, P. A C. complanata indusiis perlatis et longi-decurrentibus differt. (Alluding to the indusia being decurrent on the subtending axes.)

Similar to C. complanata in the entire indusia, deltateattenuate, 3–4-pinnate, blades 9–47 cm long, and bicolorous rhizome scales, but differs in the distinct broad, long-decurrent indusia covering the abaxial segment surfaces.

192

CHEILANTHES

entiated into false indusia; spores dark brown; n⫽2n⫽90, apogamous (Coah, USA).

Distribution. Rocky banks in montane rain forests, cloud forests; 2100–2500 m ((1500–)2800–3900 m in Guatemala). Mexico; Guat, Salv. Selected Specimens Examined. Chis (Breedlove 55788, ENCB, 55870, NY; Ghiesbreght 285, NY). Oax (Mickel 1271, NY). Guatemala (Brenckle 47-63, NY; Burger & Liesner 7441, F; Degener & Degener 26673, NY; Molina R. et al. 16485, NY, 30033, F, NY; Standley 81696, 84429, F; Tu¨rckheim 1167, NY p.p.). El Salvador (Seiler 1209, NY).

Paratypes of C. decurrens were cited by Smith (1981) as C. marginata, by Stolze (1981) as C. chaerophylla, and by Yatskievych and Moran (in Davidse et al., 1995) as C. complanata.

22. CHEILANTHES EATONII Cheilanthes eatonii Baker in Hooker & Baker, Syn. Fil. 140. 1867. Type. U.S.A. West Texas to El Paso, Wright 816 (US!; isotype NY!) Figs. 82J–N. Cheilanthes castanea Maxon, Proc. Biol. Soc. Wash. 32: 111. 1919. Cheilanthes eatonii Baker in Hooker & Baker f. castanea (Maxon) Correll, Wrightia 1: 258. 1949. Type. Mexico. Coahuila: Sierra Madre, 13 leagues south of Saltillo, Palmer 1390 (US!; isotype NY!). Myriopteris rufa, Fe´e, Me´m Foug. 8: 77. 1857, non Cheilanthes rufa, D. Don 1825. Cheilanthes schaffneri T. Moore, Index Fil. 252. 1861, non Aleuritopteris schaffneri E. Fourn., 1880, nec Cheilanthes schaffneri (E. Fourn.) Domin, 1913. Cheilanthes cinnamomea D. C. Eaton, Proc. Amer. Acad. 18: 186. 1883. Type. Mexico. [Veracruz:] Volca´n de Orizaba, pre`s de San Andres Chalahicanmecha, 2600 m, 1855, Schaffner 83. Type not seen, but a specimen at P labelled “Myriopteris rufa Fe´e, Mexico, Orizaba, Schaffner” is Cheilanthes eatonii, and the type description could apply to that species.

Rhizomes compact to short-creeping, horizontal, ca. 3 mm diam.; rhizome scales linear-lanceolate, bicolorous, with black center and tan margins, lustrous, 3–4 mm long, entire; fronds 15–38 cm long, clumped; stipes 1⁄4–1⁄3 the frond length, castaneous, terete, with abundant linear tan to white scales 1.5–2 mm long; blades narrowly lanceolate, 2–3-pinnate, 1.3–4 cm wide; rachises with loosely imbricate, appressed, linear scales mixed with hairs; pinnae 12–18 pairs, equilateral; segments oblong, obtuse; adaxial surfaces with abundant to sparse curly white to rusty hairs 0.5–1 mm long; abaxial surfaces and axes of pinnae and pinnules with lanceolate, entire scales and with a dense mat of curly hairs; laminar margins somewhat recurved, but not differ-

Distribution. Terrestrial or epipetric on limestone, less commonly basaltic slopes; 550–2850 m. Southwestern USA; Mexico; CR. Selected Specimens Examined. Ags (Rzedowski 25071a, NY). BCS (Moran 18752, ENCB p.p.). Chih (Pringle 6454, NY). Coah (Stanford et al. 202, NY). DF (Schaffner 43, NY). Dgo (Palmer 893, NY). Hgo (Pringle 11277, NY). NL (Grimes 2284, NY). Pue (Pittier 446, US). Qro (Dı´az B. 3774-9, IEB). SLP (Burgess 5432, ARIZ). Son (Hartman 360, NY). Tam (Stanford et al. 696, ARIZ). Ver (Tejero-Dı´ez 4198, IZTA). Zac (Palmer 260, NY).

Cheilanthes eatonii is a highly variable species with apogamous triploid and tetraploid cytotypes. Cheilanthes castanea has sometimes been treated as a distinct species. The type of C. castanea differs in having the segments more evident because the hairs of the abaxial surface are not as long and there are fewer scales on the costules, but most specimens of C. eatonii s.l. are intermediate in laminar pubescence. Cheilanthes eatonii reportedly crosses with C. villosa (Lellinger, 1985) in western Texas and southern New Mexico. Pinna rachis scales of C. eatonii range from narrowly lanceolate to deltate-lanceolate, whereas C. tomentosa, which is often confused with C. eatonii, has very few scales on the rachillae. Cheilanthes pinkavae T. Reeves, ined., according to Windham and Rabe (FNA Ed. Comm., 1993), and Lellinger (1985) is the name applied by Reeves (1979) to plants intermediate between C. eatonii and C. villosa, but it has not been validly published.

23. CHEILANTHES FARINOSA Cheilanthes farinosa (Forssk.) Kaulf., Enum. Filic. 212. 1824. Figs. 70A, B. Pteris farinosa Forssk., Fl. Aegypt.-Arab. 187. 1775. Aleuritopteris farinosa (Forssk.) Fe´e, Me´m. Foug. 5: 153, t. 12B, f. 1. 1852. Type. Yemen, Forsska˚l s.n. (isotype BM!). Aleuritopteris mexicana Fe´e, Me´m. Foug. 5: 153. 1852. Type. Mexico. Oaxaca: Llano Verde, Galeotti 6551 (P?, not found; isotypes BR!, K!). Pellaea pulcherrima Rovirosa, Pteridogr. Sur Me´xico, t. 14 [first of 2 plates so numbered]. 1909, nom. nud. Aleuritopteris chihuahuaensis Saiki, J. Phytogeogr. Taxon. 32(2): 85. 1984.

CHEILANTHES FEEI Type. Mexico. Chihuahua: Sierra Madre, Pringle 1441 (F; isotypes NY!, P!–3 sheets, UC!, US!).

Rhizomes compact, stout, ascending, 2–4 mm diam.; rhizome scales linear-lanceolate, concolorous, dark reddish brown to bicolorous with black center and brown margins, lustrous, margins entire, 5–6 mm long; fronds to 50 cm long, clumped; stipes 1⁄2– 2 ⁄3 the frond length, terete but grooved adaxially at junction with rachises, atropurpureous with sparse, lanceolate, dark reddish brown scales, occasionally sparsely white-farinose, abscised fronds leaving stipe bases 1.5–2.5 cm long, or not abscised; blades deltate to ovate-pentagonal, pinnate-pinnatifid, bipinnatepinnatifid proximally, basal pairs somewhat exaggerated basiscopically; pinna midveins dark to pinna tips; adaxial surfaces glabrous; abaxial surfaces white-farinose; sori at vein tips, laminar margins curved with thin, strongly modified, 1–1.5 mm wide false indusia; sporangia 64-spored; spores dark brown; 2n⫽60, 120 (Old World).

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blades under 30 cm (but not always), “tripinnate segments in lowest pair of pinnae” (none are tripinnate; the blades are exaggerated basiscopically in the basal pair of pinnae, and thus bipinnate-pinnatifid proximally. He claimed their chemistry is distinct, and the spores slightly different. We believe it is better to retain the name as C. farinosa until more careful detailed studies are made. The three species of Cheilanthes with farinose fronds in Mexico are sometimes placed in the genus Aleuritopteris, though molecular studies (Gastony & Rollo, 1998) suggest that C. aurea and C. aurantiaca, along with C. leucopoda, may belong in Notholaena.

24. CHEILANTHES FEEI Cheilanthes feei T. Moore, Index Fil. xxxviii. 1857. Figs. 83D–G. Myriopteris gracilis Fe´e, Me´m. Foug. 5: 150, t. 29, f. 6. 1852, non Cheilanthes gracilis. Type. U.S.A. [Missouri:] Hillsboro, Riehl 529 (RB; frag. US).

Distribution. Moist, wooded, rocky slopes; 1450–3100 m. Mexico; Guat, Hond, CR, Pan; Col, Peru; Africa, Asia, Pacific.

Rhizomes compact, ascending, 2–3 mm diam.; rhizome scales linear-lanceolate, some concolorous orange-brown, most bicolorous with blackish central stripe, 3–4 mm long, entire; fronds to 20 cm long, clumped; stipes 1⁄3–1⁄2 the frond length, dark brown to black, lustrous, terete, with lax to patent white to tan hairs, 1 mm long; blades narrowly oblong to lanceolate, bipinnate-pinnatifid to tripinnate, 1–3 cm wide; pinnae 3–10 pairs, equilateral, narrowly to broadly deltate; ultimate segments bead-like, largest 1–3 mm, round to slightly oblong; adaxial surfaces sparsely hirsute to glabrescent; abaxial surfaces densely villous; laminar margins curved but not differentiated, sori continuous around segment margins; sporangia 32-spored; spores brown; n⫽2n⫽90, apogamous (USA).

Selected Specimens Examined. Ags (McVaugh & Koelz 229, MICH, NY). Chih (Correll & Gentry 23025, NY). Chis (Matuda 1700, NY). DF (Lyonnet 1636, NY). Dgo (Breedlove 18833, NY). Gro (Lorea 1698, FCME). Hgo (Mickel 635, NY). Jal (Iltis 29611, NY). Me´x (Lott & Wendt P-97, NY). Mich (Hinton 12363, NY). Mor (Fisher 35420, NY). Nay (Gonza´lez 845, MEXU). Oax (Mickel 1265, NY). Pue (Arse`ne 2184, MEXU). Qro (Carranza 2839, MEXU). Sin (Pfeifer & Skog 3067, NY). SLP (Palmer 178, NY). Ver (Bourgeau 3058, NY). Zac (Rzedowski & McVaugh 949, NY). Unverified, Doubtful, or Mistaken Reports. Tam (reported by Lof, f. 27, 1980, but not verified).

This species closely resembles Notholaena candida, but C. farinosa has well developed false indusia, which are absent in N. candida. Saiki (1984a, b, c) distinguished many species from C. farinosa across its range around the world (40 species in the complex, which included much of what is currently construed as Notholaena). He applied the names Aleuritopteris mexicana and A. chihuahuaensis to the Mexican material here called C. farinosa, but distinguished these on questionable characters, such as indusia continuous versus separate (apparently a function of frond size),

Distribution. Calcareous cliffs and ledges, usually on limestone or sandstone; 1200–1400 m. W USA and Can; Mexico. Selected Specimens Examined. Chih (Correll 22635, NY, 23314, UC, US; Correll & Gentry 22718, UC, US, 23222, US; Knobloch 229, US, 409, MSC, US; Worthington 14566, NY). Coah (McCulloch 63-71-MC, MSC; Palmer 432, NY, 734, US).

194

CHEILANTHES

Cheilanthes feei is often confused with C. parryi, from which it is distinguished by having thinner, sparser pubescence and smaller, 1–3 mm, round, bead-like ultimate segments (vs. 3–5 mm, oblong segments) and tripinnate blade bases (vs. bipinnatepinnatifid). From C. lendigera, C. feei differs by the indusia being weakly differentiated (vs. strongly so) and generally having smaller fronds to 20 cm (vs. to 37 cm).

25. CHEILANTHES FENDLERI Cheilanthes fendleri Hook., Sp. Fil. 2: 103, t. 107b. 1852. Figs. 78M–P. Myriopteris fendleri (Hook.) E. Fourn., Mexic. Pl. 1: 125. 1872. Type. U.S.A. New Mexico: Fendler 1015, in 1847 (presumably K; isotype GH!, frag. US!, NY!)

Rhizomes short- to long-creeping, horizontal, 1–3 mm diam.; rhizome scales lanceolate, often deciduous with age, mostly concolorous brown, lustrous, ca. 2 mm long, entire; fronds to 30 cm long, approximate to distant; stipes ca. 1⁄2 the frond length, dark reddish brown, terete, lustrous, with scattered, loosely imbricate, linear, whitish scales 1–2 mm long; blades lanceolate to narrowly oblong, 3–4-pinnate, 1.5–5 cm wide; pinnae 10–14 pairs, equilateral; costae abaxially with scales lanceolate to deltate-lanceolate, truncate to subcordate, bases without overlapping lobes, strongly imbricate, often concealing segments, entire to denticulate, lacking cilia; ultimate segments bead-like, the largest 1.5–3 mm; adaxial surfaces glabrous; abaxial surfaces glabrous or with a few scales near segment bases; laminar margins recurved, but not differentiated; sporangia 64-spored; spores brown; 2n⫽60 (USA).

26. CHEILANTHES FIMBRIATA Cheilanthes fimbriata (A. R. Sm.) Mickel & Beitel, Mem. New York Bot. Gard. 46: 112. 1988. Figs. 72L–O. Cheilanthes microphylla (Sw.) Sw. var. fimbriata A. R. Sm., Amer. Fern J. 70: 19, f. 9–10. 1980. Type. Mexico. Chiapas: Mpio. Frontera Comalapa, 6–8 km E of Frontera Comalapa, Breedlove 39018 (DS!).

Rhizomes compact to short-creeping, horizontal, 1–2 mm diam.; rhizome scales linear, concolorous tan to orange, ca. 3 mm long, entire; fronds 8–32 cm long, clumped to approximate; stipes ca. 1⁄3 the frond length, atropurpureous to black, terete, clothed with appressed tan hairs and very narrow scales (mostly only 2 cells wide); blades lanceolate, bipinnate, rarely to bipinnatepinnatifid, subcoriaceous; segments elliptic, often auriculate to shallowly lobed; pinnae 10–17 pairs, equilateral; adaxial surfaces with lax white segmented hairs 0.3–0.5 mm long, sparse in center, more abundant near margins; abaxial surfaces with short uniform hairs 0.3–0.5 mm long and scattered linear scales on axes; sori at vein ends, laminar margins curved, fimbriate, somewhat differentiated into 0.3 mm wide thin, pale, ciliate, false indusia; spores tan.

Distribution. On limestone at lower elevation of Atlantic slope; 0–1000 m. Mexico; Guat, Hond; Cuba, Hisp. Selected Specimens Examined. Chis (Breedlove 26976, DS, 36556, DS; Mu¨nch 184, DS). Oax (Conzatti 3774, US; Hallberg 1656, NY; Mickel 1483, NY; Rose & Rose 11350, NY, US). QR (Montgomery 87-16, NY). Tam (Palmer 18, NY). Yuc (Gaumer 534, NY).

Distribution. Rocky slopes and ledges in pine-oak and pine-fir forests; found on a variety of acidic and mildly basic substrates; 1850–2400 m. Sw USA; Mexico. Specimens Examined. BCN (McGill & Pinkava P12330, ASC; Moran 15482, US, 24571, SD; Moran & Thorne 14204, ARIZ, SD, UC; Wiggins & Demaree 4956, DS, NY, 5005, DS; Yatskievych 82-191, ARIZ). Son (White 4213, MICH).

Young, sterile plants of Cheilanthes fendleri are occasionally misidentified as C. pringlei; they are distinguished from that species by having rachises that are terete (not adaxially grooved).

Unverified, Doubtful, or Mistaken Reports. Camp (Zamora 4855, UACAM, XAL, cited by Palacios Rios, 1997, but not verified). Ver (reported by Mickel and Beitel, 1988, not verified, but likely)

Cheilanthes fimbriata closely resembles C. microphylla, but has well modified, long-ciliate indusia.

27. CHEILANTHES HINTONIORUM Cheilanthes hintoniorum Mendenh. & Nesom, Sida 14: 551, f. 1. 1991. Type. Mexico. Nuevo Leo´n: Mpio. Galeana, 10 km NE

CHEILANTHES JAMAICENSIS

of Picotos, Hinton et al. 18765 (TEX!; isotypes MEXU, NY, UC). Figs. 76A–C. Rhizomes compact, horizontal, 2–3 mm diam.; rhizome scales bristle-like, 5–7 mm long, mostly only 1–2 cells wide, 3–4 cells wide at base, black, heavily indurate, entire; fronds to 17 cm long, clumped; stipes 1⁄3–2⁄5 the frond length, atropurpureous, terete, glabrous; blades pentagonal, 2–3-pinnate; pinnae 4–6 pairs, basiscopic pinnules of basal pinnae exaggerated; segments linear, mostly 10–15 ⫻ 1–1.5 mm, obtuse, apical ones mostly 12–20 (–30) mm; adaxial and abaxial surfaces glabrous, but with pebbly surface; laminar margins recurved, but not strongly modified, into 0.5–0.7 mm wide false indusia, margins minutely shortpapillate, false indusia not decurrent, vein endings swollen and usually evident; spores brown.

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tered linear-lanceolate scales 1–2 mm long, and dimorphic hairs, abaxially hairs 1 mm long, adaxially hairs 0.1 mm long, tortuous, appressed; blades linear-oblong to lanceolate, pinnate-pinnatifid to bipinnate, 1–4 cm wide; pinnae 10–15 pairs, equilateral; abaxial scales of costae lanceolate, truncate to subcordate at bases, loosely imbricate, not concealing ultimate segments, entire to slightly toothed; ultimate segments narrowly elliptic to elongatedeltate, not bead-like, the largest 3–5 mm long; adaxial surfaces scabrous with stiff, pustulate hairs; abaxial surfaces with nonpustulate hairs; laminar margins curved, not differentiated, sori continuous along segment margins; sporangia 64-spored; spores tan; 2n⫽58, 116 (NL, USA).

Distribution. Rocky slopes and ledges, usually on limestone; 100–1750 m. USA (Okla, Tex); Mexico. Distribution. Gypsum cliffs and hillsides; 965–2000 m. Mexico. Specimens Examined. NL (Cowan 3753, UC; Nesom 4308, TEX; Hinton et al. 18639, 18765, 19315, TEX; 20157, IEB, TEX, 22695, ANSM, NY, TEX; Zamudio et al. 6215, IEB, NY; Zamudio & Lux 6162, IEB, NY).

Cheilanthes hintoniorum is related to C. angustifolia and C. complanata, but has smaller plant size, hair-like rhizome scales, and occurs on gypsum (vs. neutral substrates). It is close to C. purpusii and C. pellaeopsis, but is less dissected, mostly bipinnate with linear segments, and its indusia are not decurrent. Cheilanthes apiacea is also close but has longer segments (mostly 10–15 mm long, terminal ones 20–25 mm).

28. CHEILANTHES HORRIDULA Cheilanthes horridula Maxon, Amer. Fern. J. 8: 94. 1918. Figs. 78A–D. Cheilanthes aspera Hook., Sp. Fil. 2: 111. 1852, hom. illeg., non Kaulf., 1831. Pellaea aspera Hook. & Baker, Syn. Fil. 148. 1867. Type. U.S.A. Texas: Turkey Creek, 1849, Wright 824 (K!; isotypes GH!, NY!, US!).

Rhizomes compact, horizontal, 2–3 mm diam.; rhizome scales linear-lanceolate, concolorous orange-brown or weakly bicolorous, ca. 3 mm long, entire; fronds 10–35 cm long, clumped; stipes 1 ⁄4–1⁄3 the frond length, black to dark brown, terete, with scat-

Selected Specimens Examined. Chih (Knobloch 625, MEXU; Pringle 447, NY). Coah (Palmer 1422, NY). Dgo (Dı´az Luna 7423, UC; Johnston 7793, UC). Gto (Carranza G. 3721, IEB). NL (Aguirre C. 857, ENCB). SLP (Orcutt 5392, DS). Tam (Johnston 4884, MEXU). Zac (Johnston et al. 11528B, LL, TEX).

The scabrous, pustulate hairs of Cheilanthes horridula make it one of the more distinctive species of Cheilanthes in Mexico. The species includes two sexually reproducing cytotypes in North America (Windham & Rabe in FNA Ed. Comm., 1993).

29. CHEILANTHES JAMAICENSIS Cheilanthes jamaicensis Maxon, Contr. U.S. Natl. Herb. 24: 51. 1922. Type. Jamaica. Below Cinchona, Harris 12905 (US; isotypes GH!, NY!). Figs. 82O–S. Rhizomes short-creeping, horizontal, ca. 2 mm diam.; rhizome scales linear-lanceolate, bicolorous with black central stripe and tan margins, 2–3 mm long, entire; fronds to 35 cm long, clumped; stipes 1⁄3–2⁄3 the frond length, castaneous, terete, with appressed, whitish, linear scales, 2–3 mm long, rachises with similar scales; blades lanceolate, tripinnate-pinnatifid proximally, tripinnate distally; ultimate segments distant, constricted at bases, narrowly elliptic, mostly 1–2 ⫻ 1 mm (or less); costae and costules abaxially with numerous narrow scales like those of rachises and stipes; adaxial surfaces with scattered hairs; abaxial surfaces densely to

196

CHEILANTHES

sparsely villous, hairs white, usually not obscuring laminar tissue; laminar margins recurved, slightly differentiated, whitish; sporangia 32-spored; spores brown.

of segment tips and lobes recurved and modified (occasionally only slightly so) into 0.5 mm wide entire false indusia; sporangia 32spored; spores tan to gray.

Distribution. On limestone in pine-oak forests; 1000–2400 m. Mexico; Jam, Hisp.

Distribution. Dry, rocky slopes; 600–2700 m. USA (Tex); Mexico; Guat, Hond, Salv, CR; Col, Ven.

Selected Specimens Examined. Chis (Breedlove 40736, DS; Mu¨nch 45, DS). Coah (Muller 3252, UC, US; Pennell 17347, MEXU, NY, US). Hgo (Moore 1683, UC, US). NL (Kenoyer 1100, US; Pringle 2603, UC). SLP (Pray 3062, LAM, NY, US; Whiting 829, MEXU, US, 1078, MEXU, MICH, US).

Selected Specimens Examined. Ags (McVaugh 18424, NY). Chih (Pringle 826, NY). Chis (Breedlove 21973, NY). Col (Alava & Cook 1711, MEXU). DF (Lyonnet 237, IEB, NY). Dgo (Palmer 511, NY). Gro (Mexia 9023, NY). Gto (Correll & Correll 28821, MEXU). Hgo (Herna´ndez M. 3800, MEXU). Jal (Mickel 1366, NY). Me´x (Pringle 11264, NY). Mich (Rzedowski 37390, NY). Mor (Reyes 1147, MEXU). Nay (Breedlove 44522, NY). NL (Mueller & Mueller 969, MICH). Oax (Mickel 4949, NY). Pue (Purpus 3150, NY). Qro (Dı´az B. & Zamudio 4827, IEB, MEXU). SLP (Schaffner s.n., in 1879, NY). Sin (Correll & Correll 28849, MEXU). Son (Rose 12861, NY). Tam (Kemp s.n., Jan 1902, NY). Tlax (Weber 706, ENCB). Ver (Riba 614, MEXU). Zac (Taylor & Taylor 6027, NY).

Unverified, Doubtful, or Mistaken Reports. Ver (reported by Smith, 1981, but not verified).

Cheilanthes jamaicensis has linear, white, appressed scales 4– 8 cells wide (and few if any hairs) on costae, whereas C. tomentosa has a mixture of twisted orange-tan hairs and only sparse scales mostly 2–4 cells wide. Cheilanthes jamaicensis has an unusual disjunct distribution, occurring in Jamaica and in both northern and southern Mexico.

30. CHEILANTHES KAULFUSSII Cheilanthes kaulfussii Kunze, Linnaea 13: 145. 1839. Figs. 73A–C. Cheilanthes viscosa Link, Hort. Berol. 2: 43. 1833, hom. illeg., non Carmich., 1818. Type. Cult. at B, from Mexico (B?; BM spec. “cult. in Hort. Lips., dedit. Kunze, 1840.”) Cheilanthes glandulifera Liebm., Mexic. Bregn. 258 (reprint 106). 1849. Lectotype (chosen by Smith, 1981: 72). Mexico. Oaxaca: “Inter Talea et Hac. de Sta. Gertrudes,” Liebmann s.n. [Pl. Mex. 2354] (C!–2 sheets; isotype K!).

Rhizomes compact, horizontal to ascending, 2–4 mm diam.; rhizome scales linear-lanceolate, concolorous reddish black, heavily sclerotic, lustrous, straight, 5–6 mm long, entire with a thinner, brown, short hair-tip ending in a gland cell; fronds 10–45 cm long, clumped; stipes ca. 2⁄3 the frond length, slender, atropurpureous, terete, with abundant, patent, glandular hairs ca. 0.2 mm long; blades broadly deltate to pentagonal, the lowest pinnae with strongly exaggerated basiscopic pinnules, 3-pinnate to 3-pinnate-pinnatifid (to 4-pinnate-pinnatifid); rachises and costae grooved; all surfaces covered with glandular hairs; margins

Cheilanthes kaulfussii is distinguished by the finely dissected, broadly deltate to pentagonal (rarely lanceolate) blades with abundant, short, erect glandular hairs on the stipes and blades. It is one of the most widely distributed pteridophytes in Mexico, occurring in 24 states and Distrito Federal. Some specimens from Durango have as few as one sporangium per vein end, but this is possibly a function of smaller plant size.

31. CHEILANTHES LENDIGERA Cheilanthes lendigera (Cav.) Sw., Syn. Fil. 128, 328. 1806. Figs. 73F–H, 83Q. Pteris lendigera Cav., Descr. Pl. 268. 1801. Syntypes. Ecuador. “Junto a Guaranda en el Reino de Quito,” Ne´e s.n. (MA?); Mexico. Hidalgo: “Ixmiquilpan, Nueva Espan˜a,” Ne´e s.n. (MA?). Cheilanthes minor M. Martens & Galeotti, Me´m. Foug. Mexique 75, pl. 21, f.1. 1842. Cheilanthes lendigera (Cav.) Sw. β minor (M. Martens & Galeotti) T. Moore, Index Fil. 245. 1861. Type. Mexico. Oaxaca: Rı´o de Capulalpan, Galeotti 6464 (BR!, no number on sheet marked ‘TYPUS’ photo US!; isotype P!). Myriopteris villosa Fe´e, Me´m. Foug. 5: 149, t. 28, f. 3. 1852. Type. Mexico, Galeotti 6478 (P!–2 sheets, BR!). Cheilanthes lanuginosa M. Martens & Galeotti, Me´m. Foug. Mexique 75, pl. 20, f. 2.1842. Type. Mexico, “Pre`s de Mora´n,” Galeotti 6450 (BR!, photo US!; isotype BR!, P!). Cheilanthes frigida Linden ex T. Moore, Gard. Chron. 7: 72. 1857. Myriop-

CHEILANTHES LINDHEIMERI teris frigida (Linden ex T. Moore) J. Sm., Cat. Cult. Ferns 28. 1857. Type. Cult. material, hort. Veitch. 1835 (K, photo US). Pomatophytum pocillatum M. E. Jones, Contr. W. Bot. 16: 12. 1930. Type. U.S.A. Arizona: Cochise Co., Ramsey Canyon, Huachuca Mountains, Jones 2469 (POM; isotypes GH!, NY!, UC, US).

Rhizomes long-creeping, horizontal, ca. 3 mm diam.; rhizome scales linear-lanceolate, mainly concolorous tan to orange, becoming only weakly bicolorous or dark brown concolorous with age; fronds to 37 cm long, spaced; stipes generally 1⁄2–3⁄5 the frond length, dark castaneous, terete, lustrous, with long soft orange hairs 1.5–3.5 mm long; blades deltate, tripinnate-pinnatifid to quadripinnate, the segments round, bead-like, generally ca. 1 mm across, discrete; pinnae 10–20 pairs, equilateral, narrowly deltate; adaxial surfaces glabrous; axes and abaxial surfaces with long orange hairs; sori at vein ends, laminar margins curved and strongly modified into 0.2–0.3 mm wide entire false indusia, nearly covering abaxial surfaces of segments, with sparse hairs at the indusial-laminar interface; spores dark brown; 2n⫽120 (USA).

Distribution. Moist oak woods, basaltic cliffs; (1300–)2000– 3270 m. USA (Ariz, Tex); Mexico; Guat, Hond, Nic, CR, Pan; Hisp; Col, Ven, Ec. Selected Specimens Examined. Chih (Pringle 828, NY). Chis (Breedlove 22085, NY). DF (Rusby 373, NY). Dgo (Breedlove 59128, NY). Gro (Lorea 3257, FCME). Gto (McVaugh 10405, US). Hgo (Coughlin 308, NY). Jal (Mexia 1725, NY). Me´x (Koch & Gonza´lez L. 861, NY). Mich (Pringle 3368, NY). Mor (Fisher 35413, NY). Oax (Mickel 3894, NY). Pue (Purpus 4018, NY). Qro (Rzedowski & McVaugh 432, NY). Sin (Gentry & Arguelles 18192, NY). Son (Fishbein 1744, ARIZ). Tlax (Sharp & Herna´ndez 44493, MEXU). Ver (Ventura A. 65, NY). Unverified, Doubtful, or Mistaken Reports. Ags (Siqueiros 4342, HUAA, and 14 other collections, cited by Siqueiros-Delgado & Gonza´lez-Adame, 2004, but not verified).

Cheilanthes lendigera is distinct from other members of this complex by its relatively broad false indusia with scattered hairs on the indusial-lamina, margin interface, and frond indument of hairs only 1–2 cells wide. Cheilanthes mexicana differs in having the indusia virtually absent, and Cheilanthes marsupianthes has linear stipe scales 4–5 cells wide and lacks hairs on the indusial-laminar margin interface. Reeves (1979) suggested that C. lendigera is a fertile allotetraploid of C. marsupianthes ⫻ mexicana.

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32. CHEILANTHES LEUCOPODA Cheilanthes leucopoda Link, Fil. Spec. 66. 1841. Type. Mexico. Hort. berol. (B). Figs. 83A–C. Rhizomes compact, horizontal, 4–10 mm diam.; rhizome scales linear, concolorous, dull orange-tan, entire; fronds to 35 cm long, clumped; stipes ca. 2⁄3 the frond length, stramineous, grooved distally, with a few scales at base, also with abundant patent hairs 0.5–1 mm long; blades broadly pentagonal, 4-pinnate, 5–15 cm wide; rachises grooved adaxially; basal pinnae strongly inequilateral, the basal basiscopic pinnules greatly exaggerated, costae green to stramineous; ultimate segments oblong to lanceolate, not bead-like, the largest 3–5 mm; adaxial and abaxial surfaces with scattered stiff hairs similar to those on the axes; segment tips curved and weakly differentiated as false indusia, sori discontinuous, concentrated on apical and lateral lobes; sporangia 32spored; spores dark brown; 2n⫽60 (NL, USA).

Distribution. Rocky slopes and ledges, apparently confined to limestone; (150 m in Tam) 1000–2250 m. USA (Tex); Mexico. Selected Specimens Examined. Chih (Pringle 442, NY). Coah (Stanford et al. 82, NY). Dgo (Iltis & Lasseigne 135, NY). Gto (Ventura & Lo´pez 8998, MEXU). Hgo (Chase 7234, NY). NL (Hinton et al. 21182, ARIZ; Pringle s.n., 20 Jun 1888, NY). Qro (Rzedowski 46443, MEXU). SLP (Purpus 4879, NY). Tam (Stanford et al. 2123, NY).

Cheilanthes leucopoda has 32 spores per sporangium but is a sexual diploid (Windham & Rabe in FNA Ed. Comm., 1993). This species is distinct in its broadly deltate hairy blades with stramineous axes. Molecular evidence (Cranfill, unpubl.) suggests a relationship with C. aurea in Notholaena, though it lacks the farina of that species.

33. CHEILANTHES LINDHEIMERI Cheilanthes lindheimeri Hook., Sp. Fil. 2: 101, t. 107a. 1852. Figs. 80P–V. Myriopteris lindheimeri (Hook.) J. Sm., Bot. Voy. Herald 340. 1854. Lectotype (designated here). U.S.A. Western Texas: Lindheimer 744 (K; isolectotypes GH!, P!–2 sheets, SD!, US!, YU!). Cheilanthes albida Baker, Ann. Bot. (Oxford) 5: 212. 1891. Type. Mexico.

198

CHEILANTHES

San Luis Potosı´: region of San Luis Potosı´, 1878, Parry & Palmer 999 (K; isotypes NY!–3 sheets, SD!, US!, frag. NY!).

Rhizomes long-creeping, horizontal, 1–1.5 mm diam.; rhizome scales lanceolate, concolorous tan to brown, thin to sclerotic, 1–2 mm long, entire; fronds to 25 cm long, 3–20 cm apart on the rhizomes; stipes 1⁄3–1⁄2 the frond length, blackish brown, lustrous, terete, with sparse, linear-lanceolate, tan, thin, subentire scales with occasional hair-teeth and hairs at base and some reduced nearly hair-like scales; blades narrowly lanceolate, not especially narrowed proximally, 3–4-pinnate with bead-like segments; pinnae 10–14 pairs, equilateral; adaxial surfaces glabrous but with loose, wool-like mat of white hairs 1–1.5 mm long extending beyond segment margins abaxially; abaxial surfaces with a dense covering of scales, those on the midribs linear-lanceolate, castaneous at bases, on the segments lanceolate, with long marginal cilia throughout the scale length, some scales reduced nearly to stellate hairs; laminar margins recurved but not strongly differentiated, obscured by scales and hairs; spores dark brown; n⫽2n⫽90, apogamous (USA).

what resinous, entire; fronds 12–30 cm long, clumped; stipes 1⁄3–2⁄5 the frond length, atropurpureous to black, terete, with abundant straight, patent, white to tan, gland-tipped hairs 0.5–2 (mostly ca. 1) mm long; blades narrowly deltate to lanceolate, tripinnatepinnatifid to quadripinnate; pinnae mostly opposite; segments mostly broadly adnate, nearly equilateral; adaxial surfaces densely clothed with nearly straight, somewhat appressed white hairs 1–3 mm long; abaxial surfaces densely clothed with flexuous white hairs; sori at vein tips, margin slightly or not curved, unmodified into false indusia; spores dark brown. Distribution. Terrestrial or epipetric on rocky slopes, usually in oak forests; (600–)1400–1900 m. Mexico. Key to the Mexican Varieties of Cheilanthes longipila 1. Adaxial segment hairs mostly 1–3 mm long, abaxially ca. 1 mm, flexuous; stipe hairs 0.5–2 mm; NL to Jal and Oaxaca. .......................................................................... 35a. var. longipila. 1. Adaxial segment hairs ca. 0.5–1 mm long, abaxially ca. 0.5 mm, erect, straight; stipe hairs 0.2–0.5(–1) mm; Gro. ............................................................................... 35b. var. brevipila.

34a. Cheilanthes longipila var. longipila Figs. 73D, E, 87E–H. Differing from var. brevipila by its longer blade hairs and more widespread distribution; 2n⫽60 (NL).

Distribution. Among basaltic rocks in matorral or grasslands; 1000–2300 m. USA (Ariz, N Mex, Tex); Mexico. Selected Specimens Examined. Ags (McVaugh 18310, NY). BCS (Moran 11696, ENCB). Chih (Pringle 460, NY). Dgo (Palmer 510, NY, UC). Gto (Ventura & Lo´pez 7080, MEXU). Hgo (Sa´nchez Mejorada 716, MEXU). Jal (McVaugh 16758, NY). Mich (Arse`ne 2399, MEXU). Qro (Rose 11178, NY). SLP (Schaffner 39, NY). Son (Lloyd 492, NY). Zac (Wiggins 13209, DS, MEXU, SD).

Cheilanthes lindheimeri is distinct by its long-creeping rhizome with broad, concolorous scales and the adaxial blade surfaces appearing to be densely covered with fine curly hairs, though these are actually hairs from margins of abaxial costal scales that extend beyond the segment margins.

34. CHEILANTHES LONGIPILA Cheilanthes longipila Baker, Ann. Bot. (Oxford) 5: 211. 1891. Type. Mexico. San Luis Potosı´, Parry & Palmer 989 (K!, frag. US!). Rhizomes compact, ascending, ca. 2 mm diam.; rhizome scales lanceolate, concolorous tan to orange-tan, 3–5 mm long, some-

Distribution. Dry, rocky slopes in oak forests; (600–)1450– 1900 m. Mexico. Selected Specimens Examined. Gro (Hinton et al. 14378, NY). Jal (Pringle 1863, NY; Palmer 78, NY; Pringle 5409, 2768, MEXU). Me´x (Sa´nchez Mejorada 812, MEXU). NL (Correll & Johnston 19810, ASC, LL, NY). Oax (Mickel 1669, 3912, 6317, NY). Unverified, Doubtful, or Mistaken Reports. Ags (reported by de la Cerda Lemus & Siqueiros Delgado, 1985, but not verified).

Var. longipila is distinct by the long rigid patent hairs and lanceolate to narrowly deltate blades. See C. maxoniana for comparison with that species. 34b. Cheilanthes longipila var. brevipila Mickel, var. nov. Type. Mexico. Guerrero: 2 km al SE de Amatitla´n, 1600 m, 13 Aug 1994, Soto 1052 (holotype NY!; isotype FCME!). Figs. 84N–Q, 87J–M.

CHEILANTHES LOZANOI A var. longipila pinnis alternatis et pilis laminarum brevioribus differt.

Rhizomes compact, ascending, 2–3 mm diam.; rhizome scales lanceolate, dull tan to orange, 3–4 mm long, entire; fronds to 30 cm long, clumped; stipes 1⁄3 the frond length, reddish brown, lustrous, terete, with patent glandular hairs mostly 0.2–0.5 mm long, some to 1 mm long; blades oblong to lanceolate, 2–3pinnate-pinnatifid; pinnae alternate above the basal pair, nearly equilateral; rachises flattened, costae grooved; adaxial blade surfaces with scattered white Ⳳ appressed hairs ca. 1 mm long; abaxial blade surfaces with hairs ca. 0.5 mm, erect; sori on segment lobes, laminar margins curved, but not differentiated, with marginal hairs; spores dark brown.

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laminar margins slightly curved and only somewhat modified as false indusia 0.1–0.3 mm wide, margins entire; spores tan. Distribution. Terrestrial or epipetric on dry banks and slopes; 0–1900 m. Mexico. Cheilanthes lozanoi is most closely related to C. skinneri as evidenced by the linear, bicolorous rhizome scales, chartaceous blade texture, evident veins, dull stipes and rachises, and similar scales and hairs on rachises. Key to the Mexican Varieties of Cheilanthes lozanoi 1. Blades pinnate to pinnate-pinnatifid, with 0–4 pairs of lobes with at most a single basal pair of free pinnules, terminal pinna segment distinctly larger than other pinna segments, triangular; low to middle elevations: 0– 1250(–1900) m). .............................................................. 35a. var. lozanoi. 1. Blades pinnate-pinnatifid to bipinnate (to rarely tripinnate at base); proximal 2–7 pinna pairs pinnate with 2–4 basal pairs of free pinnules, terminal pinna segment pinnatifid, similar to other segments; middle elevations: (1050–)1450–1700 m. .............................. 35b. var. seemannii.

35a. Cheilanthes lozanoi var. lozanoi

Figs. 81C–E, G.

Pinnae entire or with 1–2 pairs of lobes or single pair of basal free pinnules, terminal pinna segment triangular, distinctly larger than other pinna segments. Distribution. Tropical deciduous forest, oak woods, rocky slopes with volcanic soil; 1550–1600 m. Mexico. Specimens Examined. Gro (Antonio 339, FCME, NY; Calzada 16172, NY).

This taxon is similar to C. longipila var. longipila, but has shorter, more variable hairs on all frond surfaces.

35. CHEILANTHES LOZANOI Cheilanthes lozanoi (Maxon) R. M. Tryon & A. F. Tryon, Rhodora 83: 133. 1981. Pellaea lozanoi Maxon, Contr. U.S. Natl. Herb. 10: 500. 1908 [“lozanii”]. Type. Mexico. Guerrero: Near Iguala, Lozano s.n. [Pringle 13947] (US!).

Distribution. Dry, rocky roadbanks, streambanks, and shrubby slopes; 0–1250(–1900) m. Mexico.

Rhizomes short, compact, horizontal, 2–4 mm diam.; rhizome scales linear, 4–5 mm long, bicolorous, thickened, sclerotic, center reddish brown to black, margins thin, tan, erose-denticulate; fronds to 36 cm long, clumped; stipes ca. 1⁄3 the frond length, dull gray to grayish brown, plane adaxially with slight wings making shallow grooves in distal stipes and rachises, with scattered to dense hairs 2 mm long and linear scales; blades narrowly oblong, pinnate to bipinnate-pinnatifid, segments deltateacuminate to acute; laminae chartaceous, veins evident to obscure; adaxial surfaces glabrous or with scattered short, white, jointed hairs 0.1 mm or less long; abaxial surfaces with scattered to dense, white hairs 0.5–1 mm long, and narrow, pale brown to white, hair-like scales; sori marginal, elongate along vein tips,

Selected Specimens Examined. Chis (Purpus 6728, NY). Col (Morgan s.n., 18 Jan 1989, NY). DF (Rusby 368, NY). Dgo (Gentry 8308, UC). Gro (Hinton 9170, NY). Jal (Lott 540, NY). Me´x (Hinton 4318, LL, NY). Mich (McVaugh 17985, NY). Nay (Rodin 7015, NY). Oax (Mickel 5188, NY). Sin (Fryxell 3052, MEXU).

Cheilanthes lozanoi var. lozanoi is highly variable, and the limits of the varieties are not clearly defined. 35b. Cheilanthes lozanoi var. seemannii (Hook.) Mickel & Beitel, Mem. New York Bot. Gard. 46: 118. 1988. Figs. 81A, B, F, H, J. Pellaea seemannii Hook., Sp. Fil. 2: 141, t. 117B. 1858, non Cheilanthes seemannii Hook., Sp. Fil. 2: 85, t. 97A. 1858 [⬅ Adiantopsis seemannii (Hook.) Maxon]. Type. Mexico. Sinaloa: Mazatla´n, Seemann 1447 (K!, photo US!).

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CHEILANTHES

Similar to C. lozanoi var. lozanoi except: basal 2–7 pinnae pinnate with 2–4 basal pairs of free pinnules, pinnae each with pinnatifid terminal segment, similar to other segments; adaxial surfaces with scattered to dense white hairs and hair-like scales; 2n⫽60 (Son).

Distribution. Dry, rocky slopes, dry arroyos with cactus-acacia association, and dry thorn-scrub with a few oaks; (1050–)1450– 1700 m. Mexico.

Bonpland s.n. (P!, photos BM!, GH!; isotype B-Willd. 20126!). Figs. 85A–C, H, J, K, 86F, G. Allosorus ciliata C. Presl var. minor C. Presl, Reliq. Haenk. 1(1): 59. 1825. Type. Mexico. Haenke s.n. (PR or PRC). Cheilanthes hirsuta Link, Hort. Berol. 2: 40. 1833, non Mett., 1859. Type. No specimen cited, “hab. in Mexico,” cult. Berlin (B?, photos GH!, US!; isotype PH!), non Allosorus hirsutus C. Presl, 1825. Cheilanthes rufescens Link, Hort. Berol. 2: 39. 1833. Type. No specimen cited, cult. at B (B; isotype PH!). Cheilanthes crenulata Link, Hort. Berol. 2: 41. 1833. Type. No specimen cited, cult. at B (B; isotype PH!).

Rhizomes compact, horizontal, 2–3 mm diam.; rhizome scales linear-subulate, bicolorous, dark brown- to black-centered with brown to tan margins, lustrous, 3 mm long; fronds to 20(–35) cm long, clumped; stipes 1⁄2–2⁄3 the frond length, castaneous, lustrous, grooved, glabrous; blades ovate to deltate, 2–3-pinnatepinnatifid; segments obcuneate, obtuse; adaxial surfaces pebbly, but without papillae; abaxial surfaces glabrous; laminar margins recurved, strongly modified into 0.5 mm wide false indusia with long-fimbriate margins (hairs 0.3 mm long), otherwise glabrous, very long-decurrent to subtending segment; spores brown.

Selected Specimens Examined. BCS (Brandegee 663, NY). Chih (Palmer 69, NY, US). Dgo (Worthington 8975, NY). Gro (Martı´nez S. 1160, MEXU). Gto (Galva´n 2408B, ENCB). Jal (Palmer 300, NY). Me´x (Tejero-Dı´ez 3061, IZTA). Mich (Hinton 12077, NY). Mor (Rose 6864, NY). Oax (Mickel 6646, NY). Pue (Miranda s.n., Aug 1943, MEXU). Qro (Rose 9506, NY). SLP (Fisher s.n., 23 Aug 1937, NY). Sin (Fryxell 3052, NY). Son (Gentry et al. 19288, NY). Tam (Viereck 640, US). Tres Marı´as (Ferris 5706, DS). Unverified, Doubtful, or Mistaken Reports. Ags (Siqueiros 1564, HUAA, and five other collections, cited by Siqueiros-Delgado & Gonza´lez-Adame, 2004, but not verified). Chis (Breedlove 36811, DS, 417971, CAS; Purpus 6728, UC, US, all cited by Smith, 1981, but treated here under var. lozanoi).

Var. seemannii is the more common and widespread variety. Specimens from northwestern Mexico, especially the states of Baja California Sur, Sonora, and Sinaloa, have more remote and more pointed free segments. This seems to include the type of var. seemannii. The element with rounder segments may represent a distinct taxon. The two varieties of C. lozanoi need more study to determine if variation seen throughout Mexico (see key and descriptions) represents two or three species with hybrid intermediates, clinal variation in a single species, or one highly variable species (C. lozanoi). In this treatment, the elements are accepted on the varietal level because of the correlation of geographic and elevational separation with blade dissection, which seems to differ only in degree.

36. CHEILANTHES MARGINATA Cheilanthes marginata Kunth, Nov. Gen. Sp. 1: 22. 1815; 7: t. 669. 1825. Type. Ecuador. Quito, “prope Penipe,” Humboldt &

Distribution. Among rocks in pine-fir and pine-oak forests; 2000–4000 m. Mexico; Guat, Hond, Salv, CR, Pan; Col, Ven, Ec, Peru, Bol, n Arg. Selected Specimens Examined. Ags (McVaugh 18429, MICH). Chis (Breedlove 15267, NY). Col (Goldsmith 69, UC). DF (Rusby 366, NY). Gro (Martı´nez S. 5624, MEXU). Gto (Rzedowski 41350, IEB). Hgo (Medina-Cota & Barrios 3739, NY, TEX). Me´x (Fisher 35408, NY). Mich (Hinton 13497, NY, TEX). Mor (Pringle 15701, LL). NL (Hinton et al. 22153, TEX). Oax (Mickel 7432, NY). Pue (Ventura A. 1393, NY). Qro (Dı´az B. & Carranza 7366, IEB). SLP (Schaffner s.n., Oct 1876, NY p.p.). Ver (Seaton 278, NY).

Cheilanthes marginata is most closely related to C. pyramidalis and differs from that, as well as other members of the complex, in the non-spiculate, very long-decurrent, broad indusia with fimbriate margins. Mickel 4574 (NY) shows a few papillate hairs but not to the degree found in C. pyramidalis. See C. pyramidalis for further discussion.

CHEILANTHES MEMBRANACEA

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37. CHEILANTHES MARSUPIANTHES Cheilanthes marsupianthes (Fe´e) T. Reeves ex Mickel & A. R. Sm., comb. nov. Figs. 83M–P. Myriopteris marsupianthes Fe´e, Me´m. Foug. 5: 149, t. 12A. f. 1. 1852. Type. Mexico. Veracruz: Pic d’Orizaba, Galeotti 6256 (P! [6256bis]; isotypes BR!–3 sheets [⫹ 2 sheets 6256bis, BR!]).

Similar to C. lendigera except: stipes and rachises with scattered linear scales 4–5 cells wide, 0.2–0.4 mm wide, mixed with hairs, the blades tripinnate (vs. 3–4-pinnate), segments 1.5 mm diam. (vs. 1 mm), lacking hairs at the indusial/laminar interface; 2n⫽60 (Me´x). Distribution. Known only from the type, which lacks habitat data. Mexico. Cheilanthes maxoniana is very close to C. longipila but is distinct in the narrowly oblong blades (vs. broadly oblong to narrowly deltate), hair length 2 mm (vs. 1 mm), and hair density scattered (vs. dense).

39. CHEILANTHES MEMBRANACEA Cheilanthes membranacea (Davenp.) Maxon, Amer. Fern J. 8: 119. 1918. Figs. 71C, D, 75J–L. Distribution. Terrestrial or among rocks in pine or fir forests; 2400–3400 m. Mexico; Guat. Selected Specimens Examined. Me´x (Koch & Gonza´lez L. 861, CHAPA, MICH; Matuda 31197, UC, US; Ventura V. 1446, IEB, MEXU). Pue (Balls 4516, MICH, UC, US; Tenorio L. 15874, MEXU). Tlax (Reeves 5942, ASC). Ver (Galeotti 6256, BR; Martı´nez & Va´zquez 338, IEB; Nee 23113, UC; Nevling & Go´mez-Pompa 2038, GH, MEXU).

38. CHEILANTHES MAXONIANA Cheilanthes maxoniana Mickel, sp. nov. Type. Mexico. Tamaulipas: San Lucas, Viereck 76 (holotype US!). Figs. 87A–D. A C. longipila frondibus longioribus angusti-oblongis et pilis laminarum longioribus differt. (Dedicated to William Maxon, who first suggested the distinctness of this specimen.)

Rhizomes unknown; rhizome scales, judging from stipe bases, linear-lanceolate, orange-tan; fronds 28–40 cm long; stipes 2⁄5–1⁄2 the frond length, atropurpureous, terete, with abundant, patent, orange-tan, gland-tipped hairs 0.1–0.3 and some 1–1.5 mm long; blades narrowly oblong, bipinnate-pinnatifid to tripinnate; pinnae mostly subopposite, subequilateral; segments broadly adnate; both surfaces with scattered straight, orange-tan, spreading hairs 2 mm long, apparently not glandular; sporangia at vein tips, generally 1–3 sporangia covered by segment lobe, lacking differentiated false indusia; spores dark brown.

Pellaea membranacea Davenp., Bot. Gaz. 21: 262, t. 18, f. 5, 6. 1896. Type. Mexico. Oaxaca: Sierra de San Felipe, Pringle 5963 (GH!). Cheilanthes selinoides Mickel in McVaugh, Fl. Nov.-Gal. 17: 243. 1992. Type. Mexico. Jalisco: northeastern slopes of the Nevado de Colima, below Canoa de Leoncito, McVaugh 12869 (MICH!; isotype US!).

Rhizomes compact, horizontal to ascending, 5–8 mm diam.; rhizome scales linear-lanceolate, concolorous, castaneous, lustrous, 5–6 mm long, entire; fronds to 68 cm long, clumped; stipes ca. 2⁄3 the frond length, 1.5–3.5 mm diam., castaneous, grooved, lustrous, glabrous, tightly clustered; blades broadly deltate, quadripinnate-pinnatifid to quinquepinnate, the segments mostly elliptic, entire, discrete; adaxial and abaxial surfaces glabrous; laminar margins recurved, strongly modified into 0.8–1 mm wide false indusia, erose to short-fimbriate with hairs 0.1 mm long, very long-decurrent; spores tan.

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CHEILANTHES

Distribution. Pine-oak cloud forests; 2750–3300 m. Mexico; Guat, CR, Pan.

Distribution. Rocky slopes, cliffs, shaded, oaks; 1850–3250 m. Mexico; Guat.

Selected Specimens Examined. Jal (Boutin & Brandt 2306, NY). Oax (Hellwig 358, 359, NY; Mickel 1194, 7057, 4070, 7057, 7431, NY). Ver (Balls 4642, US).

Selected Specimens Examined. Chih (Correll 23284, NY). Coah (Wendt & Adamcewicz 521A, LL). DF (Schaffner 47, NY, P p.p., US). Dgo (Palmer 509a, US). Gto (Rzedowski 43738, ENCB, MEXU). Hgo (Arreguı´n 714, IEB). Me´x (Rzedowski 37147, IEB). Qro (Rzedowski & McVaugh 432, CAS, ENCB). SLP (Parry & Palmer 987, NY). Zac (Johnston et al. 10480K, LL).

Unverified, Doubtful, or Mistaken Reports. Chis (reported by Mickel & Beitel, 1988, but not verified). Mor (reported by Riba et al., 1996, but cited specimens are C. marginata and C. pyramidalis).

Cheilanthes membranacea is distinct in having wide, fimbriate, long-decurrent indusia and occurring at elevations over 2750 m. Cheilanthes selinoides is apparently a form of C. membranacea with entire, rather than erose to fimbriate, indusial margins. In all other characters, they seem to be identical to the former: the large ascending rhizomes, flexuous rachises, and very broad and long-decurrent indusia. Cheilanthes membranacea is distinct from C. marginata in the larger plant size, more massive rhizomes, and castaneous (vs. black) rhizome scales.

40. CHEILANTHES MEXICANA Cheilanthes mexicana Davenp., Bull. Torrey Bot. Club 15: 227. 1881. Type. Mexico. Chihuahua: Portrero Peak, Pringle 827 (GH!; isotypes BR!, DS!, NY!, P!–3 sheets, UC!, US!, YU!). Figs. 83H, J–L. Cheilanthes mexicana is similar to C. lendigera except: indusia much reduced, to ca. 0.1 mm wide, indusial/laminar interface glabrous, segments mostly ca. 1 mm diam., rachis indument of hairs 1–2(–4) cells wide; 2n⫽60 (Chih).

Some specimens of C. mexicana, e.g., Schaffner 47 (US), have a few scales.

41. CHEILANTHES MICKELII Cheilanthes mickelii T. Reeves, Brittonia 32: 502, f. 1–5. 1980. Type. Mexico. Oaxaca: Distrito Yautepec, 64 km W of Tehuantepec on Rte 190, Mickel 4210 (NY!; isotype UC!). Figs. 72E, F. Rhizomes short-creeping, compact, horizontal, ca. 1.5 mm diam.; rhizome scales linear, ca. 3 mm long, bicolorous with dark brown to black, sclerotic, lustrous streak, orange-tan margins, and hair-tips; fronds to 22 cm long, approximate; stipes ca. 1⁄3 the frond length, atropurpureous to dark-castaneous, lustrous, terete, clothed with long, jointed, tan hairs 3 mm long and sparse tan linear scales (to 4 cells wide); blades lanceolate or oblong, 2– 3-pinnate, proximal pinnae slightly reduced; adaxial surfaces with sparse, tan hairs; abaxial surfaces with scattered long, tan, curled hairs 1 mm long, laminae visible; laminar margins curved, somewhat modified, 0.3 mm wide; spores tan.

CHEILANTHES MYRIOPHYLLA

Distribution. Dry, rocky slopes and stream banks in light shade; 50–900 m. Mexico. Specimens Examined. Chis (Breedlove 30592, 36744, DS). Oax (Mickel 5194, 6068, NY; Salas M. 1848, NY). Ver (Palacios Rios 3435, XAL).

Cheilanthes mickelii is related to C. notholaenoides and C. cucullans, differing in the generally smaller size, short-creeping rhizomes, the presence of adaxial hairs, and longer abaxial hairs (ca. 1 mm). In addition, it differs from small specimens of C. notholaenoides in being chartaceous rather than coriaceous. The cited Chiapas collections were treated by Smith (1981) as Notholaena cinnamomea Baker, incorrectly.

42. CHEILANTHES MICROPHYLLA Cheilanthes microphylla (Sw.) Sw., Syn. Fil. 127. 1806. Figs. 78E–H. Adiantum microphyllum Sw., Prodr. 135. 1788. Type. Jamaica. Swartz s.n. (S!; isotype BM!). Cheilanthes pubescens Kunth, Nov. Gen. Sp. 1: 21. 1815. Type. Mexico. Veracruz: prope Xalapa, Humboldt & Bonpland s.n. (B-Willd. 20121).

Rhizomes short- to long-creeping, horizontal, 1–2 mm diam.; rhizome scales linear, concolorous orange-brown to weakly bicolorous with age; fronds to 30 cm long, approximate to 5 mm distant; stipes ca. 1⁄3 the frond length, castaneous to black, lustrous, terete, glabrate with scattered, fine, 1–3 mm golden hairs; distal stipes and rachises puberulent adaxially with 0.2–0.5 mm long hairs, hirsute abaxially with appressed, 1–3 mm long hairs; blades lanceolate, 2–3-pinnate, chartaceous; adaxial surfaces glabrous or with scattered 0.1 mm white hairs; abaxial surfaces with scattered to dense 0.1–0.2 mm white hairs along veins or rarely glabrous; laminar margins curved, modified into 0.3 mm wide false indusia, margins entire or with very short hairs; spores tan; n⫽2n⫽87 (SLP), 2n⫽116 (Jam, USA).

203

Distribution. Limestone outcrops; 100–2350 m. USA (Fl); Mexico; Guat, Bel, Hond; Bah, Gr & L Ant; Col, Ven, Ec, Peru, Bol. Selected Specimens Examined. Chih (Pringle 449, NY). Chis (Breedlove 2228, NY). Coah (Palmer 1419, NY). DF (Schaffner 99, NY). Dgo (Torres C. 3556, MEXU). Gro (Vasquez 85, NY). Hgo (Edwards 880, DS). Jal (McVaugh 24442, NY). Me´x (Rzedowski 37830, NY). Mich (Rzedowski 41767, IEB). Mor (Lyonnet 80, MEXU). NL (Correll & Johnston 19782, NY). Oax (Herna´ndez G. 924, NY). QR (Rzedowski 42943, MEXU). Qro (Servin 952, IEB, NY). SLP (Fisher 37116, US). Son (Hartman 262, NY). Tam (Hellwig 1207, NY). Ver (Purpus 2173, NY). Zac (Palmer 256, NY). Unverified, Doubtful, or Mistaken Reports. Gto (Rzedowski 39261, IEB, cited by Dı´az-Barriga & Palacios-Rios, 1992, but not verified).

Cheilanthes microphylla is most often confused with C. alabamensis, which can be distinguished by being more coriaceous, more nearly bipinnate, and the segments acute to acuminate. Cheilanthes aemula is in the same complex but has broadly deltate blades. Cheilanthes fimbriata has been included within C. microphylla but is distinct in its fimbriate indusial margins and distribution on the Atlantic slope. Cheilanthes notholaenoides and C. cucullans are closely allied, but are distinct from C. microphylla in having long, evenly distributed rachis hairs rather than dense short hairs concentrated adaxially.

43. CHEILANTHES MYRIOPHYLLA Cheilanthes myriophylla Desv., Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck. Gesammten Naturk. 5: 328. 1811. Type. Peru. s. coll. s.n. (P-Desv.!). Figs. 73J–P, 80A. Cheilanthes paleacea M. Martens & Galeotti, Me´m. Foug. Mexique 76, pl. 21, f. 2. 1842. Type. Mexico. Me´xico to Tampico, Mextitla´n, Cuesta Blanca, Galeotti 6429 (BR!; isotype BR!, frag. US!). Myriopteris intermedia E. Fourn., Bull. Soc. Bot. France 27: 328. 1880, hom. illeg., non Fe´e, 1852. Cheilanthes intermedia Baker, Ann. Bot. (Oxford) 5: 213. 1891. Cheilanthes fournieri C. Chr., Index Filic. 174. 1905. Type. Mexico. San Luis Potosı´: Schaffner 23 (P!).

Rhizomes compact, horizontal, 2–3 mm diam.; rhizome scales linear-lanceolate, bicolorous, with black to dark brown, lustrous,

204

CHEILANTHES

central streak and tan margins, ca. 5 mm long, entire; fronds to 45 cm long, clumped; stipes ca. 1⁄3 the frond length, castaneous to atropurpureous, terete, liberally clothed with appressed pale hairs 2 mm long and occasional linear scales 2 mm long; blades narrowly to ovate-lanceolate, 4-pinnate, the segments bead-like, 0.5 mm wide, discrete; adaxial surfaces glabrous or with scattered, fine, flexuous, white hairs; abaxial surfaces obscured by white to rusty brown ovate-lanceolate non-ciliate scales on axes, covering the segments, and overlying the scattered tan, flexuous hairs on the segments; laminar margins recurved, but only slightly modified into 0.1–0.3 mm wide false indusia; spores tan; 2n⫽174 (Peru), ca. 180 (Mor).

Rhizomes short-creeping, horizontal, 1–2 mm diam.; rhizome scales linear-lanceolate, concolorous dark brown to black, lustrous, ca. 3 mm long, with paler, ephemeral, kinky hair-tips; fronds to 30 cm long, approximate; stipes ca. 1⁄2 the frond length, castaneous to dark brown, terete, with abundant, lax, tangled, tan hairs; blades lanceolate to oblanceolate, bipinnate-pinnatifid to tripinnate, 1.5–5 cm wide; pinnae 8–17 pairs, equilateral to slightly inequilateral, narrowly deltate, proximal basiscopic and acroscopic pinnules often slightly enlarged (though occasionally smaller than adjacent pinnules); segments oblong to lanceolate, not bead-like, largest 3–5 mm long; adaxial surfaces tomentose; abaxial surfaces woolly with very fine hairs; laminar margins slightly curved but not differentiated, sori concentrated on apical and lateral lobes; sporangia 64- or 32-spored; spores dark brown to black; 2n⫽60 (USA).

Distribution. Dry, rocky slopes, basalt or limestone, rarely in moist oak woods; 900–3000 m. Mexico; Guat, Hond, Salv, Nic; Hisp; Col, Ven, Ec, Peru, Braz, Bol, Chile, Arg. Selected Specimens Examined. Ags (McVaugh 23709, NY). BCS (Carter 2046, DS). Chih (Pringle 829, NY). DF (Mexia 2665a, NY). Dgo (Palmer 552, NY). Gro (Lorea 593, ENCB). Gto (McVaugh 24248, NY). Hgo (Orcutt 3927, NY). Jal (Jones 27018, NY). Me´x (Ventura V. 464, NY). Mich (McVaugh 18163, NY). Mor (Ramirez R. 59, XAL). Nay (Breedlove 45562, CAS). Oax (Mexia 9118, NY). Pue (Dunn et al. 23333, NY). Qro (Sohmer 9260, NY). Sin (Breedlove 44894, ENCB). SLP (Parry & Palmer 987, NY). Son (Gentry et al. 19322, NY). Tlax (Sohns 615, MEXU). Ver (Seaton 287, NY). Zac (McVaugh 17759, NY).

Cheilanthes myriophylla resembles C. lendigera in its bead-like segments but is distinct by its dense covering of scales rather than hairs on the segments abaxially. Cheilanthes myriophylla is also similar to C. wootonii, C. covillei, and C. clevelandii in the bead-like segments and scaly blades, but the rhizomes of C. myriophylla are compact (vs. creeping) and usually with hairs on the adaxial blade surfaces.

44. CHEILANTHES NEWBERRYI Cheilanthes newberryi (D. C. Eaton) Domin, Biblioth. Bot. 20(85): 133. 1915. Figs. 84E–H. Notholaena newberryi D. C. Eaton, Bull. Torrey Bot. Club 4: 12. 1873. Syntypes. U.S.A. California: San Diego, Newberry s.n., Wood s.n.; Southern California: Temascal Range, Brewer s.n. (YU!).

Distribution. Rocky cliffs and slopes; usually on igneous substrates; 50–650 m. USA (Calif); Mexico. Selected Specimens Examined. BCN (Eastwood 12390, CAS, UC; Moran 21668, MEXU, 26503, NY, US; Raven et al. 12288, UC; Wiggins 4242, ARIZ, MICH, NY). Guad (Anthony 255, NY; Franceschi 40, UC; Newcombe 183, UC, US; Palmer 855, NY; Rose 16416, NY).

Cheilanthes newberryi is distinct by the dense mat of fine hairs and concolorous dark rhizome scales on the short-creeping rhizomes.

45. CHEILANTHES NOTHOLAENOIDES Cheilanthes notholaenoides (Desv.) Maxon ex Weath., Contr. Gray Herb. 114: 34. 1936. Figs. 72C, D, 87N. Pteris notholaenoides Desv., Me´m. Soc. Linn. Paris 6: 299. 1827. Type. Hispaniola. s. coll. s.n. (P!, photo GH). Cheilanthes aspidioides Fe´e, Me´m. Foug. 5: 157. 1852. Type. Mexico. Oaxaca: San Pedro Nolasco, Galeotti 6557 (P!; isotypes BR!–2 sheets).

Rhizomes short- to long-creeping, horizontal, ca. 2 mm diam.; rhizome scales linear-lanceolate, 4 mm long, concolorous, orangetan or becoming dark in center at base with age, margins entire, rarely the scales entirely black, sclerotic, with a few marginal hairs, some specimens with rhizome hairs showing among the scales; fronds to 40 cm long, distant; stipes 1⁄4–2⁄5 the frond length,

CHEILANTHES PELLAEOPSIS

black, lustrous, terete, with orange-tan to brown appressed filiform scales 1.5 mm long, mostly only 2 cells wide; blades lanceolate to narrowly deltate, 2-pinnate, subcoriaceous, the pinnules usually auriculate or each with a free acroscopic lobe; adaxial surfaces glabrous, bluish to whitish green, pebbly; abaxial surfaces with dense filiform scales 1–2 mm long, 2 cells wide on costae with sparse to scattered minute hairs 0.1 mm or less on segments, rarely glabrous; laminar margins somewhat curved, only slightly modified into 0.3 mm wide false indusia, slightly paler at margins, not decurrent, margins papillose; spores tan; 2n⫽174 (Oax).

205

46. CHEILANTHES PARRYI Cheilanthes parryi (D. C. Eaton) Domin, Biblioth. Bot. 20(85): 133. 1915. Figs. 84A–D. Notholaena parryi D. C. Eaton, Amer. Naturalist 9: 351. 1875. Type. U.S.A. South Utah, Parry 263 (YU!; isotypes GH!–4 sheets, US!).

Rhizomes compact, horizontal to ascending, 2–3 mm diam.; rhizome scales linear to linear-lanceolate, bicolorous with black center and orange-brown margins, some concolorous orangebrown at rhizome apices, 3–4 mm long; fronds to 18 cm long, clumped; stipes 1⁄3–1⁄2 the frond length, dark castaneous to black, lustrous, terete, with white Ⳳ patent hairs 1 mm long; blades narrowly lanceolate, tripinnate to tripinnate-pinnatifid; pinnae 5–8 pairs, deltate; adaxial surfaces tomentose; abaxial surfaces woolly, the hairs obscuring the laminar surfaces; laminar margins not curved or differentiated as false indusia, sori continuous on segment margins; sporangia 64-spored; spores black; 2n⫽60 (USA).

Distribution. Dry, rocky slopes, mostly limestone; 800–3150 m. USA (Tex); Mexico; Guat, Pan; Jam, Hisp; Ven, Peru, n Arg. Selected Specimens Examined. Chih (Pringle 449, NY). Chis (Breedlove 22391, NY). Coah (Palmer 368, NY). DF (Lyonnet 900, NY). Gro (Sharp 441392, MEXU). Gto (Rzedowski 41593, IEB). Hgo (Pringle 11263, NY). Jal (Iltis et al. 29610, NY). Me´x (Matuda 30553, MEXU). Mich (Carranza Gonza´lez 1035, IEB). Mor (Callejas 55, UAMIZ). NL (Taylor 83, NY). Oax (Mickel 3858, NY). Pue (Ugent & Flores C. 2535, NY). Qro (Rzedowski 44109, IEB). SLP (Parry & Palmer 981, NY). Tam (Stanford et al. 2618, NY). Ver (Ventura A. 318, NY). Zac (Stanford et al. 573, NY). Unverified, Doubtful, or Mistaken Reports. Ags (De la Cerda & Garcı´a 274, HUAA, and two other collections, cited by SiqueirosDelgado & Gonza´lez-Adame, 2004, but not verified). Yuc (reported by Mickel & Beitel, 1988, but not verified).

This species is confused with Cheilanthes microphylla, but the latter has white 0.2–0.5 mm abaxial laminar hairs and dense short hairs on the rachises adaxially and rachises abaxially glabrous or with only sparse hairs. Cheilanthes notholaenoides has uniform hairs adaxially and abaxially on the rachises, and the laminae are glabrous or with scattered jointed hairs 0.3–1 mm versus the short white hairs of C. microphylla. Cheilanthes notholaenoides is especially close to C. cucullans in the abundant stipe and rachis hairs and often long-ciliate rhizome scales, but the former is distinct in being only bipinnate (vs. bipinnate-pinnatifid) and coriaceous (vs. chartaceous). Not previously reported from the United States (Windham & Rabe in FNA Ed. Comm. 1993), a specimen from Texas (s. coll., s.n. “Ex herb. Bennett”) is at NY.

Distribution. Rock crevices, dry rocky granite or calcite slopes; 230–600 m. W USA (Ariz, Calif, Nev, Utah); Mexico. Selected Specimens Examined. BCN (Cota 599, US, 2374, DH; Moran 19245, ASC; Orcutt 286983, SD; Shreve 6857, MICH, US; Thorne 57804, UC; Wiggins 7610, DS, US). Son (Felger 85-727, ARIZ, SD; Felger 16896, 18680, ARIZ).

Cheilanthes parryi is characterized by abundant hairs on both laminar surfaces, tangled and long, surpassing the segments, and thus appearing like a cotton ball.

47. CHEILANTHES PELLAEOPSIS Cheilanthes pellaeopsis Mickel in McVaugh, Fl. Nov.-Gal. 17: 241. 1992. Type. Mexico. Jalisco: Mpio. Zapotitla´n, between Mexcalitos and El Chivero, northeast-facing rim of escarpment of Cerro Grande N of Los Picachos, Iltis et al. 29543 (WIS!, frag. NY!). Figs. 76G–J. Rhizomes compact, horizontal, ca. 5 mm diam.; rhizome scales linear-lanceolate, 4–5 mm long, 10–12 cells wide at base, concolorous black, lustrous, entire, often with brown marginal row

206

CHEILANTHES

of cells and brown tips; fronds to 30 cm long, clumped; stipes 1⁄2–2⁄3 the frond length, black, terete, glabrous; blades pentagonal, 3–4-pinnate, 12–15 cm wide; pinnae 8–10 pairs; segments linear, mostly 7–12 mm long, 1.1–1.3 mm wide, apical ones to 20 mm long; adaxial and abaxial surfaces glabrous, pebbly; laminar margins strongly recurved, weakly differentiated, the false indusia nearly covering the abaxial surfaces of the segments, white, firm, 0.3–0.5 mm wide, entire, papillose, slightly to strongly decurrent, with vein endings pronounced and sunken in pits, some pits with slight white farina; spores tan to light brown.

Rhizomes short-creeping, horizontal, 1 mm diam.; rhizome scales linear-lanceolate, concolorous orange-brown, weakly bicolorous with age, weakly and irregularly toothed to entire; fronds to 26(–30) cm, clumped; stipes 1⁄2 the frond length, castaneous to purplish brown, grooved, lustrous, with scattered, filiform, tortuous to straight, orange-brown scales, 1–3 mm, plus many reduced, often contorted and hair-tipped scales/hairs, 0.1– 0.3 mm; blades deltate to lanceolate, tripinnate-pinnatifid; rachises and rachillae with scales like those of stipes (but lesser axes with deltate-attenuate and tortuous scales); pinnae 5–11, equilateral except basal pair basiscopically exaggerated; adaxial surfaces green, glabrous; abaxial surfaces glabrous except for scales on dark axes; sori along segment tips only, margins curved, but not differentiated; spores tan. Distribution. Terrestrial or epipetric on rocky slopes among shrubs and in open woods; 0–1350 m. Mexico. Key to the Mexican Varieties of Cheilanthes peninsularis 1. Rachilla scales flexuous, denticulate; blades deltate, 3-pinnate-pinnatifid to 4-pinnate; pinna pairs ca. 6; BCS. ......................................................................... 48a. var. peninsularis. 1. Rachilla scales nearly straight, subentire; blades lanceolate, 2–3-pinnate; pinna pairs 7–11; Rev. ....... 48b. var. insularis.

Distribution. Terrestrial and among limestone rocks in pineoak forests; 1975–2300 m. Mexico. Specimens Examined. Gro (Anderson & Laskowski 4394, NY). Jal (only the type). Oax (Hunn OAX-998, MEXU). Tam (Diggs & Nee 2378, F; Webster & Webster 18, TEX).

The foliar axes of this species are black, similar to those of the Cheilanthes angustifolia complex, but the segments are linear with the margins strongly recurved to nearly covering the abaxial laminar surfaces. The vein endings are notably pronounced, with slight white farina. Cheilanthes pellaeopsis is closely allied to C. hintoniorum, but is larger with more pinnae pairs, farina, and broader rhizome scales. Cheilanthes pellaeopsis resembles C. decomposita in dissection and the swollen vein ends, dark axes, and black rhizome scales, but the latter has more strongly differentiated false indusia, non-papillate margins, non-decurrent indusia, broader segments (1.5–2.5 vs. 1.1–1.3 mm), indusia not covering the abaxial surfaces, black spores, more castaneous to purplish brown axes, and prominent veins. Cheilanthes pellaeopsis resembles C. purpusii, which differs in having segments mostly 3–7 mm long, apical ones 5–8(–12) (vs. segments 7–12, apical ones to 20 in C. pellaeopsis). Possibly it is an extreme form of C. purpusii. More collections are needed.

48. CHEILANTHES PENINSULARIS Cheilanthes peninsularis Maxon, Contr. U.S. Natl. Herb. 10: 496. 1908. Type. Mexico. Baja California: Cape Region, Nov 1902, Brandegee s.n. (US!, frag. NY!).

48a. Cheilanthes peninsularis var. peninsularis Figs. 79N–P. Differing from var. insularis in having flexuous, denticulate rachillae scales, deltate, tripinnate-pinnatifid to quadripinnate blades, and ca. 6 pinna pairs.

Distribution. Rocky open woods, often on shaded north slopes; 575–1350 m. Mexico. Selected Specimens Examined. BCS (Brandegee 660, GH, NY, UC; Carter & Moran 5507, UC; Jones 24559, CAS, NY; Moran 7093, DS; Porter 253, DS; Pray 1787, 1793, LAM; Wiggins 5465, CAS, 14763, 15377, DS; Wiggins et al. 495, DS, UC).

Two specimens, Pray 1760 and 1761, LAM, seem to be C. peninsularis var. peninsularis, but are labelled as coming from San Luis Potosı´, where this species is not known to occur. We suspect they were mislabelled since other specimens of C. peninsularis var. peninsularis were collected by Pray in Baja California. Cheilanthes peninsularis var. peninsularis is distinct from C.

CHEILANTHES PRINGLEI

pringlei in the former having slender purplish brown stipes with fewer (scattered) narrower scales (vs. stout reddish brown stipes with copious scales), fronds narrower ovate (vs. short, triangular or deltate-ovate), pinnae spaced (vs. close and imbricate), and rachises and rachillae with sparse yellow-brown scales (vs. with numerous broad whitish scales extending thickly even to distal parts of the pinnules, commonly obscuring the abaxial surfaces). 48b. Cheilanthes peninsularis var. insularis Weath., Amer. Fern J. 21: 25. 1931. Type. Mexico. Socorro Island, Mason 1616 (CAS!). Figs. 79Q–S. Differing from the type variety in having nearly straight, subentire rachilla scales, lanceolate, 2–3-pinnate blades, and 7–11 pinna pairs.

207

abaxially; segment lobe margins curved, but not or only slightly differentiated; spores tan. Distribution. Epipetric on dry rocky slopes; 350–2600 m. USA (Ariz); Mexico. Key to the Mexican Varieties of Cheilanthes pringlei 1. Frond scales abundant, imbricate, toothed; hairs inconspicuous; blades deltate; BCS, Chih, Son. ................................................................................. 49a. var. pringlei. 1. Frond scales scattered, not regularly imbricate; hairs conspicuous; blades oblong to deltate; Coah, SLP, Gto, Qro, Hgo, Pue, Oax, Zac. ................................. 49b. var. moncloviensis.

49a. Cheilanthes pringlei var. pringlei

Figs. 79E–H.

Cheilanthes sonorensis Goodd., Muhlenbergia 8: 93. 1912. Type. Mexico. Sonora: La Cie´nega, 1911, Goodding 942 (ARIZ).

Differing from var. moncloviensis in having abundant frond scales, inconspicuous hairs, and more westerly distribution; 2n⫽60 (USA).

Distribution. Rock crevices and volcanic talus, dry scrub; 0–1025 m. Mexico. Selected Specimens Examined. BCS (Brandegee s.n., 17 Feb 1889, NY, is somewhat intermediate). Rev (Cruz Cisneros 1603, ENCB; Dawson 13272, UC; Elmore C8, DS; Felger 15794, NY, SD, UC; Howell 8393, CAS, GH, NY; Lindsay 2836, SD; Mason 14593, CAS, MEXU, UC; Moran 5718, SD, 5928, DS, SD; Villareal 15, ENCB).

49. CHEILANTHES PRINGLEI Cheilanthes pringlei Davenp., Bull. Torrey Bot. Club 10: 61, t. 34. 1883. Type. U.S.A. Mountains of SE Arizona, 2 May 1883, Pringle s.n. (GH!; isotypes DS!, NY!–2 or 3 sheets, US!, YU!). Rhizomes long-creeping, horizontal, 1 mm diam.; rhizome scales linear-lanceolate, weakly bicolorous with orange-brown center and paler margins, 2.5–3 mm long, obscurely and irregularly toothed along margins; fronds to 13(–23) cm long, approximate to slightly spaced; stipes 1⁄2–2⁄3 the frond length, reddish brown, lustrous, grooved, with scattered to abundant (imbricate) linear whitish scales and reduced scales and contorted small hairs less than 0.1 mm; blades oblong-lanceolate or deltate to nearly pentagonal, 3–4-pinnate-pinnatifid; pinnae 3–6 pairs, basal pair exaggerated basiscopically; rachises and rachillae sparsely to densely scaly with white to tan, linear, irregularly toothed scales 1–2 mm long, otherwise glabrous adaxially and

Distribution. Dry, rocky slopes; 350–1100 m. USA (Ariz); Mexico. Selected Specimens Examined. BCS (Moran 18806, ASC). Chih (Bye 7731, GH, MEXU, MICH; Knobloch 881, DS, LL; Palmer 116, GH, MICH, NY). Son (Felger 20278, MEXU; Kennedy 7047, CAS, UC; Ownbey & Ownbey 1773, GH, NY, UC; Palmer 265, GH, NY, UC; Wiggins 6166, 7422, DS, MICH). Unverified, Doubtful, or Mistaken Reports. Qro (Zamudio 3528, IEB, cited by Dı´az-Barriga & Palacios-Rios, 1992, but not verified).

49b. Cheilanthes pringlei var. moncloviensis (Baker) Mickel, stat. nov. Figs. 79J–M. Cheilanthes moncloviensis, Baker, Ann. Bot. (Oxford) 5: 210. 1891. Type. Mexico. Coahuila: Soledad, 25 miles SW from Monclova, 1880, Palmer 1378 (K!; isotypes NY!, US!).

Differing from var. pringlei in having scattered frond scales, conspicuous hairs, and more easterly distribution.

208

CHEILANTHES

Distribution. Epipetric on basaltic rocks; 1650–2600 m. Mexico.

Distribution. Moist rocks in barrancas; pine-oak woods in arroyos, gypsum; 1550–2300 m. Mexico.

Selected Specimens Examined. Coah (only the type). Hgo (Diggs et al. 2028a, WIS; Orcutt 3930, DS). Gto (Pray 3075, LAM). Oax (Mickel 6644, NY). Pue (Purpus 4046, NY). Qro (Rose & Rose 11146, US). SLP (Parry & Palmer 990, NY p.p.; Siegler & Halstein DS-400, MEXU). Zac (Orcutt 3110, DS).

Specimens Examined. Gto (Ventura V. & Lopez P. 9059, IEB). Hgo (Kenoyer s.n., 14 Nov 1937, ARIZ, 586, F). NL (Breedlove 72040, CAS). Qro (Carranza G. 2862, 3168, IEB; Zamudio & Carranza s.n., 1–2 Apr 1997, IEB). SLP (only the type). Tam (Kemp s.n., Jan 1902, NY). Ver (Mu¨ller 3052bis, NY).

Most specimens have blades oblong-lanceolate, rather than distinctly deltate as in var. pringlei, although the Oaxaca and Coahuila specimens are somewhat deltate. The sparser scales in var. moncloviensis seem to be more diagnostic than the blade shape. They make the hairs more conspicuous, though they are present in var. pringlei too.

Cheilanthes purpusii is most closely allied to C. pellaeopsis and C. hintoniorum, differing in having smaller segments and only slightly decurrent indusia. Cheilanthes purpusii has lateral segments mostly 3–7 mm long (terminal ones 5–8(–12) mm), whereas C. pellaeopsis has lateral segments mostly 7–12 mm long (terminal ones to 20 mm), and those of C. hintoniorum are even longer. Cheilanthes purpusii resembles C. complanata, but the indusia of the former are papillate and the rhizome scales are black. Cheilanthes purpusii also resembles C. arizonica, but has indusia slightly decurrent and the stipes not grooved.

50. CHEILANTHES PURPUSII Cheilanthes purpusii T. Reeves, Rhodora 84: 293. 1982. Type. Mexico. San Luis Potosı´: Minas de San Rafael, Purpus 4881 (GH!; isotypes F, UC!). Figs. 76K–N. Rhizomes compact, horizontal, ca. 2 mm diam.; rhizome scales linear-subulate, 3–5 mm long, concolorous dark brown to reddish black, heavily indurate, lustrous, some with marginal row of brown cells, entire; fronds to 24 cm long, clumped; stipes 1⁄2–2⁄3 the frond length, atropurpureous, lustrous, terete, glabrous; blades deltate, 2–3-pinnate-pinnatifid, rachises and rachillae dark with pale green wings, deeply grooved, segments narrowly oblong to linear, mostly 3–6 mm long, terminal ones 5–8(–12) mm long, linear; adaxial surfaces pebbly; abaxial surfaces glabrous; laminar margins recurved, well differentiated into 0.5 mm wide, smooth, false indusia with short-papillate margins, slightly decurrent, covering the abaxial surfaces, vein ends swollen and conspicuous on adaxial surfaces; spores tan.

51. CHEILANTHES PYRAMIDALIS Cheilanthes pyramidalis Fe´e, Me´m. Foug. 7: 38, t. 25, f. 3.1857. Syntypes. Mexico. Valley of Mexico, Schaffner 88; Guatimalpan, Schaffner 304 (P!); San Agostı´n, Schaffner 305 (isosyntype K!). Figs. 70G–K, 86A–E. Pellaea angustifolia (Kunth) Baker var. elongata Rovirosa, Pteridogr. Sur Me´xico 130. 1909. Type. Mexico. Chiapas: Mesa de Coapilla, Rovirosa 1059 (isotype PH!, frag.US!, photo US!).

Rhizomes compact, horizontal, 2–3 mm diam.; rhizome scales linear-subulate, bicolorous, dark brown to black with narrow light brown entire margins, lustrous, ca. 3 mm long; fronds 15– 45 cm long, clumped; stipes 1⁄2–2⁄3 the frond length, castaneous, lustrous, grooved, glabrous; blades 3–4-pinnate-pinnatifid, ovate to deltate, rarely narrowly oblong; segments narrowly obcuneate to linear, obtuse, often strongly drooping; adaxial surfaces with 0.1–0.3 mm long white papillae at segment apices and on false indusia; abaxial surfaces glabrous; laminar margins recurved, strongly modified into 0.3–0.5 mm wide false indusia covering most of abaxial laminar surfaces, margins long-fimbriate (hairs 0.3 mm), generally long-decurrent along pinnule axes; spores grayish brown; n⫽2n⫽87, apogamous.

CHEILANTHES SKINNERI

209

52. CHEILANTHES SKINNERI Cheilanthes skinneri (Hook.) R. M. Tryon & A. F. Tryon, Rhodora 83: 133. 1981. Figs. 69A, B. Pellaea skinneri Hook., Sp. Fil. 2: 141, t. 118B. 1858. Type. Guatemala. Skinner s.n. (K!). Pellaea flavescens E. Fourn., Mexic. Pl. 1: 119. 1872, hom. illeg., non Fe´e, 1869. Pellaea fournieri Baker in Hooker & Baker, Syn. Fil. 486. 1874. Doryopteris fournieri (Baker) C. Chr., Index Filic. 244. 1905. Type. Mexico. [Morelos:] Cuernavaca, Bourgeau s.n. (P!).

Distribution. Grassy slopes and rocks in dry pine-oak woods to moist woods; 850–3000 m. Mexico; Guat, Hond, Salv, Nic, CR. Selected Specimens Examined. Ags (McVaugh & Koelz 212, MICH). BCS (Carter & Ferris 3360A, MEXU). Chih (Pringle 1442, NY). Chis (Breedlove 22395, NY). Col (Reko 4847, US). DF (Rusby 365, NY). Dgo (Pennell 18342, NY). Gro (Hinton 9650, NY). Gto (Dugas s.n., in 1880, NY). Hgo (Rose 9205, NY). Jal (Mexia 1513, NY). Me´x (Mickel 697, NY). Mich (Hinton 13421, NY). Mor (Clark 7286, NY). Nay (Norris & Taranto 14636, MICH). Oax (Mickel 3891, NY). Pue (Riba et al. 324, MEXU). Qro (Dı´az B. 4972a, IEB). SLP (Wendt & Chiang 12273d, NY). Sin (Breedlove 16748, NY). Son (Gentry et al. 19321, NY). Tlax (Vibrans 400D, MEXU). Ver (Dorantes et al. 05113, NY). Zac (McVaugh 17760, NY).

Cheilanthes pyramidalis represents one of the most difficult complexes in Mexican pteridophytes. It is one of the most widespread ferns in Mexico, occurring in 23 states and the Distrito Federal, and displaying a wide range of variation. Typically, the species has fimbriate, decurrent indusia with spicules on the indusial surfaces and along the laminar-indusial interfaces or even onto the adaxial blade surfaces (Figs. 70J, K). The segments may be narrowly oblong or, on occasion, linear and drooping (Figs. 70H, 86C–E). Some specimens lack spicules, or the indusia may not be decurrent (Figs. 86A, B). The blades are often lanceolate but may be broadly deltate. The name C. hirsuta Link has been generally applied to this species in recent years, but the isotype at PH has the longdecurrent indusia of C. marginata. As the specific epithet suggests, it is unique in having hairs on the segments, not only at the indusial margin but especially along the indusial-laminar interfaces (Fig. 86F), with some of the hairs irregularly broadened (scale-like). Such hairs are not found in specimens of C. pyramidalis, but are occasional in C. marginata. Thus, we consider C. hirsuta a synonym of C. marginata. Cheilanthes marginata is clearly allied and often confused with C. pyramidalis. Typically, it too has fimbriate, long-decurrent indusia, but it generally lacks spicules, the marginal fimbriae are longer, and the indusia are broader (0.5 mm) and longerdecurrent, not just to the subtending axis, but to the next segment. Blades of C. marginata are usually deltate (occasionally lanceolate) with less elongate segments, and the species generally occurs at higher elevations than does C. pyramidalis.

Rhizomes compact, horizontal, 2–4 mm diam.; rhizome scales linear, 4–5 mm long, bicolorous with dark reddish brown to black center and tan margins, the margins proximally with long hairs (1 mm); fronds to 50 cm long, approximate; stipes 1⁄2–2⁄3 the frond length, green to stramineous, grooved, sparsely clothed with pale linear scales and hairs; blades 2–4-pinnate-pinnatifid, broadly deltate to ovate-lanceolate, basal pair of pinnae exaggerated basiscopically, pinnae alternate to subopposite at blade base, often resulting in somewhat flexuous rachises, pinnule rachises and blade rachises with green stripes adaxially; segments deltatelanceolate; adaxial surfaces glabrous; abaxial surfaces glabrous except for sparse hairs or narrow scales (to 1 mm) on costae and major veins; veins conspicuous; sori marginal, with distinct false indusia 0.5 mm wide, broken into flaps 0.5–1 mm long; spores tan.

Distribution. Moist, exposed or shaded banks; 50–2450 m. Mexico; Guat, Salv, Nic, CR, Pan; Col. Selected Specimens Examined. BCS (Pray 1781, LAM, NY). Chih (Benı´tez 2007, MEXU). Chis (Breedlove 36706, DS). Col (McVaugh 15921, NY). DF (Rusby 370, NY). Dgo (Ibana Garcı´a 792, US). Gro (Hinton 9476, MEXU, NY). Jal (Pringle 2586, MEXU, NY). Me´x (Tejero-Dı´ez 2409, IZTA). Mich (Dı´az B. & Garcı´a 6409, IEB). Mor (Lyonnet 863, MEXU, NY). Nay (Feddema 448, NY). Oax (Mickel 6902, NY). Sin (Breedlove 18047, NY). Son (Sanders 12525, UC). Ver (Ventura A. 8841, MEXU, NY).

Cheilanthes skinneri is distinct in its deltate blades, stramineous stipes and rachises, and the indusia often interrupted into 0.5–1 mm long flaps. It is remarkable, considering the great difference in soral configuration, that C. skinneri apparently crosses with Hemionitis subcordata to form a fertile diploid hybrid, Hemionanthes gryphus (Mickel, 1987), which see for discussion.

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53. CHEILANTHES SPICULATA Cheilanthes spiculata Mickel, sp. nov. Type. Mexico. Me´xico: Temascaltepec, Cuentla, Hinton et al. 6837 (holotype NY!; isotypes F!–2 sheets, LL!, TEX!). Figs. 85D–G. A C. marginata laminis angusti-dentatis et paginis adaxialiter spiculatis abstans. (Alluding to the abundant spicules near the adaxial segment margins and the indusial-laminar interface.)

scattered linear scales and some jointed hairs; pinnae 13–25 pairs, equilateral; segments oval to oblong, rounded, pouch-like, the largest 1–2 mm diam.; adaxial surfaces tomentose with fine curved hairs; abaxial surfaces with costal scales linear, truncate at base, inconspicuous, loosely imbricate, not concealing segments, entire, not ciliate; segments abaxially densely tomentose; laminar margins recurved, slightly differentiated as false indusia, not decurrent; sporangia 32-spored; spores brown. n⫽2n⫽90, apogamous (NL, Tam, USA).

Rhizomes compact, horizontal to ascending, 1–2 mm diam.; rhizome scales linear-subulate, bicolorous, black with narrow brown to tan margins, lustrous, 3 mm long; fronds 16–32 cm long, clumped; stipes 1⁄2–3⁄4 the frond length, atropurpureous, lustrous, grooved, glabrous; blades narrowly deltate, mostly bipinnate, to tripinnate at base; segments elliptic, acute; adaxial surfaces spiculate near the margins; abaxial surfaces glabrous; laminar margins recurved, strongly modified into 0.4 mm wide false indusia with long-fimbriate margins (hairs 0.3 mm long), very long-decurrent to next segment with spicules along indusiallaminar interface; spores brown.

Distribution. Rocky slopes and ledges, on a variety of substrates including limestone and granite; 600–2400 m. USA (southern and eastern states); Mexico. Selected Specimens Examined. Chih (Knobloch 5946, NY). Coah (Palmer 1391, NY; Wynd & Mueller 337, NY). Hgo (Orcutt 6253, DS). NL (Pringle 2776, MEXU). SLP (Ortega O. 00359, NY). Son (Hartman 360, UC). Tam (Stanford et al. 2619, NY). Ver (Conant 715, MEXU; Faden et al. 76/65, UC).

Distribution. Habitat data absent. Mexico. Known only from the type. Cheilanthes spiculata resembles C. marginata in the broad, long-decurrent indusia but is distinct in the narrowly deltate blades and abundant spicules on the adaxial blade margins.

54. CHEILANTHES TOMENTOSA Cheilanthes tomentosa Link, Hort. Berol. 2: 42. 1833. Figs. 82A–E. Myriopteris tomentosa (Link) J. Sm., Hist. Fil. 280. 1875. Type. Cultivated at Berlin (B, Tryon photo NY!, frag. of B at US!; isotype PH!).

Rhizomes compact, horizontal, 2–3 mm diam.; rhizome scales linear-lanceolate, mostly bicolorous, with dark brown to black center and narrow orange-tan margins, lustrous, 3–4 mm long, entire; fronds to 45 cm long, clumped; stipes ca. 1⁄3 the frond length, castaneous, terete, with orange-tan jointed hairs and filiform scales ca. 1 mm long; blades narrowly lanceolate to narrowly oblong, 3–4-pinnate, 1.5–8 cm wide; rachises terete, with

Cheilanthes tomentosa is closely related to C. eatonii, but differs by having narrower (filiform), inconspicuous costal scales (these more evident in C. eatonii).

55. CHEILANTHES VILLOSA Cheilanthes villosa Davenp. ex Maxon, Proc. Biol. Soc. Wash. 31: 142. 1918. Type. U.S.A. Arizona: Chiricahua Mts, Sep 1881, Lemmon s.n. (GH!, frag. US!). Figs. 80B–H. Rhizomes compact, horizontal, 2–3 mm diam.; rhizome scales linear-lanceolate, mostly bicolorous, with distinct black center and narrow light brown margins, 4–5 mm long with longattenuate tips, lustrous, entire; fronds to 30 cm long, clumped; stipes dark reddish brown, terete, densely clothed with scales imbricate, ascending, scattered, tan, lanceolate, 3–4 mm long, mixed with abundant reduced linear scales; blades narrowly lanceolate to ovate, 3–4-pinnate, 1.5–5 cm wide; pinnae 8–15 pairs, equilateral; abaxial scales deltate or ovate to lanceolate, shallowly cordate at base, often with overlapping basal lobes, conspicuous, strongly imbricate and spreading laterally, far exceeding the segment margins and conspicuous from the adaxial side, usually concealing ultimate segments abaxially, whitish with orange-

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brown centers, erose-dentate, not ciliate; ultimate segments round to oval, bead-like, mostly less than 1 mm long; adaxial surfaces with scattered to dense curly white hairs; abaxial surfaces nearly glabrous except for a few curly white hairs; laminar margins recurved, but only weakly differentiated; sporangia 32-spored; spores brown. n⫽2n⫽90, apogamous (USA).

Distribution. Cliffs and rocky slopes, usually on igneous substrates; 750 m. USA (Calif); Mexico. Specimen Examined. BCN (Wiggins 13044, DS).

Distribution. Cliffs and rocky slopes, usually on limestone; 1450–2400 m. USA (Ariz, N Mex, Tex); Mexico. Selected Specimens Examined. Chih (Correll 22687, NY; Pringle 459, NY, UC). Coah (Cowan 3630, UC; Stanford et al. 202, ARIZ, NY, UC, US; Pray 2095, NY). Dgo (Chiang et al. 10004, CAS, LL, NY). Hgo (Pray 3099, LAM). NL (Chiang et al. 8017, CAS, LL, MEXU). Son (Baker 10424, ARIZ). Zac (Moreley 638, UC). Unverified, Doubtful, or Mistaken Reports. Ags (reported by de la Cerda Lemus & Siqueiros, 1985, but not verified).

Cheilanthes villosa is distinct by its large deltate scales on costae abaxially (2 mm wide and spreading, visible from adaxial side) and scattered curly hairs adaxially.

56. CHEILANTHES VISCIDA Cheilanthes viscida Davenp., Bull. Torrey Bot. Club 6: 191. 1877. Syntypes. U.S.A. [California:] Sierra Nevada near San Gorgorio Pass, Parry & Lemmon 427, April 1876 (GH [type with no. 272, with 427 crossed out]; isosyntype NY!); California/Nevada: Downieville Buttes & bluffs of White Water Canyon in the Colorado Desert, Apr–May [no year], Lemmon s.n. (isosyntype NY!). Figs. 84J–M. Rhizomes compact, ascending, 2–3 mm diam.; rhizome scales linear, concolorous reddish orange, dull or slightly viscid, flexuous, ca. 3 mm long, entire; fronds to 20 cm long, clumped; stipes 1⁄3–1⁄2 the frond length, dark reddish brown to atropurpureous, lustrous, shallowly grooved, with short, erect, glandular hairs ca. 0.1 mm long; blades narrowly oblong, bipinnatepinnatifid to tripinnate; pinnae 5–9 pairs, deltate; adaxial surfaces viscid, occasionally sessile glands visible, often with soil particles adhering; abaxial surfaces viscid with short glandular hairs 0.1 mm; sori on segment lobes, laminar margins curved, but not differentiated; sporangia 64-spored; spores brown.

Cheilanthes viscida is distinct in its glandular hairs, viscid laminar surfaces, and oblong blade shape. In its glandular hairs it most closely resembles C. kaulfussii, which is widespread in Mexico and has broadly pentagonal blades and mostly erect glandular hairs (rather than narrowly oblong blades and mostly sessile glands).

57. CHEILANTHES WOOTONII Cheilanthes wootonii Maxon, Proc. Biol. Soc. Wash. 3: 146. 1918. Type. U.S.A. Arizona: Santa Rita Mts, Madero Canyon, 21 Sep 1914, Wooton s.n. (US!). Figs. 81K–P. Rhizomes long-creeping, horizontal, 1–3 mm diam.; rhizome scales ovate-lanceolate, concolorous brown or dark brown to weakly bicolorous, ca. 2 mm long, lustrous, entire, loosely appressed and deciduous with age; fronds to 35 cm long, distant; stipes ca. 1⁄2 the frond length, dark reddish brown, terete, with abundant, loosely imbricate, linear, tan to whitish, spreading scales to 3 mm long; blades oblong-lanceolate, 3–4-pinnate, 2–5 cm wide; pinnae 10–18 pairs, equilateral; costal scales abaxially lanceolate-ovate, truncate to subcordate at base without overlapping lobes, strongly imbricate, often covering the segments, ciliate with coarse cilia confined to basal half; ultimate segments bead-like, mostly less than 1 mm; adaxial surfaces glabrous; abaxial surfaces glabrous or with a few small scales at base; laminar margins recurved, weakly differentiated, entire; sporangia 32spored; spores dark brown; n⫽2n⫽90, apogamous (USA), 2n⫽116 (Tex).

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Distribution. Oak-grassland, pine and oak, among rocks, rock crevices, granite boulders, in pine forest; 1400–2650 m. Sw USA; Mexico. Selected specimens examined. BCN (Orcutt s.n., 10 Apr 1886, UC; Wiggins 13044, DS; Moran 14204, ASC; 19171, ARIZ, 20448, SD; Moran & Thorne 14371, ENCB). Chih (Knobloch 793, US). Son (Druery s.n., “about 1895,” US; Harvey 1696, LL; Wiggins 11692, TEX, US).

Distribution. Igneous rocks, alkaline rocks, oak woodland, on and at base of rocks, oak grassland, limestone ledges, grassy rocky hillsides; 925–1800 m. USA (Ariz, N. Mex, Tex); Mexico.

Unverified, Doubtful, or Mistaken Reports. Ags (reported by de la Cerda Lemus & Siqueiros Delgado, 1985, but not verified).

Selected Specimens Examined. BCS (Carter 2016, DS, MEXU, UC; Jones 27016, UC). Chih (Knobloch 145, MEXU; Pringle 445, NY). Coah (Johnston et al. 11974, NY; Wynd & Mueller 472, NY). Dgo (Maysilles 8302, MICH). Son (Gentry et al. 19329, NY; Lloyd 498, NY, UC; White 4047, MICH).

Cheilanthes wootonii closely resembles C. yavapensis, which see for discussion.

Cheilanthes wrightii is distinct by its small size, grooved stipes, and nearly glabrous blades.

58. CHEILANTHES WRIGHTII

59. CHEILANTHES YATSKIEVYCHIANA

Cheilanthes wrightii Hook., Sp. Fil. 2: 87, t. 110A. 1858. Type. U.S.A. Western Texas to El Paso, New Mexico, 1849, Wright 823 (K; isotypes GH!, NY!, US!). Figs. 79A–D.

Cheilanthes yatskievychiana Mickel, sp. nov. Type. Mexico. Sonora: west side of Sierra del Aliso, N of Cerro El Halco´n, 28⬚ 38' 15" N, 109⬚ 43' 30" W, Burquez M. 96-302 (holotype MO!). Figs. 74F–K.

Rhizomes short- to long-creeping, horizontal, 1 mm diam.; rhizome scales lanceolate, concolorous dull orange-tan, 1–2 mm long, entire; fronds 8–22 cm long, approximate to spaced; stipes 1 ⁄2 the frond length, castaneous, lustrous, grooved, with scattered to sparse whitish hairs and linear scales to 1.5 mm long; blades narrowly oblong to lanceolate, bipinnate-pinnatifid to tripinnate; pinnae 5–8 pairs, equilateral; adaxial and abaxial surfaces glabrous; laminar margins curved, but not differentiated; spores brown; 2n⫽60 (USA).

A C. bonariense pilis abaxialiter albis rectis, pilis stipitum rachidum catenatis rectis, squamis rhizomatum concoloribus brunneis pallidis non-lustratis diversa. (Dedicated to Dr. George A. Yatskievych, who called this specimen to our attention.)

Rhizomes slender, 1 mm diam (4 mm incl. scales and stipe bases), compact, horizontal; rhizome scales linear-lanceolate, concolorous pale brown, dull, ca. 3 mm long, entire; fronds 10–15 cm long, clumped; stipes 1⁄4–1⁄3 the frond length, dark reddish brown to black with age, terete, ca. 0.6 mm diam. with abundant, lax, catenate hairs to 1 mm long, hairs deciduous with age; blades linear, 6–8 mm wide; pinnae 16–22 pairs, oblong, shallowly lobed, articulate, short-stalked ca. 1 mm; both surfaces densely clothed with straight white hairs, these obscuring the surfaces; sporangia at vein ends, laminar margins slightly curved, undifferentiated; spores tan.

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Distribution. In Platanus gallery forest of deep canyon; 800 m. Mexico. Known only from the type collection.

Distribution. Rock crevices, north-facing slopes; 1520–1650 m. USA (Ariz, N Mex, Tex); Mexico.

This species superficially resembles the widespread C. bonariensis in the linear blades with shallowly lobed pinnae and abundant laminar hairs, but C. yatskievychiana can be readily distinguished by the abaxial hairs white and straight (vs. rusty and tangled), stipe and rachis hairs catenate, straight, nonglandular (vs. curved at base and glandular), rhizome scales concolorous pale brown and dull (vs. strongly bicolorous and lustrous), and fronds only 10–15 cm long (vs. to 75 cm); it has been found only in Sonora (vs. distribution throughout Mexico except for the Yucatan Peninsula and northern Baja California).

Specimens Examined. BCN (Chambers 645, DS). BCS (Moran 18738, 23871, SD).

60. CHEILANTHES YAVAPENSIS Cheilanthes yavapensis T. Reeves ex Windham, Contr. Univ. Michigan Herb. 19: 32. 1993. Type. U.S.A. Arizona: Yavapai Co., small east-wall tributary of Black Canyon at a point 3.09 km SSW of Sheep Gulch Spring and 1.13 km NE of the confluence of Black Canyon and Sycamore Creek, Windham 202 (UT; isotypes ASC!, ASU!, US!). Figs. 80W–Bb. Rhizomes long-creeping, horizontal, 1–1.5 mm diam.; rhizome scales ovate-lanceolate, subconcolorous blackish, lustrous, 2–3 mm long, entire; fronds to 22 cm long, distant; stipes 1⁄3–1⁄2 the frond length, castaneous, terete, lustrous, with abundant linear whitish scales and hairs; blades narrowly lanceolate, 4-pinnate; pinnae 12–15 pairs, equilateral; segments bead-like, 0.5–1 mm long; adaxial surfaces glabrous; abaxial surfaces with ovatelanceolate, mostly castaneous scales, margins ciliate the entire length; laminar margins recurved, weakly differentiated; sporangia 32-spored; spores tan; n⫽2n⫽120, apogamous (USA).

Cheilanthes yavapensis is thought to be the hybrid between C. covillei and C. lindheimeri (Gastony & Windham, 1989), but has most often been treated as part of C. wootonii. It is distinct from the latter species in having: rhizome scales dark brown to bicolorous, tightly appressed, and persistent (vs. brown, loosely appressed, and somewhat deciduous); adaxial blade surfaces sparsely pubescent (vs. glabrous); abaxial blade scales ciliate most of their length (vs. ciliate only in the basal half); n⫽2n⫽120, apogamous tetraploid (vs. n⫽2n⫽90, apogamous triploid). Name of Uncertain Application Myriopteris cheiloglyphis Fe´e, Me´m. Foug. 8: 77. 1857. Type. Mexico. Veracruz: Orizaba, 2500 m, Schaffner 91 (RB). From the type description, this seems to apply to the Cheilanthes lendigera group, but whether it is C. lendigera itself or C. marsupianthes is not clear; the former is much more common.

2 6 . C H E I L O P L E C T ON Cheiloplecton Fe´e, Me´m. Foug. 7: 33. 1857. Type: Cheiloplecton rigidum (Sw.) Fe´e [⬅ Pteris rigida Sw.]. Terrestrial or epipetric; rhizomes short, compact, ascending, clothed with narrow scales; fronds small, erect, monomorphic, multiple abscission zones at stipe bases; stipes brown to black, with scales and hairs; blades 1–2-pinnate-pinnatifid; veins prominulous below, free, forking; laminae subcoriaceous to coriaceous, with short hairs adaxially, hairs and scales abaxially; sori marginal, continuous around segment margins; indusia well developed, continuous, broad, strongly recurved, hidden under recurved margin (Fig. 88D); spores tetrahedral-globose, tan, brown and black (mixed); x⫽30 Cheiloplecton is a small genus of two or three species of mesic areas (roadbanks and slopes, often among rocks) of southern Mexico and northern Central America. It is very close to Cheilanthes, but has been commonly placed in Pellaea. Gastony and

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Rollo (1998) showed Cheiloplecton to be sister to Notholaena. We are maintaining it as a marginally distinct genus because of its broad, doubly recurved margin/indusium.

1. CHEILOPLECTON RIGIDUM Cheiloplecton rigidum (Sw.) Fe´e, Me´m. Foug. 7: 34, t. 20, f. 3. 1857. Pteris rigida Sw., Syn. Fil. 104, 299. 1806. Pellaea rigida (Sw.) Hook., Sp. Fil. 2: 144. 1858. Cheilanthes rigida (Sw.) Mett., Abh. Senckenberg. Naturf. Ges. 3: 92. 1859. Type. Mexico. Me´xico: Chalma, Ne´e s.n. (MA).

Rhizomes stout, ascending; rhizome scales concolorous, light brown, becoming dark brown to black with age, linear, 6–12 ⫻ 0.8–1 mm, varying in size down to hair-like scales 3–4 mm ⫻ 0.1 mm, somewhat oily-viscid; fronds to 40 cm tall, clumped; stipes 1⁄4–1⁄2 the frond length, castaneous, with multiple abscision zones at base, with scattered light brown, linear, filiform-tipped scales 4–8 ⫻ 1 mm, each with slightly auriculate to hastate base and long, multicellular hairs often with enlarged apical cell; blades broadly deltate to lanceolate, pinnate-pinnatifid except at base where often bipinnate-pinnatifid, pinnae with sparse nonarticulate hairs 0.5–1 mm long adaxially, mostly along margins and midribs, sparse to dense hairs and scales abaxially 2–5 ⫻ 0.1– 1 mm; vein ends enlarged and conspicuous adaxially, especially visible on the recurved edges of pinnae; laminae coriaceous, margins strongly recurved; indusia strongly recurved, margins entire to papillate; spores tan, brown and black (mixed).

Key to the Mexican Varieties of Cheiloplecton rigidum 1. Blades bipinnate-pinnatifid at base, deltate, the basal pinna pair strongly exaggerated basiscopically; rhizome scales becoming dark brown with age, 6–7 mm long. .................................................................................. 1a. var. rigidum. 1. Blades pinnate-pinnatifid, lanceolate; rhizome scales becoming black with brown base with age, 10–12 mm long. ........................................................................... 1b. var. lanceolatum.

The existence of plants with intermediate blade form and well formed spores suggests that these extremes might be treated as varieties, rather than distinct species. 1a. Cheiloplecton rigidum var. rigidum.

Figs. 88B–F.

Pteris cartilaginea C. Presl, Reliq. Haenk. 1(1): 57, t. IX, f. 3. 1825. Type. Mexico. Haenke s.n. (PRC).

Similar to var. lanceolatum except: rhizome scales smaller (6– 7 mm long) and dark brown at maturity; fronds often larger (to 40 cm long); blades deltate, bipinnate-pinnatifid at base; hairs adaxially 0.5–0.8 mm long; laminae generally less hairy beneath (occasionally subglabrous); scales abaxially 2 ⫻ 0.1–0.3 mm.

Distribution. Rocky slopes, woods by river, mossy ledges, deciduous woodlands, some calcareous, gypsum and slate, igneous rocks; 100–1900 m. Mexico; Guat, Salv. Selected Specimens Examined. Chih (Palmer s.n., US). Chis (Breedlove 36530, MEXU). Col (McVaugh 15523, MICH). Gro (Mexia 8826, NY). Gto (Ventura y Lo´pez 7331, MEXU). Hgo (Fisher s.n., 11 Aug 1937, NY). Jal (Pringle 2602, MICH). Me´x (Mickel 686, NY). Mich (Hinton 12229, NY). Mor (Storer 94, UC). Nay (Norris & Taranto 14042A, MICH). NL (Aguirre C. 510, ENCB). Oax (Mickel 6264, NY). Pue (Purpus 4177, NY). Qro (Rzedowski 51531, IEB). SLP (Pringle 3298, NY). Son (Martin et al. s.n., 13 Oct 1988, ARIZ). Tam (Yatskievych 83-331, ARIZ). Ver (Seaton 343, NY). Zac (Johnston et al. 12233, NY). Unverified, Doubtful, or Mistaken Reports. Ags (Garcı´a 4802, HUAA, and one other collections, cited by Siqueiros-Delgado & Gonza´lez-Adame, 2004, but not verified).

Some broad-bladed specimens have longer black rhizome scales and more heavily clothed abaxial blade surfaces. Whether these characters are variable or whether they represent intergradations with var. lanceolatum is not known. 1b. Cheiloplecton rigidum var. lanceolatum C. C. Hall, Mickel & Beitel, Mem. New York Bot. Gard. 46: 124. 1988. Type. Mexico. Oaxaca: Distrito Huajuapan, 9 km NW of Huajuapan de Leo´n, Mickel 4093 (NY!). Fig. 88A. Rhizome scales concolorous, light brown, becoming black with brown base with age, sclerotic, linear, 10–12 ⫻ 0.8 mm, varying in size down to hair-like scales 4 ⫻ 0.1 mm, somewhat oily-viscid; fronds to 28 cm tall, clumped; stipes 1⁄4–1⁄2 the frond length, with multiple abscision zones at base, black, stout, clothed with light brown, linear, filiform-tipped scales 8 ⫻ 1 mm with slightly hastate base and long multicellular hairs, often with enlarged apical cell; blades pinnate-pinnatifid, narrowly oblong to lanceolate, broadest at the middle; pinnae with hairs and narrow scales abaxially 4–5 ⫻ 1 mm, sparse hairs 0.8–1 mm long adaxially, non-articulate; veins barely visible, vein endings enlarged and conspicuous adaxially, especially visible on the recurved edges of pinnules; laminae coriaceous, margins strongly recurved; indusia strongly recurved, top and bottom portions each 0.8 mm wide, margins fimbriate (to papillose), cells along recurved margins whitened; spores tan, brown and black (mixed); n⫽2n⫽90 (Pue), apogamous.

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215

1. Blades chartaceous; sori (2–)4–5 pairs per segment, often directed laterally, usually slightly spaced; stems rhizomatous or with a trunk to 1 m; Ver. ...................... 2. C. schiedei.

1. CIBOTIUM REGALE Cibotium regale Verschaff. & Lem., Ill. Hort. 15: under pl. 548, f. 5. 1868. Type. Cultivated material, collected in Mexico. Chiapas: Ghiesbreght 351 (GH; isotypes K!, PH!, YU!). Fig. 89D.

Distribution. Rocky slopes, in open to lightly wooded, dry to moist areas; 1050–1850 m. Mexico. Selected Specimens Examined. Gro (Orcutt 4229, US; Rowell 4002, US). Jal (Puga 58, ENCB, 3313, IBUG). Oax (Breedlove 59778, CAS; Mickel 6289, NY). Pue (Purpus 1153, NY; Rzedowski 27891, ENCB, NY). SLP (Takaki 1751, UAMIZ). Sin (Breedlove 43884, MEXU).

Cibotium guatemalense Rchb. f. ex Kuhn, Linnaea 16: 152. 1869. Type. Guatemala. Wendland s.n. (B!). Cibotium wendlandii Mett. ex Kuhn, Linnaea 36: 151. 1869. Type. Guatemala. Wendland s.n. (B).

Similar to Cibotium schiedei except: trunks to 1.5 m (vs. rhizomatous to trunk-forming), blades subcoriaceous to coriaceous (vs. chartaceous), and sori (4–)6–8 pairs per segment, erect, and approximate (vs. (2–)4–5 pairs per segment, often directed laterally, and usually slightly distant).

The lanceolate, pinnate-pinnatifid blades distinguish this variety from var. rigidum. Black rhizome scales over 10 mm long are characteristic of var. lanceolatum, but are also found in some specimens of var. rigidum, which see for further discussion. Intermediates with somewhat reduced blade bases or plants with mixed reduced and non-reduced blades are known from Hidalgo.

27. C I B O T I UM Cibotium Kaulf., Berlin. Jahrb. Pharm. Verbundenen Wiss. 21: 53. 1820. Type: Cibotium chamissoi Kaulf. Terrestrial; stems stout, horizontal to erect, forming stout trunks (ca. 1.5 m tall in ours, erect and taller in Asian and Pacific species, to 15 m), densely clothed with long, soft, golden hairs; fronds large, with dense masses of golden hairs at stipe bases, bipinnate-pinnatifid to tripinnate, coriaceous to chartaceous, glabrous, glaucous or green beneath; veins free; sori marginal, indusia bivalvate, the lower valve narrower, fitting inside the upper, both valves different color from laminar tissue; annuli oblique; x⫽68. Cibotium comprises about 10 species of eastern Asia, Hawaii, Mexico, and Central America. It occurs in middle elevation rain forests in the New World.

Distribution. Montane rain forests; 1140–2200 m. Mexico; Guat, Bel (Munro et al. 3277, MO), Hond, Salv. Specimens Examined. Chis (Bachem C. et al. 542, UAMIZ; Breedlove 21761, MEXU, 31656, CAS, MEXU, 32516, MEXU; MacDougall s.n., 22 Oct s. ann., MEXU; Pe´rez Farrera 150a, UAMIZ; Purpus 9195, UC, US; Riba 1410, ENCB, MEXU).

Stolze (1976) discussed the variation of characters in Cibotium and reasons for combining C. guatemalense and C. wendlandii under C. regale.

2. CIBOTIUM SCHIEDEI

Reference Maxon, W. R. 1912. Studies of tropical American ferns. No. 3. Contr. U.S. Natl. Herb. 16: 54–58 [Cibotium].

Key to the Mexican Species of Cibotium 1. Blades subcoriaceous to coriaceous; sori (4–)6–8 pairs per segment, erect, approximate; trunks to 1.5 m; Chis. ....................................................................................... 1. C. regale.

Cibotium schiedei Schltdl. & Cham., Linnaea 5: 616. 1830. Figs. 89A–C. Dicksonia schiedei Baker in Hooker & Baker, Syn. Fil. 50. 1866. Type. Mexico. Veracruz: Hacienda de la Laguna, Schiede 801 (B!; isotype US!).

Trunks rhizomatous to erect, to 1 m tall; fronds 2–4 m, chartaceous; blades broadly ovate, nearly tripinnate to tripinnatepinnatifid; stipes and rachises smooth, pinnae lanceolate; costae

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and costules sparsely to densely pubescent; segments dark green adaxially, paler to glaucous abaxially, the margins revolute; sori (2–)4–5 pairs per segment, more or less erect to directed laterally, slightly spaced along segment margins.

stalked; rachises non-alate; sori 0.5–0.8 mm diam.; indusia brown. ................................................................. 1. C. apiculata. 1. Basal veins regularly united by a transverse veinlet, forming a single series of costal areoles; margins lightly and remotely crenate-serrate; pinnae decurrent; rachises alate; sori 1 mm diam.; indusia tan. .............................. 2. C. decurrens.

1. CNEMIDARIA APICULATA Cnemidaria apiculata (Hook.) Stolze, Fieldiana, Bot. 37: 40. 1974. Figs. 90A–C. Hemitelia apiculata Hook. in Hooker & Baker, Syn. Fil. 29. 1868. Cyathea aristata Domin, Acta Bot. Bohem. 9: 93. 1930, nom. nov., non Cyathea apiculata Domin, 1929. Type. Mexico. Oaxaca: Sierra San Pedro Nolasco, Talea, etc., Ju¨rgensen 873 (K!, frags. NY!, US!; isotype P!, photo F).

Distribution. Deciduous forests; montane cloud forests, oak forests; 600–1400 m. Mexico. Selected Specimens Examined. Ver (Bellerman s.n., May 1886, US; Bourgeau 2378, B; Copeland herb. 73, B, MEXU, MICH, UC, US; Galeotti 6458, BR; Hutchison s.n., UC; Purpus 1976, MICH, UC, US; Schaffner s.n., B; Serres s.n., 31 Oct 1912, US; Ventura A. 235, MICH, 4841, ENCB, MEXU, 13380, 13394, IEB).

Maxon (1912) reported C. schiedei from Veracruz and Oaxaca, but cited specimens only from the former.

2 8 . C N E M I DA R I A

Stems short, erect, hardly emerging from soil, clothed with bicolorous scales; fronds ca. 2 m long; stipes 1⁄3–1⁄2 the frond length, stramineous, brown towards base, spines lacking or present at base, 1 mm long, scales dark brown with margins pale, erose; blades pinnate-pinnatifid, rachises non-alate, terminal portion gradually pinnatifid; pinnae sessile to slightly stalked, to 32 cm long, lobes slightly falcate, serrulate toward apices, sharply cuspidate, apices acuminate, dark spot at base of each pinna abaxially; costae with a few bicolorous scales and minute brown, amorphous scales; veins mostly 1–2-forked, mostly free, basal veins connivent at sinuses; sori supramedial (2⁄3–3⁄4 the way from costae to margins), almost submarginal on veins, not necessarily at the vein fork, 0.5–0.8 mm in diam.; indusia brown, subentire, 0.5 ⫻ 0.3 mm, hood-like, on proximal side of sori.

Cnemidaria C. Presl, Suppl. Tent. Pterid. 56. 1836. Type: Cnemidaria speciosa C. Presl. For synonymy, see Stolze (1974). Terrestrial; stems erect, to 0.5 m long, apex scaly; fronds to 3.5 m long; blades pinnate to pinnate-pinnatifid; stipes smooth or spiny, spines to 7 mm long, stipe scales marginate, commonly bicolorous; axes and midribs glabrous adaxially, laminae glabrous to hairy with multicellular trichomes; basal adjacent veins forming costal areoles, or connivent at sinuses; sori subglobose, receptacles raised; indusia present, hemitelioid (saucer-shaped); spores 64 per sporangium, trilete with one large pore near center of each face on or near equator; x⫽69. Cnemidaria is a neotropical genus of about 25 species with two in Mexico. The genus is distinct among cyatheoid tree ferns by the pinnate-pinnatifid fronds, marginate stipe scales, glabrous adaxial costae, hemitelioid indusia, and triporate spores. Reference Stolze, R. G. 1974. A taxonomic revision of the genus Cnemidaria (Cyatheaceae). Fieldiana, Bot., n.s. 37: 1–98.

Key to the Mexican Species of Cnemidaria 1. Veins mostly free, rarely forming basal arches along the costae; margins serrulate; pinnae sessile to slightly

Distribution. Terrestrial in wet montane forests; 1100–1700 m. Mexico. Specimens Examined. Oax (Hallberg 1557, NY; Mickel 994, 5676, 5717, 5935, NY). Ver (Lira 30, MEXU, XAL; Pe´rez Garcı´a et al. 165, MEXU).

Cnemidaria apiculata is distinct by its serrate pinna margins and mostly free veins. The dark spot at the base of each pinna, possibly a nectary or aerophore, deserves study regarding its function and origin.

COCHLIDIUM

2. CNEMIDARIA DECURRENS Cnemidaria decurrens (Liebm.) R. M. Tryon, Contr. Gray Herb. 200: 52. 1970. Figs. 90D, E. Hemitelia decurrens Liebm., Mexic. Bregn. 286 (reprint 134). 1849. Hemistegia decurrens (Liebm.) E. Fourn., Mexic. Pl. 1: 135. 1872. Cyathea decurrentiloba Domin, Acta Bot. Bohem. 9: 110. 1930, nom. nov., non Cyathea decurrens (Hook.) Copel., 1929. Type. Mexico. Oaxaca: Distr. Chinantla, pr. Lobani, Liebmann s.n. [Pl. Mex. 2089, Fl. Mex. 912] (C!, frag. US!). Hemitelia mexicana Liebm., Mexic. Bregn. 287 (reprint 135). 1849. Hemistegia mexicana (Liebm.) E. Fourn., Mexic. Pl. 1: 135. 1872. Cyathea liebmanii Domin, Pterid. Dominica 264. 1929, nom. nov., non Cyathea mexicana Schltdl. & Cham., 1830. Type. Mexico. Oaxaca: Distr. Chinantla, pr. Lacoba, Liebmann s.n. [Pl. Mex. 2105, Fl. Mex. 909, 910, 911]; three parts of one frond (Stolze, 1974) (C!, frag. US!; isotype BM!). Hemistegia lucida Fe´e, Me´m. Foug. 5: 351. 1852. Hemitelia lucida (Fe´e) Maxon, Contr. U.S. Natl. Herb. 16: 39. 1912. Cyathea lucida (Fe´e) Domin, Pterid. Dominica 264. 1929. Type. Mexico. Oaxaca: Galeotti 6537 (BR!; isotypes BM!, BR!–13 sheets, P!–5 sheets, US!). Hemistegia elegantissima Fe´e, Me´m. Foug. 8: 110. 1857. Cyathea elegantissima (Fe´e) Domin, Acta Bot. Bohem. 9: 113. 1930. Type. Mexico. Linden s.n. Specimen unknown, according to Stolze (1974). Hemitelia guatemalensis Maxon, Contr. U.S. Natl. Herb. 16: 40. 1912. Cyathea guatemalensis (Maxon) Domin, Pterid. Dominica 264. 1929. Type. Guatemala. Alta Verapaz: Salvin s.n. (US!; isotypes GH!, K!, US!).

Stems stout, erect, to 0.5 m tall; fronds 2–3 m long, from a compact crown; stipes green to stramineous or light brown, dark brown at base, armed with short spines, 3–4 mm long, scurfy with matted hairs and small scales, larger scales dark brown with margin narrow, pale, erose; blades 1-pinnate, rachises alate; pinnae entire to crenate, narrowly lanceolate, apices gradually pinnatifid, sessile or subsessile, bases broadly cuneate, the distal pinnae broadly adnate and decurrent on the rachises, lacking dark dots at base; laminae glabrous on both surfaces, but with twisted, matted hairs and scattered scales abaxially on costae and sometimes on veins; veins forming costal areoles, others free, connivent to sinuses; sori medial on veins, 1 mm diam.; indusia thin, tan, small, 0.3 mm long, to one side at base of sori.

Distribution. Terrestrial in wet montane forests at lower middle elevations of the Atlantic slope; 100–1350 m. Mexico; Guat, Hond. Selected Specimens Examined. Chis (Mu¨nch 4, DS, US). Oax (Martı´nez C. 515, MEXU, XAL; Mickel 5889, NY, 5890, MEXU, NY;

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Wendt et al. 4747, MEXU). Ver (Beaman 06110, MEXU; Calzada 8369, MEXU, XAL; Lira 104, MEXU, XAL; Nee et al. 24995, MEXU; Pe´rez Garcı´a 163, MEXU).

Cnemidaria decurrens is distinguished from C. apiculata by having decurrent pinnae, less incised pinna margins, and regular costal areoles.

29. COCHLIDIUM Cochlidium Kaulf., Berlin. Jahrb. Pharm. Verbundenen Wiss. 21: 36. 1820. Type: Cochlidium graminoides (Sw.) Kaulf. [⬅ Grammitis graminoides Sw.]; also see Bishop, 1978. Xiphopteris Kaulf., Berlin. Jahrb. Pharm. Verbundenen Wiss. 21: 35. 1820. Lectotype (chosen by J. Sm., Hist. Fil. 179. 1875): Xiphopteris serrulata (Sw.) Kaulf. [⬅ Acrostichum serrulatum Sw.]. For additional synonymy, see L. E. Bishop (1978).

Small epiphytes; rhizomes ascending to short-creeping; rhizome scales orangish to tan, concolorous, linear-lanceolate, entire or nearly so; fronds clumped; stipes not articulate; blades undivided or occasionally furcate, firm, chartaceous to subcoriaceous, glabrous or with scattered minute catenate hairs, lacking reddish brown setae, margins entire or sinuate; veins obscure or occasionally visible, simple or forked, sometimes united to form costal areoles without free veinlets; sori either linear, consisting of a line of sporangia borne in a medial groove and immersed or superficial, or sori intermittent or round and borne at the base of lateral veins, often confluent at maturity; indusia and paraphyses absent; sporangia glabrous with vertical annulus; spores green, tetrahedral-globose; x⫽33, 34?, 35 (counts of 2n⫽ca. 148, based on 37, are now thought to be erroneous). According to Bishop (1978), Cochlidium comprises 16 neotropical species, one of which extends into Africa and adjacent islands; these are included in Grammitis by some workers. Three species of Cochlidium are found in Mexico, where they grow in montane rain forests. The type of Xiphopteris, X. serrulata, has been placed in Grammitis s.l. by numerous authors, but Bishop (1978) included X. serrulata in Cochlidium on the basis of the pale concolorous scales, lack of branched trichomes, a stellar type similar to that of Cochlidium, and the conduplicate nature of the fertile region. He believed that the almost-fused nature of the receptacles suggested a derivation from a coenosoral ancestor, although the series suggests to us a trend in the opposite direction (separate sori to coenosori) with equal or greater justification. Preliminary molecular data suggest that Cochlidium is monophyletic (Ranker et al., 2004) and the sister group to the blackmargined, simple bladed species of Grammitis (Grammitis s.str.) allied to the group of G. bryophila (Maxon) F. Seym. in the Neotropics; this group also includes several closely related species in East Africa and Madagascar to Polynesia. The clade Grammitis s.str. plus Cochlidium, in turn, appears to be the sister group of the peculiar endemic Hawaiian genus Adenophorus (Ranker et al., 2003).

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COCHLIDIUM References

Bishop, L. E. 1978. Revision of the genus Cochlidium (Grammitidaceae). Amer. Fern J. 68: 76–94; Copeland, E. B. 1952. The American species of Xiphopteris. Amer. Fern J. 42: 41–52, 92–110; Maxon, W. R. 1914. Notes upon Polypodium duale and its allies. Contr. U.S. Natl. Herb. 17: 398–406; Ranker, T. A., J. M. O. Geiger, S. C. Kennedy, A. R. Smith, C. H. Haufler & B. A. Parris. 2003. Molecular phylogenetics and evolution of the endemic Hawaiian genus Adenophorus (Grammitidaceae). Molec. Phylogen. Evol. 26: 337–347; Ranker, T. A., A. R. Smith, B. S. Parris, J. M. O. Geiger, C. H. Haufler, S. C. K. Straub, & H. Schneider. 2004. Phylogeny and evolution of grammitid ferns (Grammitidaceae): a case of rampant morphological homoplasy. Taxon (53(2): 415–428).

Key to the Mexican Species of Cochlidium 1. Blades pinnatifid; sori on lateral veins, coalescing at maturity into a coenosorus on each side of the midrib. .................................................................................... 3. C. serrulatum. 1. Blades simple, entire; sorus in a single groove along midrib. 2. Hydathodes conspicuous adaxially; sporangial capsules less than 250 ␮m long; spores less than 43 ␮m diam. ............................................................... 1. C. linearifolium. 2. Hydathodes absent adaxially; sporangial capsules more than 250 ␮m long; spores more than 43 ␮m diam. ........................................................................... 2. C. rostratum.

1. COCHLIDIUM LINEARIFOLIUM Cochlidium linearifolium (Desv.) Maxon ex C. Chr., Dansk Bot. Ark. 6(3): 23. 1929. Figs. 91A–C. Monogramma linearifolia Desv., Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck. Gesammten Naturk. 5: 302. 1811. Grammitis linearifolia (Desv.) Steud., Nomencl. Bot. 2: 187. 1824. Type. French Guiana. s. coll. s.n. (P).

Selected Specimens Examined. Chis (Breedlove 30161, DS, MEXU, NY, 36965, DS; Heath & Long 300, CAS, MEXU; Martı´nez S. et al. 25269, MEXU). Gro (Lorea 4580, FCME). Oax (Breedlove 64731, CAS; Mickel 3800, 5953, NY, UC). Ver (Lira 27, MEXU, XAL; Vasquez 557, XAL).

This species is closely related to C. rostratum, and differs primarily by the presence of relatively conspicuous hydathodes. Throughout most of its range, C. linearifolium occurs at low elevations, usually below 1000 m, and is separated elevationally from C. rostratum, which generally occurs at elevations above 1500 m. But in Mexico, this distinction apparently does not hold, as specimens with visible hydathodes occur from 1000– 3100 m. Specimens determined by Mickel and Beitel (1988) and Palacios-Rios (1992) as C. rostratum have all been re-examined and are all C. linearifolium; both species are now known from Chiapas. Mickel collections cited were all determined by Bishop, in 1989.

2. COCHLIDIUM ROSTRATUM Cochlidium rostratum (Hook.) Maxon ex C. Chr., Dansk Bot. Ark. 6(3): 23. 1929. Figs. 91D, E. Monogramma rostrata Hook., Sp. Fil. 5: 122. 1864 as “Monogramme.” Grammitis rostrata (Hook.) R. M. Tryon & A. F. Tryon, Rhodora 84: 129. 1982. Type. Nicaragua. Island on the Lake Omotepec [Isla de Ometepe], Wright s.n. (K!; isotype US!).

Differs from C. linearifolium in the absence of hydathodes (as viewed at 30 times magnification) and in the larger sporangia (according to Bishop, nearly twice the size of those of C. linearifolium); spores more than 43 ␮m diam.; 2n⫽66 (CR).

Rhizomes ascending, or very short-creeping; rhizomes scales tan to orange-brown, ca. 2 mm long; fronds clumped; stipes absent; blades 45–50(–90) ⫻ 1.5–2 mm, entire, decurrent to base of frond, thick, glabrous; veins obscure; hydathodes visible as rounded dots adaxially at ends of veins (only slightly visible in some specimens); sori elongate along the distal 1⁄4–1⁄2 of midvein, in a deep groove; spores less than 43 ␮m diam.

Distribution. Uncommon epiphyte in cloud forests or montane forests; 1300 m. Mexico; Guat, Hond, Salv, Nic, CR, Pan; Gr & L Ant; Col, Ven. The specimen cited as this species by Bishop (in Davidse et al., 1995) from Belize (Allen 15304, MO!) is C. linearifolium. Specimens Examined. Chis (Breedlove 32226, DS, LL, MEXU, 35108, DS, 38888, DS, MEXU, 53296, CAS).

Distribution. Epiphytic in lowland and montane rain forests; 150–3100 m. Mexico; Guat, Bel, Hond, Nic, CR, Pan; Col, Ven, Guy, Sur, Fr Gui, Ec, Braz.

Cochlidium rostratum is generally distinguished from C. linearifolium by the larger spores and absence of hydathodes (see Bishop, 1978). All specimens identified as this species from Oaxaca (Mickel & Beitel, 1988) and Veracruz (Palacios Rios, 1992)

CTENITIS

have hydathodes, and so are referrable to C. linearifolium. Specimens from Chiapas cited by Smith (1981) are mostly C. linearifolium, except those from the vicinity of Lagunas de Monte Bello and Laguna Tsiskaw, near the Guatemala border, where C. rostratum appears to be more common (all four collections cited above); however, one collection of C. linearifolium (Breedlove 36965, DS) also occurs here. Further work, especially including cytological studies, is needed to clarify the reputed differences between these two species. If they prove to be conspecific, C. linearifolium is the correct name.

3. COCHLIDIUM SERRULATUM Cochlidium serrulatum (Sw.) L. E. Bishop, Amer. Fern J. 68: 80. 1978. Figs. 91F, G. Acrostichum serrulatum Sw., Prodr. 128. 1788. Grammitis serrulata (Sw.) Sw., J. Bot. (Schrader) 1800(2): 18. 1801. Xiphopteris serrulata (Sw.) Kaulf., Wesen Farrenkr. 87. 1827. Polypodium serrulatum (Sw.) Mett., Fil. Hort. Bot. Lips. 30. 1856, hom. illeg., non Sw., 1802. Polypodium duale Maxon, Contr. U.S. Natl. Herb. 16: 61. 1912. Type. Jamaica. Swartz s.n. (S, isotype US!). For additional synonymy, see L. E. Bishop (1978).

Rhizomes ascending; rhizome scales orange-brown, 1–1.3 mm long; stipes to ca. 1.5 cm long, dark brown at very base, green distally, glabrous; blades dimorphic, sterile blades shorter, deeply pinnatifid, 10–20 ⫻ 2–3 mm, segments 1–1.5 mm long; fertile blades longer, 35–60 mm long, fertile in distal half, basal sterile part pinnatifid, 25–35 ⫻ 1–1.5 mm, the segments ascending, deltate, 0.8–1 mm long, chartaceous, glabrous; veins evident on both sides of blades; hydathodes none or not readily visible; sori basal on unbranched veins, appearing nearly costal but not in deep grooves, the blades somewhat folded (conduplicate); 2n⫽ca. 105 (Jam, Trin), 140 (Jam).

Distribution. Epiphytic or terrestrial on mossy banks of the Atlantic slope; 150–1850 m. Mexico; Guat, Bel, Hond, Salv (Seiler 1004, NY), Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Guy (Gillespie 2764, TEX), Ec, Galapagos, Peru, s Braz, Bol; tropical Africa, Madagascar, Mauritius. Selected Specimens Examined. Chis (Breedlove 53298, 56522, CAS; Dressler 1641, MEXU, NY, UC, US). Oax (Mexia 9124, ARIZ, F, GH, LL, MO, NY, UC; Mickel 898, 4731, UC). Pue (Sharp 441735, MEXU, US). Ver (Nee et al. 24973, F, NY; Purpus 3021, MO, UC, US, 8689, UC).

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This species is readily distinguishable from other species of Cochlidium by the pinnatifid sterile blades and the toothed or pinnatifid sterile portions of fertile blades. In Mexico, it is most likely to be confused with certain species of Lellingeria, particularly L. hellwigii and L. prionodes, which differ in the broader blades (2.5–4.5 mm wide in fertile portion), darker, clathrate rhizome scales, bifurcate hairs along the abaxial costae, and the flattish (rather than more-or-less conduplicate) fertile part of blade.

30 . C T E N I T I S Ctenitis (C. Chr.) C. Chr. in Verdoorn, Man. Pterid. 544. 1938. Dryopteris Adans. subg. Ctenitis C. Chr. in Rosenvinge, Biologiske Arbejder Tilegnede Eug. Warming, 77. 1911. Type: Ctenitis distans (Brack.) Ching [⫽ Aspidium ctenitis Link] Terrestrial; rhizomes mostly suberect to erect, sometimes short-creeping, scaly, with old stipe bases and roots frequently forming stout caudices to 4 cm diam.; fronds small to mediumsized to large, usually clumped; blades herbaceous to chartaceous, pinnate-pinnatifid to quadripinnate or more divided; midribs, especially the adaxial rachises, costae, and costules, with short, reddish to red-brown, jointed hairs (ctenitoid hairs) 0.2–0.8 mm long; rachises and costae abaxially with few to many scales, often these often inrolled, sometimes clathrate, flat or plane, also often with glands and jointed hairs; veins free, simple or forking in the ultimate segments; sori abaxial, round, indusia reniform, attached at sinuses, sometimes greatly reduced or even absent altogether; spores bilateral, spinulose; x⫽41. Ctenitis is a genus of about 150 species, about evenly distributed between the Old and New World wet tropics, mostly of low and middle elevations. It has usually been thought to be related to the tectarioid ferns and perhaps allied to Tectaria and Lastreopsis. However, unpublished molecular data indicate that this relationship may not be as close as once thought (Hasebe et al., 1995; Cranfill, unpubl. data); rather, Ctenitis may be more intimately related to the dryopteroid clade (rather than the tectarioid clade). Twenty-one species of Ctenitis are known from Mexico, most of them restricted to only the southernmost part of the country. Apparently, southern Mexico and Guatemala, especially in calcareous habitats, are a major center of diversity for the genus in the Neotropics. Almost all species of Ctenitis have distinctive jointed hairs on the blades, especially on the adaxial rachises, costae, and minor axes. These hairs are usually short, less than 0.5 mm long and often much shorter than that, with very short cells and reddish joints. This kind of hair, often called a ctenitoid hair, is named for the genus, and is an easy character to use in distinguishing Ctenitis from most other genera of ferns, particularly Dryopteris and Thelypteris, with which Ctenitis has historically been combined and confused. Similar hairs are found in Tectaria. Reference Rojas, A. F. 2001. Ocho nuevas especies y nuevos a´mbitos geogra´ficos de helechos de la familia Tectariaceae (Filicales) en el Neotro´pico. Revista Biol. Trop. 49: 467–488.

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CTENITIS

Key to the Mexican Species of Ctenitis 1. Blades pinnate-pinnatifid to bipinnate, if bipinnate then pinnules entire, serrate, or crenate at margins, never cut more than half their width. 2. Indusia present. 3. Costae and laminae glandular-pubescent between veins abaxially, as well as on indusia. 4. Costal scales strongly inrolled at their bases; blades bipinnate proximally, the pinnules entire to crenate or serrate. 5. Pinnules crenate to serrate. ...................................................................................................................................................... 2. C. bullata. 5. Pinnules entire. ....................................................................................................................................................................... 18. C. strigilosa. 4. Costal scales flat; blades deeply pinnate-pinnatifid at bases, the segments entire. 6. Indusia large, persistent; segments rounded at the tips. ............................................................................................ 13. C. microchlaena. 6. Indusia small, ephemeral and not easily seen in mature fronds; segments apiculate at tips. .............................. 19. C. submarginalis. 3. Costae and laminae without glandular hairs between veins abaxially, indusia glabrous 7. Veins reaching segment margins (as viewed adaxially); segments entire or minutely toothed only at tips; costal scales linear-lanceolate, not inrolled at bases, flattish. ............................................................................................................... 20. C. thelypteroides. 7. Veins stopping short of segment margins (as viewed adaxially); segments serrate or obtusely lobed; costal scales strongly inrolled at bases, saccate. 8. Blades bipinnate proximally, pinnules obtusely lobed; costal scales blackish, strongly clathrate. ........................... 10. C. lanceolata. 8. Blades deeply pinnate-pinnatifid with crenately to serrately incised pinnules; costal scales tan, not clathrate. ........ 16. C. salvinii. 2. Indusia absent. 9. Costal scales reddish, flat, lanceolate; segments Ⳳ entire, rounded to truncate at apices. 10. Pinnae incised 1⁄3–2⁄3 their width; rachises and costae abaxially with dense, spreading, stiff scales. ................................ 15. C. refulgens. 10. Pinnae incised 3⁄4 their width, or more; rachises and costae abaxially with scattered, Ⳳ appressed, lax scales. .... 19. C. submarginalis. 9. Costal scales somewhat to decidedly inrolled, or with expanded, subclathrate bases and darkened, subulate tips; segments often toothed toward tips, acutish to mucronate at apices. 11. Sori inframedial; rhizomes suberect, fronds fasciculate; proximal pinnae deflexed; tubular, yellowish glands absent or sparse on laminae abaxially; pinna segments approximate or imbricate. .............................................................................. 1. C. baulensis. 11. Sori medial; rhizomes short-creeping, fronds not fasciculate; proximal pinnae not deflexed; tubular, yellowish, appressed glands often on laminae abaxially; pinna segments separated by wide sinuses. ............................................. 14. C. nigrovenia. 1. Blades bipinnate-pinnatifid (at least at bases) to tripinnate-pinnatifid. 12. Costal scales decidedly inrolled (saccate) basally. 13. Blades tripartite, with proximal pinna pair much larger than more distal pinna pairs; pinnae above the proximal pair deeply pinnatifid or barely pinnate basally; indusia present or absent. 14. Indusia present; proximal pinna pair ca. 25 ⫻ 15 cm, basal basiscopic pinnule, and sometimes also the next pair, only 1 ⁄4 the length of longest pinnule on the same pinna. ................................................................................................................ 5. C. erinacea. 14. Indusia absent; proximal pinna pair (5–)10–16 ⫻ (3–)8–12 cm, basal basiscopic pinnule usually 3⁄4 or more the length of next pair. ................................................................................................................................................................................ 8. C. hemsleyana. 13. Blades not tripartite, gradually reduced and less dissected from bases to apices; pinnae above the proximal pair pinnate or more divided; indusia present. 15. Stipes and rachises moderately scaly; indusia with a much darkened, almost blackish, center. .................................. 11. C. melanosticta. 15. Stipes and rachises very densely scaly; indusia uniformly tan or slightly darkened toward center. .................................... 21. C. ursina. 12. Costal scales flat or merely vaulted (with slightly incurved margins) basally. 16. Blades shorter than 25 cm; indusia absent; blades densely glandular on both sides. .............................................................. 3. C. chiapasensis. 16. Blades usually longer than 50 cm; indusia present or absent; blades glandular or eglandular. 17. Stipes and rachises with dark brownish, hair-like, patent or retrorse scales; costal scales uniformly castaneous; indusia large, persistent, ciliate-glandular. ............................................................................................................................................ 7. C. grisebachii. 17. Stipes and rachises with reddish to somewhat darkened, thin, often lax scales, those of stipe bases forming a large woolly tuft; indusia usually small and caducous, or absent. 18. Proximal pinnae not or only slightly basiscopically enlarged; costal scales not aggregated in pinna axils, tan, not clathrate, not vaulted. ............................................................................................................................................................ 9. C. interjecta. 18. Proximal pinnae strongly basiscopically enlarged; costal scales most numerous in pinna axils, tan to blackened, often clathrate and vaulted. 19. Segment margins eciliate; indusia absent; 50–800(–1400) m. ...................................................................................... 6. C. excelsa. 19. Segment margins ciliate; indusia present (sometimes very small and obscure in mature sori) or absent. 20. Stipe base scales filiform, orangish or reddish tan; blades abaxially with yellowish, cylindrical glands; 350 m. ................................................................................................................................................................................ 17. C. sloanei. 20. Stipe base scales brownish; blades eglandular or sparsely glandular abaxially; 600–2700 m. 21. Scales in pinna axils brownish, strongly clathrate; veins ending at margins (often in marginal teeth) or nearly so; indusia usually present, often small and hidden in mature sori; (700–)1300–2700 m. ...... 4. C. equestris. 21. Scales in pinna axils stramineous, not or weakly clathrate; veins ending 0.1–0.2 mm before margins; sori exindusiate; 600–1800 m. .................................................................................................................... 12. C. mexicana.

CTENITIS BULLATA

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1. CTENITIS BAULENSIS

2. CTENITIS BULLATA

Ctenitis baulensis A. R. Sm., Proc. Calif. Acad. Sci., ser. 4, 40: 213, f. 2I–K. 1975. Type. Mexico. Chiapas: Mpio. Cintalapa, 16 km NW of Rizo de Oro, SE of Cerro Baul, Breedlove 21812 (DS! isotype NY!) Figs. 93L, M.

Ctenitis bullata A. R. Sm., Proc. Calif. Acad. Sci., ser. 4, 40: 213, f. 2E–H. 1975. Type. Mexico. Chiapas: Mpio. La Trinitaria, Lagos de Monte Bello, Breedlove 25339 (DS!; isotypes MEXU!, NY!). Figs. 94C, D.

Rhizomes suberect, the caudices 2–3 cm diam.; fronds ca. 40 cm long; stipes darkened at bases, stramineous or faintly greenish distally, ca. 1⁄2 the frond length, to 20 cm ⫻ 2 mm, bases densely scaly, scales to 3 ⫻ 1.5 mm, flattened to somewhat inrolled, tan, but darkened toward the attenuate tips; blades green, ovatelanceolate, deeply pinnate-pinnatifid, to 20 ⫻ 13 cm; rachises abaxially with scales similar to those of stipes, but shorter; pinnae to ca. 10 pairs, sessile, largest 7 ⫻ 2.5 cm, deeply incised nearly to costae, proximal pinnae narrowed at their bases; segments approximate or even imbricate, entire to denticulate, tips rounded or subacute, margins ciliate with jointed hairs; veins simple, to 9 pairs per segment, reaching margins above sinus; indument on costae abaxially of sparse glands and dense, tan scales, these strongly inrolled at bases, darkened and attenuate at tips; laminae between veins glabrous on both sides of blades, or abaxially with sparse appressed, glandular hairs; sori inframedial, to 1 mm diam., exindusiate.

Rhizomes suberect, caudices ca. 2.5 cm diam.; fronds to ca. 90 cm long; stipes brownish, ca. 1⁄2 the frond length, to ca. 40 cm ⫻ 4 mm, toward the bases with patent or reflexed hair-like scales to 1 cm long; blades dark green, ovate-lanceolate, to ca. 45 ⫻ 25 cm, bipinnate or bipinnate-pinnatifid at bases, deeply pinnatepinnatifid distally; rachises abaxially with scales similar to those of stipes, but shorter, 2–2.5 mm long; pinnae to ca. 20 pairs, stalked to 6 mm, to ca. 13 ⫻ 3.3 cm, broadest at bases, equilateral, distal pinnae sessile, pinnules stalked to 1 mm, serrate or shallowly lobed, pinnules of more distal pinnae sessile, serrate or entire; segments entire or often toothed, tips acute, margins sparingly ciliate with jointed hairs; veins simple or forked, to 10 pairs per segment, ending at margins above the sinuses; indument on costae abaxially of numerous tan, inrolled scales, 1–1.5 mm long jointed hairs 0.3–0.7 mm, and short glandular hairs ca. 0.1 mm, also with jointed hairs on costae and veins; laminae between veins on both sides glandular; sori inframedial, ca. 1 mm diam., indusiate, indusia persistent, tan, round-reniform, ca. 0.6–0.8 mm diam., glandular, erose or glandular-ciliate at margins.

Distribution. Terrestrial in montane rain forests; 1600 m. Mexico; Belize (reported by Rojas, 2001, but specimens not seen). Specimen Examined. Chis (Breedlove 31328, DS).

This seems most closely related to Ctenitis tonduzii (Christ) R. M. Tryon & A. F. Tryon, from Costa Rica and perhaps Nicaragua (fide Stolze, 1981: 162), differing from that in having inframedial, exindusiate sori, approximate segments, and smaller, more decidedly inrolled scales on the blades. Moran (in Davidse et al., 1995) synonymized C. tonduzii under C. nigrovenia, but C. baulensis differs from C. nigrovenia in the more or less glabrous blade surfaces, inframedial sori, and approximate segments.

Distribution. Terrestrial in montane rain forests, in ravines and depressions; 1150–1600 m. Mexico. Specimens Examined. Chis (Breedlove 14968, DS, ENCB, US, 22331, DS, MEXU, NY; 38909, DS, 56218, CAS, ENCB, NY).

Ctenitis bullata is closely related to C. strigilosa and is distinguished by its larger glandular indusia, inrolled scales 2–2.5 mm long on the rachises, similar scales (1–1.5 mm long), jointed hairs 0.3–0.7 mm long, glands along the costae abaxially, scattered jointed hairs 0.5–0.8 mm long adaxially on the glandular costae, and scattered jointed hairs and rare glandular hairs on both sides of the blades, between and on the veins.

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3. CTENITIS CHIAPASENSIS

4. CTENITIS EQUESTRIS

Ctenitis chiapasensis (Christ) A. R. Sm., Proc. Calif. Acad. Sci., ser. 4, 40: 229. 1975. Figs. 95C, D.

Ctenitis equestris (Kunze) Ching, Sunyatsenia 5: 250. 1940.

Aspidium chiapasense Christ, Bull. Herb. Boissier, se´r. 2, 5: 727. 1905. Dryopteris chiapasensis (Christ) C. Chr., Index Filic. 257. 1905. Type. Mexico. Chiapas: Baduitz, Mu¨nch 117 (P, not found; isotypes DS! frag. NY!, US!).

Rhizomes suberect, caudices ca. 1.5 cm diam.; fronds to ca. 35 cm long; stipes stramineous to tan, ca. 1⁄2 the frond length, to 15 cm ⫻ 0.5–1 mm, bases darkened and with castaneous, lanceolate scales ca. 5 ⫻ 1 mm, distally the scales scattered, lighter, and rather inconspicuous; blades green, deltate-ovate, to 18 ⫻ 12 cm, bipinnate-pinnatifid to nearly tripinnate proximally, becoming bipinnate and then pinnate-pinnatifid distally; rachises abaxially with scattered, tan, subclathrate, lanceolate to ovate-lanceolate scales to ca. 2 mm, also with dense jointed hairs ca. 0.3 mm long and with cylindrical glands; pinnae 6–8 pairs, upcurved, proximal pair the longest, inequilaterally deltate, to 8 ⫻ 5 cm, stalked ca. 5 mm, basiscopically enlarged, middle pinnae equilateral, becoming sessile and then adnate distally; segments entire, rounded at tips, margins with jointed hairs; veins mostly simple, occasionally forked, reaching the margins above the sinuses; indument on costae and costules abaxially of scattered scales (like those of rachises) and cylindrical glands, jointed hairs few or absent; laminae between veins of yellowish, cylindrical glands ca. 0.1 mm long on both sides of blades; sori inframedial, exindusiate.

Aspidium equestre Kunze, Linnaea 18: 347. 1844. Dryopteris equestris (Kunze) C. Chr., Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 8, 6: 54. 1920. Type. Mexico. Leibold 42 (LZ, destroyed; isotype B!; Leibold 42bis at HBG–photo LL!).

Rhizomes erect, stout, caudices to ca. 4 cm diam.; fronds large, to ca. 150 cm long or more; stipes stramineous to tan, ca. 1⁄3–1⁄2 the frond length, 30–75 cm ⫻ 3–8 mm, bases with dense brown to orange-brown scales 8–15 ⫻ 0.8–1.5 mm, above the bases sparsely to moderately scaly, the scales lanceolate, 3–6 mm long, clathrate, black-walled in middle with pale margins; blades green to yellow-green, deltate, mostly 3–4-pinnate-pinnatifid proximally, 2–3-pinnate-pinnatifid distally, to 25–80 ⫻ 30–90 cm; rachises abaxially with scales similar to those of stipes but shorter, especially concentrated at pinna bases; pinnae ca. 15 pairs, proximal pair the largest, stalked to 2–8 cm, 17–50 ⫻ 12–40 cm, inequilaterally deltate, pinnules much more developed basiscopically, largest pinnules with pinnulets pinnate to pinnatisect; segments entire to crenulate, rounded to acute at tips, margins ciliate with usually minute hairs 0.1–0.2 mm; veins simple or forked, reaching margins above the sinuses; indument on costae and costules abaxially of scattered, orangish, red-brown, or brown-black, often clathrate and iridescent, ovate to ovate-lanceolate, flat or vaulted scales 1–2 ⫻ 0.3–0.5 mm, also sometimes with jointed hairs 0.1 mm long; laminae between veins glabrous on both sides, or abaxially with appressed glandular hairs 0.1–0.3 mm, occasionally with minute erect glands or glandular hairs 0.1 mm long; sori medial, exindusiate or with small, caducous indusia, these often hidden in mature sori. From other bipinnate-pinnatifid species of Ctenitis, C. equestris differs by the brown rhizome scales, flattish, dark, clathrate costal scales, and veins reaching the ciliate segment margins. The most similar species in Mexico are C. excelsa, which differs in the eciliate segment margins, and C. mexicana (q.v.). These three species and C. sloanei, together with a number of additional species from Central America and South America, form a related group.

Distribution. Terrestrial or possibly epipetric on calcareous rocks in montane rain forests; elevation unknown. Known only from the type collection. Ctenitis chiapasensis differs from other bipinnate-pinnatifid or more divided species by the small blades, exindusiate sori, and densely glandular blades on both surfaces. The nearest relative is perhaps Ctenitis chiriquiana (C. Chr.) Lellinger, from Costa Rica and Panama.

Distribution. Terrestrial in wet montane forests; 700–2700 m. Mexico; Guat, Hond, Salv, Nic, CR, Pan. Key to the Mexican Varieties of Ctenitis equestris 1. Scales of rachises and pinna axils brown-black, linearlanceolate, subentire, scattered, vaulted, spreading, not concealing the axes. ....................................................... 4a. var. equestris. 1. Scales of rachises and pinna axils orangish tan, ovate, erose-denticulate to short-ciliate, flat, appressed, imbricate, concealing the axes. .........................................4b. var. erosa.

4a. Ctenitis equestris var. equestris

Figs. 96C, D.

Lastrea ciliata Liebm., Mexic. Bregn. 273 (reprint 121). 1849. Type. Mexico. Veracruz: San Antonio Huatusco, Liebmann s.n. [Pl. Mex. 2394] (C!–2 sheets; frag. US!).

CTENITIS ERINACEA Polypodium alsophiloides Liebm., Mexic. Bregn. 208 (reprint 56). 1849. Lectotype (chosen by Smith, 1981: 80). Mexico. Oaxaca: “Trapiche de la Concepcion,” Liebmann s.n. [Pl. Mex. 2406, Fl. Mex. 199] (C! photo UC!). Aspidium bourgeaui E. Fourn., Mexic. Pl. 1: 98. 1872 [“bourgaei”]. Type. Mexico. [Veracruz:] “In Valle Cordobensi,” Bourgeau 1839 (P!, photo UC!; isotypes BM, C-photo NY!; isotype US! frag. UC!). Aspidium scabriusculum Davenp., Bot. Gaz. 21: 255. 1896, hom. illeg., non Mett. in Salomon, 1883. Dryopteris davenportii C. Chr., Index Filic. 260. 1905. Type. Mexico. Veracruz: Above Orizaba, Pringle 6132 (GH; isotypes B, K, LL!, MEXU!–3 sheets, NY!, P!, S, UC!, US!). Dryopteris equestris (Kunze) C. Chr. var. mentiens C. Chr., Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 8, 6: 56. 1920. Lectotype (inferentially chosen by Stolze, Fieldiana, Bot., n.s., 6: 147. 1981. Guatemala. Alta Verapaz: Coba´n, Tu¨rckheim II.2118 (US!).

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but the costae abaxially have many scales that are decidedly inrolled and rather broad, more like C. melanosticta or perhaps C. interjecta. The sori are exindusiate. If not a hybrid involving some combination of these three species (some sporangia appear abortive), it may represent an undescribed taxon. 4b. Ctenitis equestris var. erosa Stolze, Amer. Fern J. 67: 41. 1977. Type. Guatemala. San Marcos: Cerro Tumbador, Williams et al. 23084 (F; isotypes US!–2 sheets). Figs. 96E, F. Differs from var. equestris by the scales of rachises and pinna axils orangish tan, ovate, erose-denticulate to short-ciliate, flat, appressed, imbricate, concealing the axes; 2n⫽82 (Oax).

Differing from var. erosa in the scales of rachises and pinna axils brown-black, linear-lanceolate, subentire, scattered, vaulted, spreading, not concealing the axes.

Distribution. Terrestrial in wet montane forests; 1850–2700 m. Mexico; Guat, Hond. Distribution. Terrestrial in wet montane forests; 700–2300 m. Mexico; Guat, Hond, Salv, Nic, CR, Pan. Cited from Belize (Moran in Davidse et al., 1995: 198) on the basis of Schipp 276 (MO), but this gathering in UC is Ctenitis excelsa. Selected Specimens Examined. Chis (Breedlove 22309, NY). Dgo (Tenorio L. 761, MEXU, UC). Gro (Lorea 2411, FCME, IEB). Hgo (Flores 202, ASU, IEB, XAL). Jal (Morones 282, NY). Me´x (Hinton 7361, ARIZ, LL, NY, TEX, US). Mich (Pringle 13919, CAS, US). Mor (Lyonnet 776, DS, NY, US). Nay (McVaugh 18841, NY). Oax (Mickel 6014, CAS, UC). Rev (Flores Palacios 738, 777, UC). Sin (Lehto 24358, ASU, ENCB). Tam (Sharp 50229, US). Ver (Copeland herb. 20, BM, MICH, UC).

The types of Polypodium alsophiloides and Aspidium bourgeaui were placed in synonymy under var. erosa by Mickel and Beitel (1988), but we now believe these are both referable to var. equestris. The type of P. alsophiloides was from 3000 feet, much below the elevational range of var. erosa; moreover, the type of A. bourgeaui has the costal scales typical of var. equestris. The sole specimen seen from Durango lacks marginal hairs and so will not key to this species. The two specimens cited from Isla Socorro are also atypical in being densely glandular between the veins, on both abaxial and adaxial surfaces, and in the smaller ultimate and penultimate segments. Fink 37 (MEXU), from Veracruz, is unusual in the relatively large, persistent indusia. Breedlove 57212 (CAS), from near the Guatemalan border in Chiapas, is perhaps most like C. equestris in size and dissection,

Selected Specimens Examined. Chis (Breedlove 34700, MEXU p.p.; Little & Sharp 9938, UC, US; Martı´nez S. 19631, MEXU). Gro (Lorea 3430, FCME). Hgo (Sa´nchez Mejorada 224, MEXU). Oax (Mickel 4164, 4377, 5352, NY, UC; Smith 504, UC).

In Mexico this well marked variety occurs at higher elevations than var. equestris. Citation of var. erosa from Michoaca´n by Dı´az-Barriga & Palacios-Rios (1992) is based on Martı´nez L. 924 (ENCB!, IEB), which is var. equestris. The collection cited from Hidalgo is poor and incomplete, but seems to be var. erosa. Inclusion of Veracruz in the range by Mickel and Beitel (1988) and by Palacios Rios (1992) is the result of misidentification of the type of Aspidium bourgeaui.

5. CTENITIS ERINACEA Ctenitis erinacea A. R. Sm., sp. nov. Type. El Salvador. Chalatenango: East slope of Los Esesmiles, ca. 2160 m, Tucker 1140 (holotype UC!; isotypes LL!, US!). Figs. 94A, B. A C. hemsleyana stipitibus rachidibus et costis perspicue squamatioribus squamis persistentibus usque 1.2 cm longis, frondibus grandioribus, 1 ⫹ m longis, pari proximali pinnarum grandioribus, usque 25 ⫻ 15 cm, pinnulis basiscopicis usque 10 cm longis, indusiis persistentibus fulvis ad marginem fimbriatis differt. (L., erinaceus, hedgehog, alluding to the to numerous, persistent, dark, spreading, bristle-like scales on the stipes, rachises, and costae.)

Rhizomes erect, stout, caudices 1.5–3.5 cm diam.; fronds 1 m or more long; stipes light to dark brown, nearly equalling or slightly

224

CTENITIS

exceeding the blades, to 75 cm ⫻ 2–5 mm, throughout with dark brown to blackish, spreading scales 5–12 ⫻ 0.3–0.8 mm, these strongly inrolled and clathrate at the bases; blades dark green, deltate, tripartite, (14–)22–60 ⫻ (8–)16–45 cm, proximal pinna pair pinnate-pinnatisect, exaggerated basiscopically, suprabasal pinnae merely pinnatisect; rachises abaxially with dense, dark, hair-like scales to 8 mm long, and sparse to dense jointed hairs, 0.1 mm long; pinnae 7–15(–20) pairs, proximal pair pinnatepinnatisect, inequilaterally deltate, to 25 ⫻ 15 cm, stalked to 5 mm, basiscopically enlarged, largest basiscopic pinnules to 10 cm; segments subentire to crenulate, often curved, acute at tips, margins eciliate; veins unbranched, ending close to margins above the sinuses; indument on costae abaxially of numerous, dark brown, inrolled scales to 6 mm long and jointed hairs mostly 0.1 mm long; laminae between veins glabrous or sparsely hairy on both sides, hairs 0.1 mm, somewhat appressed; sori inframedial or subcostular, with indusia tan, fimbriate at margins.

6. CTENITIS EXCELSA Ctenitis excelsa (Desv.) Proctor, Rhodora 63: 34. 1961. Figs. 92J-M. Polypodium excelsum Desv., Me´m. Soc. Linn. Paris 6: 243. 1827. Dryopteris excelsa (Desv.) C. Chr., Index Filic. 264. 1905. Type. West Indies. Locality and collector unknown (P, photo UC!, frag. B).

Rhizomes erect, stout, caudices to 3 cm diam.; fronds large, mostly 100–250 cm long or more; stipes stramineous, ca. 1⁄3–1⁄2 the frond length, to 75 ⫹ cm ⫻ 10 mm, bases with a dense mat of orangish, linear or filiform scales 10–30 ⫻ 0.2–0.6 mm; blades dark green, deltate, 3–4-pinnate-pinnatifid proximally, 2pinnate-pinnatifid distally, to 75 ⫻ 60 cm; rachises abaxially glabrescent or with sparse linear scales like those of stipes, but shorter; pinnae to ca. 15 pairs, proximal pairs the longest, stalked to 9 cm, 20–55 ⫻ 10–30 cm, basiscopically enlarged, largest basiscopic pinnules to 20 ⫻ 10 cm, middle pinnae equilateral, pinnae becoming sessile distally; segments entire to crenulate, often subfalcate, acutish at tips, margins eciliate; veins simple or forked, reaching the margins above the sinuses; indument on costae abaxially of spreading to appressed, dark brown, clathrate, lanceolate, plane to saccate scales 10–30 ⫻ 0.2–0.5 mm, also with a few glandular hairs 0.1 mm; laminae between veins glabrous on both sides, or nearly so; sori medial, exindusiate.

Distribution. Terrestrial in cloud forests; 1300–2000 m. Mexico; Guat, Hond, Salv. Specimens Examined. Chis (Breedlove 23193, DS, MEXU, 34700, DS p.p.; Croat 46382, MO, UC; Gliessman CH-6, DS, MEXU, TEX; Magan˜a 284, MEXU). Hgo (Gimate L. 957, CHAPA, MEXU, NY). Oax (Wendt et al. 5028, CAS, CHAPA). Pue (Ventura A. 384, ARIZ, ASU, ENCB, IEB, NY, TEX). Ver (Go´mez P. & Riba 355, XAL). Guat (Steyermark 29854, US). Hond (Molina R. 459, US; Standley & Williams 773, US; Molina R. et al. 31402, US). Salv (Seiler 479, NY, 890, UC; Croat 42398, CAS, MEXU).

Ctenitis erinacea is similar to C. hemsleyana, but differs in having stipes, rachises, and costae more decidedly and persistently scaly, with scales to 12 mm long; fronds larger, 1 m long or more; proximal pair of pinnae larger, to 25 ⫻ 15 cm, the basiscopic pinnules to 10 cm long; and indusia present, persistent, tan, fimbriate at the margins. Ctenitis erinacea is similar in scaliness to C. bullata and C. ursina, but differs from both of those by the tripartite blades. Ctenitis erinacea apparently grows at higher elevations than C. hemsleyana. The sole specimen seen from Veracruz differs in having very small indusia (and so resembles C. hemsleyana), but it is like C. erinacea in size of blades and very reduced pinnules at the basiscopic bases of the proximal pinnae.

Distribution. Terrestrial in wet forests; 50–800(–1400) m. Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; L Ant. Selected Specimens Examined. Chis (Croat 43846, CAS; Martı´nez S. 8138, NY). Gro (Lorea 2919, FCME, IEB; Tenorio L. 462, ENCB, MEXU). Oax (Hammel & Merello 15548, UC). Ver (Orcutt 3214, US; Purpus 2932, BM, NY, UC, 4253, US;).

Ctenitis excelsa is similar to Ctenitis equestris, but differs in the rhizome apices and stipe bases with a dense mat of orangish, linear or filiform scales 10–30 ⫻ 0.2–0.6 mm, ultimate segments oblong, oblique, often subfalcate and acutish at the tips, segment margins eciliate, sori exindusiate, and occurrence at lower elevations. From C. mexicana it can be distinguished by the darker, more decidedly clathrate scales in pinna axils and the veins ending at the segment margins, or nearly so. From other flat-scaled, bipinnate-pinnatifid (or more divided) taxa, C. excelsa can be

CTENITIS HEMSLEYANA

distinguished by the eciliate blade margins and orangish filiform rhizome scales.

7. CTENITIS GRISEBACHII Ctenitis grisebachii (Baker) Ching, Sunyatsenia 5: 250. 1940. Figs. 94E, F. Nephrodium grisebachii Baker in Hooker & Baker, Syn. Fil. 285. 1867. Dryopteris grisebachii (Baker) Kuntze, Revis. Gen. Pl. 2: 812. 1891. Type. Cuba. [Oriente:] Wright 1055 (K!–3 sheets; isotype US!). Ctenitis molinae Stolze, Amer. Fern J. 67: 40. 1977. Type. Guatemala. Chimaltenango: Volca´n Fuego, Steyermark 52120 (US!–2 sheets; isotype F).

Rhizomes suberect, stout, caudices to 3 cm diam.; fronds large, to 150 cm long or more; stipes stramineous to brown, ca. 1⁄3–1⁄2 the frond length, to 60⫹ cm ⫻ 7 mm, throughout with brown stiff spreading lustrous, hair-like, flat scales 5–9 ⫻ 0.2–1 mm, underlain by glandular hairs 0.1 mm long and sometimes a few jointed hairs; blades light green, deltate-ovate, 2–3-pinnatepinnatifid, to ca. 75 ⫻ 70 cm; rachises abaxially with scales similar to those of stipes but shorter and less numerous; pinnae to ca. 15 pairs, proximal pair the longest, to 38 ⫻ 12 cm; segments entire to crenulate, rounded to acute at tips, margins with jointed hairs 0.1–0.5 mm long; veins simple or the basal ones sometimes forked, ending at the margins above the sinuses; indument on costae and costules abaxially of flat, brown scales 0.8–1 ⫻ 0.3– 0.5 mm, also with appressed jointed hairs 0.1–0.3 mm long and small glands 0.1 mm on costae, costules, and veins, adaxially with sparse hairs 0.3 mm on veins; laminae between veins on both sides with appressed glandular hairs 0.1–0.2 mm long, or glabrescent adaxially; sori medial, with indusia brown to tan, 0.5– 0.8 mm wide, persistent, glandular; 2n⫽164 (Jam).

Distribution. Terrestrial in wet montane forests; 1300–1950 m. Mexico; Guat, Hond, Nic, CR; Cuba, Jam, Hisp. Specimens Examined. Chis (Breedlove 32211, DS; Herna´ndez M. 513, MEXU, NY; Miranda 6973, MEXU). Gro (Lorea 1216, 1219, FCME). Ver (Finck 56, NY; Mu¨ller 1843, NY; Ventura A. 10731, ARIZ, XAL).

This species is similar in size to C. equestris, but differs in the segment margins with jointed hairs 0.1–0.5 mm long; stipe and costal scales narrower, lustrous, stiff, and uniformly dark brown;

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costules with glandular hairs and flat, brown scales 0.8–1 ⫻ 0.3– 0.5 mm; pinnae glandular on laminae and veins abaxially; sparse jointed hairs 0.3 mm on veins adaxially; and sori with brown to tan, persistent indusia 0.5–0.8 mm wide. The combination of the ciliate segment margins, flat costal scales, dark brown, flat, stiff, spreading stipe scales, and large, persistent, glandular indusia distinguishes C. grisebachii from other bipinnate-pinnatifid (or more divided) species in Mexico. Citation of this species from Nayarit by Mickel and Beitel (1988) was based on misidentified specimens of Ctenitis equestris.

8. CTENITIS HEMSLEYANA Ctenitis hemsleyana (Baker ex Hemsl.) Copel., Gen. Fil. 124. 1947. Figs. 96A, B. Polypodium hemsleyanum Baker ex Hemsl., Biol. Cent.-Amer., Bot. 3: 660, t. 108. 1885. Dryopteris hemsleyana (Baker ex Hemsl.) C. Chr., Index Filic. 270. 1905. Type. Guatemala. Chilasco´, Godman & Salvin s.n. (K!). Ctenitis costaricensis R. C. Moran, Brenesia 21: 450, f. 2. 1983. Type. Costa Rica. Cartago: Tablazo, Go´mez 21295 (CR; isotypes BM, F, G, MO, US!).

Rhizomes erect, stout, caudices 1.5–3.5 cm diam.; fronds (25–)50–90 cm long; stipes light to dark brown, nearly equalling the blades, to 45 cm ⫻ 2–4 mm, bases with dark brown to black, spreading scales 5–10 ⫻ 0.3–0.8 mm, these strongly inrolled and clathrate at the bases; blades dark green, deltate, tripartite, proximal pinna pair pinnate-pinnatisect, exaggerated basiscopically, suprabasal pinnae merely pinnatisect, (14–)25–40 ⫻ (8–)16–40 cm; rachises abaxially with scattered to dense, dark, hair-like scales 3–4 mm long and sparse to dense jointed hairs 0.1 mm long; pinnae 7–12(–16) pairs, proximal pair bipinnatepinnatisect, inequilaterally deltate, the longest (5–)10–16 ⫻ (3–) 8–12 cm, stalked to 5 mm, basiscopically enlarged, largest basiscopic pinnules to 8 ⫻ 1.8 cm, suprabasal pinnae pinnatisect; segments subentire to crenulate, often curved, acute at tips, margins eciliate; veins unbranched, ending very close to margins above the sinuses; indument on costae abaxially of scattered, dark brown, inrolled scales 1–1.5(–2) mm long and jointed hairs 0.1– 0.3 mm long; laminae between veins glabrous on both sides; sori inframedial or subcostular, exindusiate, or nearly so.

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CTENITIS

acute at tips, margins eciliate; veins unbranched, ending just short of the margins above the sinuses; indument on costae and costules abaxially of scattered, light tan, flattish to basally incurled scales to 1.5 ⫻ 0.3 mm, also with a few glandular hairs 0.1– 0.2 mm; laminae between veins glabrous on both sides, or with sparse glands abaxially; sori medial, with very small (0.2–0.3 mm diam.) tan indusia, often hidden in mature sori, indusia minutely glandular at the margins, or indusia seemingly absent.

Distribution. Terrestrial in wet montane forests; 900–1900 m. Mexico; Guat, Hond, Salv, Nic, CR, Pan. Selected Specimens Examined. Chis (Breedlove 35186, DS, MEXU; Croat 46382, UC). Hgo (Luna et al. 809, FCME). Oax (Mickel 952, 6608, ENCB, NY). Pue (Rzedowski 29994, ENCB, NY, 32425, ARIZ, CHAPA, ENCB). Ver (Ortiz J. 13, CHAPA, ENCB; Purpus 6060, UC, US).

The exaggerated basal pinnae in otherwise pinnatepinnatisect blades and the strongly inrolled scales distinguish C. hemsleyana from all other Mexican Ctenitis except C. erinacea. From that species, C. hemsleyana differs in having smaller fronds, with usually fewer than 10 pinna pairs above the proximal pinnae, and exindusiate sori (indusia to 0.5 mm diam. in C. erinacea). Ctenitis erinacea also has more elongate blades above the proximal pinnae with more pinna pairs (12 or more) that are at right angles to the rachis. A few specimens from Veracruz, e.g., Ramı´rez 687 (XAL), have very small, ciliate-margined indusia, rather scaly stipes and rachises, and reduced pinnules at the bases of proximal pinnae and thus vary in the direction of C. erinacea.

9. CTENITIS INTERJECTA Ctenitis interjecta (C. Chr.) Ching, Sunyatsenia 5: 250. 1940. Figs. 95A, B. Dryopteris interjecta C. Chr., Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 8, 6: 43. 1920. Type. Guatemala. Alta Verapaz: Cubilquitz, Tu¨rckheim s.n. [Donn. Sm. 8049] (US!; isotypes B, NY!, P, US!).

Rhizomes suberect to erect, caudices stout, to 3 cm diam.; fronds to ca. 100 cm long; stipes tan, ca. 1⁄3–1⁄2 the frond length, to ca. 40 cm ⫻ 5 mm, bases with a dense mat of orangish tan, non-clathrate, linear, twisted, tangled, and flaccid scales 10–25 ⫻ 0.2–0.5 mm; blades green, broadly ovate, 2-pinnate-pinnatisect, proximal pinna pair only slightly exaggerated basiscopically, 30– 50⫹ ⫻ 30–40 cm; rachises abaxially with scattered tan scales to ca. 2 ⫻ 0.3 mm, also with a few jointed hairs and some even shorter glandular hairs ca. 0.1–0.2 mm long; pinnae ca. 10–12 pairs, 15–25 ⫻ 6–8 cm, the proximal and longest pairs stalked to 1.5 cm, pinnules to ca. 6 ⫻ 1.8 cm, the largest pinnatifid to pinnatisect; segments entire to minutely toothed, rounded to

Distribution. Terrestrial in wet forests; 150–1000(–1450) m. Mexico; Guat, Bel, Hond, CR (Standley & Valerio 46461, CR). Specimens Examined. Chis (Breedlove 34078, DS; Martı´nez S. 17172, MEXU, 24928, UC; Palacios-Rios 2844A, UC, XAL). Oax (Hallberg 1609, NY; Herna´ndez G. 1250, CHAPA, NY; Mickel 7164, 7176, NY). Ver (Nee & Calzada 22752, NY).

Ctenitis interjecta is similar to C. equestris, but differs in the rhizome apices and stipe bases with dense mat of orangish, linear or filiform scales, eciliate ultimate segments, veins ending before reaching the margins, rachis and costal scales tan, concolorous, not heavily clathrate, somewhat vaulted and with erosedenticulate margins, and pinnae adaxially with sparse, jointed hairs 0.3–0.5 mm long on the veins. It also occurs at generally lower elevations than C. equestris. From C. melanosticta, C. interjecta differs in the orangish rhizome scales 0.1–0.3 mm wide, tan flat costal scales, and exindusiate sori (or indusia small and caducous). From C. excelsa, with which it agrees in having eciliate segment margins, C. interjecta differs in the proximal pinnae stalked usually less than 1 cm (vs. 3–10 cm) and not exaggerated basiscopically, pinnules sessile or often partially adnate (vs. shortstalked), and lighter colored, narrower scales not conspicuously aggregated in the pinna axils. Ventura A. 10731 (XAL), cited as this species by Palacios-Rios (1992), is C. grisebachii.

10. CTENITIS LANCEOLATA Ctenitis lanceolata (Baker) A. R. Sm., Proc. Calif. Acad. Sci., ser. 4, 40: 229. 1975. Figs. 93G, H. Nephrodium lanceolatum Baker in Hooker & Baker, Syn. Fil., ed. 2, 498. 1874. Dryopteris lanceolata (Baker) Kuntze, Revis. Gen. Pl. 2: 813. 1891. Type. Guatemala. Alta Verapaz: Coba´n, Salvin & Godman s.n. (K!).

CTENITIS MELANOSTICTA Nephrodium tricholepis Baker in Hemsley, Biol. Cent.-Amer., Bot. 3: 651. 1885. Dryopteris tricholepis (Baker) C. Chr., Index Filic. 298. 1905. Dryopteris lanceolata (Baker) Kuntze var. tricholepis (Baker) C. Chr., Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 7, 10: 107. 1913. Type. Guatemala. Alta Verapaz: Coba´n, Salvin s.n. (K; frag. US!). Dryopteris lanceolata (Baker) Kuntze var. deltoideolanceolata C. Chr., Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 7, 10: 107. 1920. Type. Guatemala. Alta Verapaz: Coba´n, Tu¨rckheim II.1621 (US!).

Rhizomes suberect, caudices to ca. 1 cm diam.; fronds to ca. 25(–50) cm long; stipes brownish, ca. 1⁄3–1⁄2 the frond length, to 7(–18) cm ⫻ 1 mm, bases and also distal parts with dark brown to blackish, clathrate scales to 5 ⫻ 0.5 mm; blades thinly herbaceous to chartaceous, drying dark green or brownish, lanceolate, bipinnate to bipinnate-pinnatifid, to 15 ⫻ 4(–8) cm; rachises abaxially with hair-like scales to 2 mm long; pinnae to ca. 15 pairs, proximal pairs slightly reduced or not, short-stalked, 3–4 ⫻ 1–1.5 cm, pinnules oblong, oblique, often deeply and obtusely lobed at bases, lobes of acroscopic side larger, basiscopic side subentire to shallowly lobed, decurrent onto costa; segments entire, rounded to acute at tips, margins eciliate; veins simple or forked, not reaching the margins; indument on costae abaxially of tan to blackish scales with strongly inrolled bases and hairlike tips; laminae between veins glabrous or nearly so on both sides of blades; sori medial, with large tan, persistent, glabrous indusia ca. 1 mm diam.

227

Aspidium melanostictum Kunze, Linnaea 13: 148. 1839. Dryopteris melanosticta (Kunze) Kuntze, Revis. Gen. Pl. 2: 813. 1891. Type. Mexico. [Veracruz:] Chiconquiaco, Schiede [765] (isotypes B!, LE!). Aspidium extensum Fe´e, Me´m. Foug. 5: 294. 1852, hom. illeg., non Blume, 1828. Aspidium expansum Fe´e, Me´m. Foug. 10: 42. 1865, hom. illeg., non Willd., 1810, nec Desv., 1811. Type. Mexico. Linden 6457 [an error for Galeotti 6457] (BR!, P?).

Rhizomes suberect, stout, caudices to 3⫹ cm diam.; fronds to 130 cm long; stipes stramineous to brownish, ca. 1⁄3–1⁄2 the frond length, to 60 cm ⫻ 8 mm, bases with dark brown, linear scales 7–15 ⫻ 0.5–1 mm, distally the scales reduced, hair-like, 1–2 ⫻ 0.1–0.5 mm; blades dark green, deltate, 2–3-pinnate-pinnatifid proximally, Ⳳ equilateral, proximal pinnae only slightly if at all exaggerated basiscopically, 60–80 ⫻ 40–60 cm; rachises abaxially with brown hair-like scales 1–2 mm long; pinnae to ca. 15 pairs, proximal pair stalked to ca. 10 mm, 3-pinnate-pinnatifid, 17–30 ⫻ 9–17 cm, narrowed proximally, pinnules ca. 4–10 ⫻ 1.5–2 cm, bases strongly adnate and decurrent, especially distally, on the pinnae and toward blade apices; segments crenulate to falcately lobed or incised, acute at tips, margins eciliate; veins simple or forked, ending just short of the margins above the sinuses; indument on costae and costules abaxially of scattered to dense light tan to brownish scales 0.5–2 mm long, scales with strongly inrolled margins, also the costae and costules with scattered to dense glandular hairs; laminae between veins glabrous on both sides; sori medial, with large persistent indusia, these with dark red-brown centers and tan borders, 0.8–1 mm wide, ciliateglandular; 2n⫽82 (Chis).

Distribution. Terrestrial or epipetric in montane rain forests; 1150–1250 m. Mexico; Guat. Specimens Examined. Chis (Breedlove 38989a, DS, 56244, CAS).

This differs from other small, pinnate-pinnatifid and bipinnate species of Ctenitis by the large indusia, blades lacking glandular hairs abaxially, and the blackish, strongly clathrate costal scales. The blades are dark green and tend to be fully bipinnate proximally and the pinnules obtusely lobed. Nearest allies are uncertain.

11. CTENITIS MELANOSTICTA Ctenitis melanosticta (Kunze) Copel., Gen. Fil. 124. 1947. Figs. 92F–H.

Distribution. Terrestrial or epipetric in wet lowland and montane forests; 100–1600 m. Mexico; Guat, Bel, Hond, Salv?, Nic, CR. Selected Specimens Examined. Chis (Spellman et al. 176, UC). Hgo (Mickel 623, ISC). Oax (Mickel 7280, UC). Pue (Ramı´rez 30, 32, XAL). Qro (Rzedowski 46563, IEB, MEXU). Tam (Dressler 1950, MEXU, UC). SLP (Pringle 3825, LL, MEXU, UC, US). Tab (Cowan 3943, ENCB, MEXU, TEX, UC). Ver (Copeland herb. 21, UC, US).

Among the highly dissected (bipinnate-pinnatifid or more divided) species of Ctenitis, C. melanosticta is distinct by its darkcentered, large indusia. The scales of C. melanosticta are darker, narrower, more inrolled, and more spreading than in C. equestris and allies (q.v.).

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CTENITIS

12. CTENITIS MEXICANA

portion) scales on the costae and costules abaxially, and veins ending well before the segment margins (as seen adaxially).

Ctenitis mexicana A. R. Sm., sp. nov. Type. Mexico. Veracruz: Mpio. Yecuatla, plan de Almansa, 600 m, Ventura A. 3308 (holotype NY!; isotypes ENCB!, MEXU!, XALU). Figs. 95J, K.

13. CTENITIS MICROCHLAENA

A C. equestri var. erosa indusiis carentibus, paleis planioribus et pallidioribus (stramineis et non clathratis praeter basaliter) costarum, venis adaxialiter ante marginas segmentorum terminantibus differt. (Latinization of the adjectival form of Mexico, where the species is restricted.)

Rhizomes unknown, probably erect, stout; fronds large, to 200 cm long; stipes tan to brown, ca. 1⁄3–1⁄2 the frond length, 30–50+ cm ⫻ 5–8 mm, bases with dense brown scales 5–12 ⫻ 0.5–1 mm, above the bases sparsely to moderately scaly, the scales ovate, appressed, 0.5–2 mm long, non-clathrate; blades green, deltate, mostly 3–4 pinnate-pinnatifid proximally, 2–3-pinnatepinnatifid distally, to 100⫹ ⫻ 60⫹ cm; rachises abaxially with scales similar to those of stipes but shorter, especially concentrated at pinna bases; pinnae ca. 15 pairs, proximal pair the largest, stalked to ca. 5 cm, to 50 ⫻ 30 cm, inequilaterally deltate, pinnules much more developed basiscopically, pinnules pinnate, pinnulets pinnate to pinnatifid; segments entire, acute at tips, margins with minute scattered hairs 0.1–0.15 mm; veins simple or forked, ending well before the margins; indument on costae and costules abaxially of scattered, appressed, stramineous, nonclathrate (or weakly clathrate at center-base), not iridescent, ovate to ovate-lanceolate, flat scales 0.5–0.15 ⫻ 0.3–0.8 mm, also with a few jointed hairs 0.1–0.2 mm long; laminae between veins glabrous on both sides, or abaxially with appressed glandular hairs 0.1–0.2 mm; sori medial, exindusiate.

Ctenitis microchlaena (Fe´e) Stolze, Fieldiana, Bot., n.s., 27: 9. 1991. Figs. 93C, D. Aspidium microchlaena Fe´e, Me´m. Foug. 8: 102. 1857. Dryopteris microchlaena (Fe´e) C. Chr., Index Filic. 278. 1905. Type. Mexico. [Veracruz:] Orizaba, Schaffner 459 (not found at P; isotype K!). Aspidium microcarpon Fe´e, Me´m. Foug. 8: 105. 1857, hom. illeg., non Blume, 1828. Type. Mexico. [Veracruz:] pre`s de Cordoba, Schaffner 214 (not located). Aspidium karstenii A. Braun, Ind. Sem. Hort. Berol. App. 3. 1867. Dryopteris karstenii (A. Braun) C. Chr., Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 7, 10: 98. 1913. Type. Cultivated Hort. berol. from spores of plant collected in Venezuela. Karsten s.n. (B, frag. BM; isotype K).

Rhizomes suberect to erect, stout, caudices to 4 cm; fronds 70–100 cm long; stipes stramineous to brownish, ca. 1⁄3 the frond length, to 30(–40) ⫻ 5 mm, bases with a dense mat of orangebrown, non-clathrate, hair-like scales 10–20 ⫻ 0.5–1 mm; blades green, pinnate-pinnatifid, ovate-lanceolate, 50–70 ⫻ 15–20 cm; rachises abaxially with scattered brown, non-clathrate scales mostly 1–2 mm long, also with dense jointed hairs 0.2–0.5 mm; pinnae 22–33 pairs, the proximal ones stalked 1–3 mm, 7–12 (–16) ⫻ 1.5–2.5 cm, broadest at the bases, deeply incised to about 1–2 mm from costae; segments entire or faintly toothed near the rounded tips, not or only slightly falcate, margins eciliate; veins simple, 6–10 pairs per segment, ending at the margins above the sinuses; indument on costae abaxially of scattered, tan to reddish, flat to twisted, lanceolate scales, also of short glandular hairs 0.1– 0.2 mm long; lamina between veins glabrescent or with spreading to appressed glandular hairs abaxially, margins ciliate with jointed hairs 0.1–0.3 mm long; sori medial to supramedial, indusiate, the indusia thin, tan, glabrous or minutely hairy.

Distribution. Terrestrial in lower montane and montane rain forests; 600–1800 m. Mexico. Specimens Examined. Pue (Ventura A. 496, CHAPA, ENCB, IEB, MEXU, NY, XAL). Qro (Rzedowski 46577, IEB, UC). Ver (Ballesteros & Ballesteros 356, XAL; Sousa 3494, CAS).

Ctenitis mexicana is most similar to C. equestris in size and dissection of blades, and in having minute hairs along segment margins, but is distinguished by the absence of indusia, flatter, lighter colored (stramineous and non-clathrate, except in basal

Distribution. Terrestrial in montane rain forests; 1000–1500 m. Mexico; Hond, Nic (Rojas, 2001), CR; Col, Ven, Peru. Specimens Examined. Chis (Breedlove 27456, 32934, DS, MEXU, NY, 33083, DS, 38041, DS, MEXU, 49045, CAS, ENCB; Palacios-Rios 2904, UC). Ver (Ventura A. 5423, CHAPA, ENCB, MEXU, NY).

CTENITIS REFULGENS

229

This is most similar to C. submarginalis, differing from that in the rather large, persistent indusia and the non-apiculate segments that are rounded at the tips. A collection from the Eastern Highlands of Chiapas (Breedlove 33084, DS), Mpio. Ocosingo, 800 m, appears intermediate between this species and possibly C. interjecta or C. equestris. It has large indusia, bipinnate fronds at the blade bases, and pinnules adnate nearly their width (or more) with crenate to dentate margins. The sporangia appear to be abortive, and it seems likely that the specimen represents a hybrid in the C. submarginalissloanei group, between a pinnate-pinnatifid species and a more dissected species.

14. CTENITIS NIGROVENIA Ctenitis nigrovenia (Christ) Copel., Gen. Fil. 124. 1947. Figs. 92D, E. Nephrodium nigrovenium Christ in Donnell Smith, Bot. Gaz. 20: 545. 1895. Dryopteris nigrovenia (Christ) C. Chr., Index Filic. 279. 1905. Type. Honduras. Santa Ba´rbara: San Pedro Sula, Thieme s.n. [Donn. Sm. 5646] (P!; isotype US!).

Rhizomes short-creeping to suberect, caudices 1.5–3 cm diam.; fronds clumped or somewhat spaced, 50–100 cm long; stipes stramineous to tan, 1⁄4–1⁄2 the frond length, 15–30 cm ⫻ 2–3 mm, bases with spreading to lax brown scales 4–5 ⫻ 1–2 mm, distally the scales becoming somewhat inrolled, clathrate, 1–2 mm long, underlain with dense glandular hairs; blades green, pinnatepinnatisect, ovate, 30–50 ⫻ 16–24 cm; rachises abaxially with scattered dark brown clathrate scales mostly 0.5–2 mm long; pinnae 10–15 pairs, the proximal ones stalked to 3 mm, 10–15 ⫻ 1.5–3 cm, deeply incised to about 1 mm from costae, broadest in the middle, usually somewhat narrowed at their bases; segments crenulate, slightly falcate, acute to sometimes apiculate at the apices, margins eciliate; veins simple, ending at the margins (or nearly so), in small teeth, above the sinuses; indument on costae abaxially of scattered dark brown, flat or basally inrolled scales 0.8–2 mm long, costae also with appressed to spreading, yellowish glandular hairs 0.1–0.2 mm; laminae between veins with scattered yellowish to tan glandular hairs 0.1–0.3 mm long on laminae (also with similar glandular hairs on veins) adaxially glabrous, margins eciliate; sori medial, exindusiate, but often with yellowish, glandular hairs (paraphyses) arising from receptacles.

Distribution. Terrestrial in wet montane forests; 50–120 m. Mexico; Guat, Hond, Salv (Seiler 817, NY), CR, Pan; Col, Ven, Peru, Bol. Also cited from Nicaragua by Moran (in Davidse et al., 1995). Selected Specimens Examined. Chis (Breedlove 33888, CHAPA, DS, LL, NY, US, 42648, DS, atypical). Oax (Herna´ndez G. 1425, CHAPA, NY, 1786, CAS, CHAPA, NY; Mickel 7215, ENCB, UC, US). Qro (Rzedowski 43285, IEB). SLP (Pringle 3957, LL, MEXU, NY, UC, US). Ver (Delgadillo 12275, GUADA, UC).

Among the pinnate-pinnatifid species in Mexico, the combination of veins reaching the margins and the lack of marginal hairs will distinguish Ctenitis nigrovenia. These characters are also found in C. thelypteroides and C. baulensis, which see for differences. Breedlove 42648 (DS, MEXU), from Chiapas, is atypical in having eglandular costae and blade tissue abaxially, costal scales less abruptly tapered into hair-like tips, and more deeply incised pinnae. In its scales and glabrous blades it is more like C. baulensis.

15. CTENITIS REFULGENS Ctenitis refulgens (Klotzsch ex Mett.) C. Chr. ex Vareschi, Fl. Venez. 1(1): 404. 1969. Figs. 93A, B. Phegopteris refulgens Klotzsch ex Mett., Ann. Sci. Nat. Bot., se´r. 5, 2: 240. 1864. Polypodium refulgens (Klotzsch ex Mett.) Hook. & Baker, Syn. Fil. 307. 1867. Dryopteris refulgens (Klotzsch) C. Chr., Index Filic. 288. 1905. Lectotype (chosen by Lellinger, Proc. Biol. Soc. Wash. 89: 721. 1977). Colombia. Choco´: Mouth of Rı´o Nercua, Schott 7 (US; presumed isolectotype B).

Rhizomes short-creeping to ascending, stout, caudices 2–3 cm diam.; fronds 60–100 cm long; stipes stramineous to tan, ca. 1⁄5–2⁄5 the frond length, to 15–45 cm ⫻ 4–6 mm, along their whole length with rather dense hair-like scales, scales reddish brown, 4–10 ⫻ 0.2–0.3 mm, entire, saccate basally; blades dark green, pinnate-pinnatifid, broadly ovate, 40–60 ⫻ 23–35 cm; rachises abaxially with spreading hair-like scales like those of stipes, but shorter and less numerous; pinnae to ca. 15 pairs, sessile or proximal ones stalked to 3 mm, 12–20 ⫻ 2–4 cm, incised 1⁄3–2⁄3 their width, proximal pinna segments somewhat reduced; segments entire, 6–7 mm wide, rounded to acute at tips, margins ciliate with

230

CTENITIS

jointed hairs 0.1–0.2 mm long; veins simple, 7–12 pairs per segment, proximal 1 or usually 2(–3) pairs from adjacent segments running to sinuses; indument on costae abaxially of reddish brown, spreading, hair-like scales 2–3 mm long, also with a few glands less than 0.1 mm long; laminae between veins glabrous on both sides, or with scattered appressed, yellowish glands 0.1 mm abaxially; sori medial, exindusiate.

Distribution. Terrestrial in lowland rain forests; 350 m. Mexico; Guat (Lundell 18108, LL), Pan; Col, Ven, Guy, Sur, Fr Gui, Col, Ec, Peru, n Braz, Bol. Specimens Examined. Chis (Breedlove 33376, DS, MEXU, NY).

This is similar to C. microchlaena, but differs in the longer, broader, less incised pinnae, stipes more densely scaly along their whole length, rachises and costae abaxially with rather dense reddish brown, stiff, hair-like scales, broader segments 6–7 mm wide, basalmost 1–2(–3) pairs of veins from adjacent segments running to the sinus, and exindusiate sori.

16. CTENITIS SALVINII Ctenitis salvinii (Baker) Stolze, Amer. Fern J. 67: 43. 1977. Figs. 93E, F. Nephrodium salvinii Baker in Hooker & Baker, Syn. Fil. 274. 1867. Dryopteris salvinii (Baker) Kuntze, Revis. Gen. Pl. 2: 813. 1891. Type. Guatemala. Choctum, Salvin & Godman s.n. (K!). Aspidium lindenii Kuhn, Linnaea 36: 116. 1869. Dryopteris lindenii (Kuhn) Kuntze, Revis. Gen. Pl. 2: 813. 1891. Ctenitis lindenii (Kuhn) A. R. Sm., Proc. Calif. Acad. Sci., ser. 4, 40: 229. 1975. Type. Mexico. Tabasco: Teapa, Linden 1489 (B, photos BM!, NY!; isotypes K!, P, W!, frag. BM!).

Rhizomes suberect to erect, caudices ca. 1–1.5(–2) cm diam.; fronds 30–60(–100) cm long; stipes stramineous to tan, ca. 1⁄3 the frond length, 8–20(–30) cm ⫻ 1–2(–3) mm, bases with spreading, castaneous, lanceolate, flattish scales ca. 4–6(–8) ⫻ 0.5–1 mm; blades thinly herbaceous, green, deeply pinnate-pinnatifid to bipinnate at the bases, ovate-lanceolate, 15–40(–60) ⫻ 6–12(–20) cm; rachises abaxially with tan basally inrolled scales 0.5–2 mm long; pinnae 15–25 pairs, the proximal ones stalked 1–3 mm, somewhat shorter than the next pair, Ⳳ equilateral, 4–10 ⫻ 1– 2 cm, broadest at the middle; segments distinctly serrate, the

largest incised halfway to costules, rounded at the tips, margins eciliate; veins simple to 1(or 2)-forked, not reaching the margins; indument on costae and sometimes costules abaxially of numerous, tan, non-clathrate, strongly inrolled (saccate) scales 0.3–1.5 mm long, adaxially the costae and costules with jointed hairs mostly 0.2–0.5 mm long; laminae between veins on both sides glabrous or sparsely glandular abaxially; sori medial with large, tan, persistent, glabrous or sparsely glandular-hairy indusia 0.5– 0.8 mm diam.

Distribution. Terrestrial or on calcareous rocks, in wet montane forests; 100–1000 m. Mexico; Guat, Bel. Selected Specimens Examined. Chis (Breedlove 48268, 49593, 57236, 57755, CAS, 57987, CAS, ENCB, NY; Carlson 2110, US). Tab (Cowan 2219, CAS, 2058, MEXU, NY). Ver (Va´zquez T. V-2454, CHAPA, CIB, NY, XAL; Wendt & Villalobos 2545, CHAPA, ENCB, NY).

Ctenitis salvinii differs from other pinnate-pinnatifid and bipinnate species by the indusiate sori, lack of glandular hairs on the laminae abaxially, lobed pinnules, and tan, non-clathrate, strongly inrolled costal scales. Along with C. lanceolata, it is one of the smallest Mexican species in the genus. Usually it grows on limestone rocks. Outside of northern and eastern Chiapas (23 collections seen) and eastern Guatemala, the species is rare. Aspidium ameristoneuron Fe´e (Me´m. Foug. 8: 104. 1857) was said by Fe´e to be from Cuba, but Christensen (1920: 44), citing a Mettenius annotation in B, thought the type likely to have been collected in “Tabasco. 1849. Jurgensen,” and possibly an earlier name for Ctenitis salvinii. Most likely, the Berlin specimen annotated by Mettenius is Linden 1489 (Linden did collect in Tabasco; as far as we know, Ju¨rgensen did not). The holotype is possibly in RB (Windisch, Amer. Fern J. 72: 57. 1982). Linden 1489 is the type of Ctenitis lindenii. Fe´e’s description of Aspidium ameristoneuron seems to fit plants treated here, but until the identity of the type is settled, we prefer to maintain traditional usage.

17. CTENITIS SLOANEI Ctenitis sloanei (Poepp. ex Spreng.) C. V. Morton, Amer. Fern J. 59: 66. 1969. Figs. 95E–H.

CTENITIS STRIGILOSA Polypodium sloanei Poepp. ex Spreng., Syst. Veg., ed. 16, 4(1): 59. 1827. Type. Cuba. Near Cahoba, Poeppig s.n. (LZ destroyed, frag. BM; isotypes B, HBG, L, P, frag. US!).

Rhizomes erect, caudices 4–5 cm diam.; fronds 1–1.5 m long; stipes stramineous to tan, 1⁄3–1⁄2 the frond length, to 70 cm ⫻ 10 mm, bases with dense, orangish to reddish tan, woolly, tangled, hair-like scales mostly 10–40 ⫻ 0.5–1 mm; blades chartaceous, yellowish green, deltate, 4-pinnate-pinnatifid at bases, 2-pinnate-pinnatifid distally, broadly deltate, 50–100 ⫻ 40–120 cm; rachises abaxially with scattered, stramineous to tan, lanceolate, flattened to somewhat inrolled scales mostly 1–2 mm long; pinnae to ca. 12 pairs, proximal ones stalked to 4 cm, the largest to 65 ⫻ 25 cm, basiscopically enlarged (unequally deltate, basiscopic pinnules longer than corresponding acroscopic ones), the largest basiscopic pinnules to 30 ⫻ 18 cm; segments entire or the largest ones serrate, subfalcate, rounded or subacute at tips, margins glandular-ciliate with hairs 0.1–0.2 mm; veins simple or 1forked, reaching margins above the sinuses; indument on costae and costules abaxially of tan, lanceolate, flattish or sometimes basally inrolled, appressed scales 1–2 ⫻ 0.2–0.5 mm, most numerous in axils of pinnae and pinnules; laminae on and between veins abaxially with yellowish cylindrical and capitate glands ca. 0.1 mm long, adaxially glabrous or with sparse glands or glandular hairs; sori medial, exindusiate or with small, tan, fugacious, glandular indusia, often hidden in mature sori, sporangia often bearing a small glandular hair ⬍ 0.1 mm long below annuli; 2n⫽82 (Jam), 164 (Trin).

231

sloanei cited by Palacios-Rios, 1992, from Veracruz as this species are C. excelsa.

18. CTENITIS STRIGILOSA Ctenitis strigilosa (Davenp.) Copel., Gen. Fil. 125. 1947. Figs. 94K, L. Aspidium strigilosum Davenp., Bot. Gaz. 256. 1896. Dryopteris strigilosa (Davenp.) C. Chr., Index Filic. 295. 1905. Type. Mexico. Veracruz: Barranca of Metlac, Pringle 6077 (GH!; isotypes MEXU!, NY!, UC!, US!). Dryopteris strigilosa (Davenp.) C. Chr. var. cookii Maxon ex C. Chr., Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 7, 10: 105. 1913. Type. Guatemala. Alta Verapaz: Finca Sepacuite, Cook & Griggs 507 (US!).

Rhizomes suberect to erect, caudices ca. 1.5 cm diam.; fronds 25–40 cm long; stipes brownish, 1⁄3–1⁄2 the frond length, 7–20 cm ⫻ 1–1.5 mm, bases with dark, spreading, lustrous, hair-like scales 5–9 ⫻ 0.2–1 mm, scales slightly inrolled at the margins, underlain by dense glandular hairs 0.1–0.2 mm long; blades chartaceous, green, bipinnate at the bases, deeply pinnate-pinnatisect distally, ovate, to ca. 20 ⫻ 10 cm; rachises abaxially with somewhat inrolled, dark brown scales 3–5 mm long, underlain with dense jointed hairs 0.1–0.3 mm long; pinnae to ca. 10–12 pairs, sessile or stalked to ca. 3 mm, to ca. 5 ⫻ 2 cm, the pinnatisect ones incised to within 1 mm of costae; segments entire or the larger ones minutely toothed, especially near their tips, which are sometimes apiculate, margins with jointed hairs to 0.3 mm; veins simple, reaching the margins above the sinuses; indument on costae abaxially of inrolled scales 2–3 mm long, these underlain by dense glandular and jointed hairs 0.1–0.3 mm; laminae between veins on both sides with dense to scattered glandular hairs mostly ca. 0.1 mm long; sori inframedial to subcostular throughout segments, with tan, persistent, glandular indusia 0.3–0.5 mm diam.

Distribution. Terrestrial in lowland rain forests; 350 m. USA (Fla); Mexico; Nic, CR, Pan; Cuba, Jam, Hisp, L Ant; Col, Ven, Trin, Ec, Peru, Bol. Also cited from Honduras and El Salvador by Moran (in Davidse et al., 1995), but the identity of these needs verification. Specimens Examined. Chis (Breedlove 33125, DS, MEXU, NY).

This differs from C. equestris and C. mexicana by the narrower, orangish to reddish tan stipe bases and rhizome scales, similar to those found in C. interjecta and C. excelsa, and also by the glandular blade surfaces abaxially. Ctenitis interjecta differs by the more nearly equilateral proximal pinnae, while C. excelsa differs by the eciliate segment margins. Both collections of C.

Distribution. Terrestrial or on calcareous cliffs in wet pine-oak forests, lower montane rain forests; 550–1600 m. Mexico; Guat. Specimens Examined. Chis (Breedlove 33050, DS, 47795, 48304, CAS, 57179, ENCB, CAS). Oax (Schultes & Reko 275, US; Ingela 95-M032, NY). Ver (Copeland herb. 19, BM, MEXU, MICH, UC, US).

232

CTENITIS

Ctenitis strigilosa is distinguished by the pinnate-pinnatifid to usually barely bipinnate blades, numerous dark hair-like scales on the stipes and rachises, small indusia, and densely glandular rachises, costae, veins, and laminae (both sides of blades). It is closely related to C. bullata, which see.

1091, ISC, US). Qro (Rzedowski 46569, IEB, XAL). Ver (Finck 36, MEXU; Langman 3431, MEXU, US; Mu¨ller 142, NY; Purpus 16477, US; Va´squez T. 369, XAL).

Ctenitis submarginalis (Langsd. & Fisch.) Ching, Sunyatsenia 5: 250. 1940. Figs. 92A–C.

From other pinnate-pinnatifid species, C. submarginalis differs by the exindusiate sori or sori with very small, ephemeral indusia, reddish, flattened (vs. inrolled) costal scales, and pinnae incised 3⁄4 their width or more. All but one of the specimens cited by Smith (1981: 84) from Chiapas as this species are C. microchlaena, a close relative (which see). In Mexico, C. submarginalis is relatively common only in Veracruz; elsewhere it is rarely collected or seemingly absent.

Polypodium submarginale Langsd. & Fisch., Pl. Voy. Russes Monde 1: 12, t. 13. 1810. Dryopteris submarginalis (Langsd. & Fisch.) C. Chr., Index Filic. 296. 1905. Type. Brazil. “Insula Catharinae,” s. coll. s.n. (LE).

20. CTENITIS THELYPTEROIDES

19. CTENITIS SUBMARGINALIS

Rhizomes erect, stout, caudices to 3 cm diam.; fronds to ca. 100 cm long; stipes stramineous to tan, 1⁄4–2⁄5 the frond length, 20–30(–45) cm ⫻ 3–5 mm, scales at bases brown or orangebrown, linear, 10–20 ⫻ 0.5–1.5 mm; blades green, 1-pinnatepinnatifid, oblong-lanceolate, ca. 50–70 ⫻ 20–35 cm; rachises abaxially with twisted, flat scales 1–4 ⫻ 0.1–0.3 mm; pinnae to ca. 20–25 pairs, the proximal ones stalked to ca. 2(–6) mm, the distal ones sessile, mostly 10–18 ⫻ 1.5–2.5(–3) cm, linear-oblong, not basiscopically enlarged, incised 1–3 mm from costae; segments entire or minutely crenulate toward tips, often apiculate, margins ciliate with jointed hairs to 0.5 mm long; veins simple, 8–15 pairs per segment, reaching the margins above the sinuses or the lowermost pair from adjacent segments running to the sinus; indument on costae abaxially of scattered, twisted, appressed, subentire or denticulate scales 1–4 ⫻ 0.1–0.3 mm and also minute, glandular hairs less than 0.1 mm long; laminae between veins abaxially with a few lax hair-like scales 0.1–0.3 mm long, also with appressed to spreading, yellowish, cylindrical or capitate glandular hairs less than 0.1 mm long; sori medial, exindusiate, or indusia very small, minutely ciliate, caducous or generally completely hidden by mature sporangia; 2n⫽82 (USA).

Distribution. Terrestrial in wet montane forests; 1000–1600 m. USA (Fla, La); Mexico; Guat, Hond, Nic, CR; Hisp; Col, Ven, Ec, Peru, Braz, Parag, n Arg, Uru. Selected Specimens Examined. Chis (Breedlove 31352, DS). Hgo (Copeland herb. 18, MEXU, UC, US; Moore 5087, UC, US). Oax (Mickel

Ctenitis thelypteroides A. R. Sm., Proc. Calif. Acad. Sci., ser. 4, 40: 215, f. 2A–D. 1975. Type. Mexico. Chiapas: Mpio. Las Margaritas, western side of Laguna Miramar E of San Quintı´n, Breedlove 33280 (DS! isotypes MEXU!, NY!). Figs. 93J, K. Rhizomes short-creeping to obliquely erect, to 4 mm diam., mostly obscured by stipe bases; fronds to ca. 60 cm long; stipes tan, ca. 1⁄2 the frond length, to 28 cm ⫻ 2 mm, bases with castaneous, spreading, lanceolate scales to 5 ⫻ 1 mm; blades green, pinnate-pinnatifid, ovate, to ca. 30 ⫻ 16 cm; rachises abaxially with scales similar to those of stipe bases but smaller; pinnae to 15 pairs, proximal ones stalked to 2 mm, to 9 ⫻ 2 cm, deeply incised to within 1 mm of costae, proximal pinnae somewhat narrowed at their bases; segments entire or minutely toothed, tips acute to obtuse, margins eciliate; veins simple, to 8 pairs per segment, reaching the margins above the sinuses; indument on costae abaxially of castaneous, linear-lanceolate, flattened, entire scales to 1.5 mm, also bearing minute glands ca. 0.1 mm; laminae between veins abaxially glabrous or with inconspicuous, scattered, yellow, appressed, tubular glands, glabrous adaxially; sori medial, mostly confined to tips of segments, with tan, persistent glabrous indusia ca. 0.3 mm diam.

Distribution. Terrestrial in lowland rain forests; 200 m. Mexico; Guat. Specimen Examined. Chis (Sohns 1696, US).

CULCITA

This differs from C. nigrovenia, its closest relative, by the smaller, indusiate sori localized at the tips of the segments and the proximally narrowed pinnae. Stolze (1981: 155) and Moran (in Davidse et al., 1995: 199) reduced C. thelypteroides to synonymy under C. nigrovenia, but the two species seem adequately distinct to us.

21. CTENITIS URSINA Ctenitis ursina A. R. Sm., Proc. Calif. Acad. Sci., ser. 4, 40: 216, f. 3C–E. 1975. Type. Mexico. Chiapas: Mpio. La Trinitaria, E of Lago Tziscao, Monte Bello National Park, Breedlove 32274 (DS!–2 sheets). Figs. 94G, H, J. Rhizomes not known, but probably suberect, stout; fronds large, to ca. 130 cm long or more; stipes tan, 1⁄3–1⁄2 the frond length, to ca. 50⫹ ⫻ 1 cm, bases and also distal part very densely covered with spreading, lustrous, brown, lanceolate scales to 10 ⫻ 1 mm; blades dark green, 3-pinnate-pinnatifid, broadly deltate, to 75 ⫻ 70 cm; rachises abaxially with scales similar to those of stipes, but smaller and lighter colored; pinnae ca. 10–12 pairs, proximal ones stalked 1 cm, to ca. 37 ⫻ 13 cm, equilateral, with pinnules to ca. 20 pairs per pinna, to 6 ⫻ 1.5 cm, adnate and increasingly decurrent towards pinna apices, slightly reduced at pinna bases; segments entire to deeply serrate, tips acute, margins eciliate; veins mostly simple, ending before the margins; indument on costae and costules abaxially of scales similar to those of rachises, but smaller and less dense; laminae between veins glabrous on both sides; sori medial, indusiate, indusia persistent, tan with slightly darker centers, erose to glandular-ciliate on margins.

233

This is said by Fe´e to be arborescent with a spiny trunk; Fournier (1872) treated it as a synonym of Ctenitis equestris. Christensen (1920: 125) regarded it as of uncertain identity, perhaps a species of Alsophila. Aspidium obtusilobum Fe´e, Me´m. Foug. 8: 105. 1857, hom. illeg., non Willd., 1810. Dryopteris huatuscensis C. Chr., Index Filic. 271. 1905. Type. Mexico. Veracruz: Huatusco, Schaffner 105 (P?); Smith (1981: 84) cited a possible authentic specimen, Schaffner s.n. (P!), without locality data. This name was treated by Smith (1981) and also by Mickel and Beitel (1988) as a synonym of Ctenitis submarginalis, but the holotype has apparently not been seen by anyone in the last century. It is also possible that the name is a synonym of C. microchlaena, a species not recognized in the earlier floristic accounts for Chiapas and Oaxaca.

31 . C U L C I T A Culcita C. Presl, Suppl. Tent. Pterid. 135, t. 5, f. 5. 1836. Type: Culcita macrocarpa C. Presl [⬅ Dicksonia culcita L’He´r.]. Terrestrial; rhizomes horizontal, stout, short, not forming trunks, apices with dense, long, golden or castaneous hairs; fronds large, monomorphic; stipes hairy; blades quadripinnate, glabrous to slightly hairy, coriaceous; veins free; sori marginal, indusia bivalvate, lower valve fitting inside outer (upper) valve, outer valve greenish, inner valve whitish to tan; sporangia with oblique annuli; spores tetrahedral; x⫽58, 66. Culcita is a genus of two species and is distinct from Dicksonia and Cibotium by its 4–5-pinnate blades and erect stems not forming a distinct trunk. Culcita has a sole neotropical species, and one species in Macaronesia (Madeira). Seven similar species from Polynesia, Australia, and Malesia are now generally included in the genus Calochlaena (White & Turner, 1988). The New World species occurs primarily in evergreen cloud forests at high elevations. References Gastony, G. J. 1981. Spore morphology in the Dicksoniaceae. I. The genera Cystodium, Thyrsopteris, and Culcita. Amer. J. Bot. 68: 808–819; Maxon, W. R. 1922. The genus Culcita. J. Wash. Acad. Sci. 12: 454–460; White, R. A. & M. D. Turner. 1988. Calochlaena, a new genus of dicksonioid ferns. Amer. Fern J. 78: 86–95.

1. CULCITA CONIIFOLIA Distribution. Terrestrial in montane rain forests; 1300 m. Mexico. Specimen Examined. Chis (Breedlove 35261, DS).

This is closely related to C. melanosticta, differing by the densely scaly stipes and by the less distinctly bicolored indusia. Unplaced Names Phegopteris inaequalis Fe´e, Me´m. Foug. 8: 91. 1857. Syntypes. Mexico. [Veracruz:] Huatusco, Schaffner 240, 241a; Totutla, Schaffner 241b (RB, cited by Windisch, 1982).

Culcita coniifolia (Hook.) Maxon, Annual Rep. Board Regents Smithsonian Inst. 1911: 488, t. 13C. 1912. Figs. 97A–D. Dicksonia coniifolia Hook., Sp. Fil. 1: 70, t. 24A. 1844. Type. Venezuela. Caracas, Linden 538 (K!; isotype BR!, BM! photo ex BR, UC! photo ex BR).

Rhizome apices and stipe base hairs dense, castaneous, 1–3 cm long; fronds 1–3 m long; blades broadly deltate, 4–5-pinnatepinnatifid; ultimate segments oblique; axes hairy (especially in axils) to glabrescent; indusial valves about twice as wide as long; 2n⫽132 (CR).

234

CYATHEA Trichipteris excelsa C. Presl ⫽ T. corcovadensis (Raddi) Copel. ⬅ Cyathea corcovadensis (Raddi) Domin. (Additional synonymy in Barrington, 1978.) Sometimes treated as Trichopteris. For discussion of viewpoints of Trichipteris vs. Trichopteris, see Gastony (Amer. J. Bot. 66: 1258-1260. 1979), Morton (Amer. Fern J. 61: 142–143. 1971), and Lellinger (1987).

Distribution. Terrestrial in cloud forests on Atlantic slope, evergreen cloud forests, rain forests, Quercus-Podocarpus forests, cloud forests with much Drimys; 2000–3000 m. Mexico; Guat, Hond, Salv, Nic, CR, Pan; Cuba, Jam, Hisp; Col, Ven, Ec, Peru, s Braz, Bol. Selected Specimens Examined. Chis (Breedlove 6117, 12922, US, 22007, CAS, MEXU, MO, NY; Martı´nez S. et al. 3285, MO; Matuda 5280, US; Mickel 1252, ENCB, NY, ISU, UC, US; Mu¨nch s.n., May 1902, US). Oax (Diggs et al. 3930, MEXU, NY; Mickel 917, ENCB, ISC, MEXU, NY, US, 4884, ENCB, MEXU, NY, UC, US).

Culcita resembles Dicksonia in the coriaceous fronds with marginal, bivalvate sori and indument of hairs but is distinct in being trunkless and having 4-pinnate (vs. 2-pinnate-pinnatifid to 3-pinnate) blades.

3 2 . C YA T H E A Cyathea Sm., Me´m. Acad. Roy. Sci. (Turin) 5: 416. 1793. Type: Cyathea arborea (L.) Sm. [⬅ Polypodium arboreum L.]. Hemitelia R. Br., Prodr. 158. 1810. Type: Hemitelia multiflora (Sm.) Spreng. ⬅ Cyathea multiflora Sm. Trichipteris C. Presl in J. Presl & C. Presl, Delic. Prag. 1: 172. 1822. Type:

Terrestrial; stems erect, usually forming massive trunks to 20 m tall, 10 cm diam. (without the occasional mantle of wiry adventitious roots), scaly at apices; fronds large, to 4 m long; stipes stout, scaly, scales conform or marginate, concolorous to bicolorous, without dark apical setae, cells of the margins lightly to strongly different from those of the center in size, shape, and orientation; blades 2–3⫹-pinnate, chartaceous to coriaceous; axes with long, curved, acicular hairs adaxially, often with bullate scales abaxially, the rachises and costae rarely with spines; veins free, simple or forked; sori round, medial, with raised receptacles, indusia arising from beneath the sori and either completely enclosing them or forming cups or flat unequal-sided saucers, or absent; sporangia with slightly oblique annuli; spores tetrahedralglobose, 64 per sporangium; x⫽69. Cyathea is a genus of about 115 species, 8 in Mexico. The scaly tree ferns have been treated as a single genus (Holttum & Sen, 1962) and as six genera (R. Tryon, 1970). We are treating them here in four genera, following Lellinger (1987) and Moran (in Davidse et al., 1995). References Barrington, D. S. 1978. A revision of the genus Trichipteris. Contr. Gray Herb. 208: 1–93; Gastony, G. J. 1979. Spore morphology in the Cyatheaceae. III. The genus Trichipteris. Amer. J. Bot. 66: 1238–1260; Holttum, R. E. & U. Sen. 1962. Morphology and classification of the tree ferns. Phytomorphology 11: 406–420; Lellinger, D. B. 1987. The disposition of Trichopteris (Cyatheaceae). Amer. Fern J. 77: 90–94; Maxon, W. R. 1909. Cyatheaceae. N. Amer. Fl. 16: 65–88; Maxon, W. R. 1922. Studies of tropical American ferns. No. 7. The North American species of Alsophila grouped with A. armata. Contr. U.S. Natl. Herb. 24: 33–46; Riba, R. 1967. Revisio´n monogra´fica del complejo Alsophila swartziana Mart. (Cyatheaceae). Anales Inst. Biol. Univ. Me´xico, Bot. 38: 61–100; Riba, R. 1969. The Alsophila swartziana complex (Cyatheaceae). Rhodora 71: 7–17; Riba, R. 1981. Cyatheaceae of Veracruz. Flora de Veracruz 17: 1–42; Tryon, R. M. 1976. A revision of the genus Cyathea. Contr. Gray Herb. 206: 19–98.

Key to the Mexican Species of Cyathea 1. Stipe scales conform, castaneous with short marginal teeth; indusia absent; rachises and costae sparsely scaly. ........................... 7. C. myosuroides. 1. Stipe scales marginate, the cells of the margin decidedly different from those of the scale center in size, shape, and orientation, not ciliate, but often erose. 2. Indusia present, globose. 3. Abaxial surfaces of penultimate and ultimate segment midribs with pale brown bullate scales and hairs; pinnules sessile to short-stalked 1–2 mm. ............................................................................................................................................................................... 4. C. fulva. 3. Abaxial surfaces of penultimate and ultimate segment midribs with scattered dark brown, generally flat scales, especially abundant in pinna axils, hairs lacking; pinnules short- to long-stalked 1–6 mm. ............................................ 3. C. divergens var. tuerckheimii. 2. Indusia absent, though sori in Cyathea costaricensis subtended by 1 to several pale bullate scales. 4. Blades with long, sharp spines on rachises and pinna axes. ........................................................................................................ 6. C. microdonta. 4. Blades without spines, or if present, limited to stipes. 5. Bullate scales on ultimate segment midribs dark brown; stipe scales concolorous, brown to dark brown; ultimate segments rounded at tips. ............................................................................................................................................................ 8. C. schiedeana. 5. Bullate scales on ultimate segment midribs pale tan to white; stipe scales bicolorous or concolorous, pale tan to whitish, often with brown centers; ultimate segments broadly to sharply acute at tips.

CYATHEA BICRENATA

235

6. Sori subtended by 1-several bullate scales; rachises and costae glabrous adaxially; penultimate segment midribs (costae) with trichomidia only; stipe spines absent. ....................................................................................................... 2. C. costaricensis. 6. Sori without subtending scales; axes with large pale scales and/or long hairs; costules with broad pale scales and/or long hairs; stipe spines small to stout. 7. Costae and costules with broad, pale scales, long hairs, and trichomidia; sori only at bases of ultimate segments; stipe scales concolorous, pale; stipe spines 1 mm long or less. .................................................................................... 5. C. godmanii. 7. Costae and costules with long hairs and trichomidia only; sori along the length of the ultimate segments; stipe scales bicolorous each with broad to narrow, dark brown central stripe and pale margins; stipe spines 1–5 mm long. ..................................................................................................................................................................................... 1. C. bicrenata.

Key to Sterile or Fragmentary Specimens of Scaly Mexican Tree Ferns 1. Blades pinnate or pinnate-pinnatifid; trunks less than 0.5 m tall. 2. Veins mostly free, only very rarely forming basal arches along the costae; margins coarsely serrate. .................................. Cnemidaria apiculata. 2. Basal veins regularly united by a transverse veinlet, forming a single series of costal areoles; margins lightly and remotely crenate-serrulate. ................................................................................................................................................................................Cnemidaria decurrens. 1. Blades 2–3-pinnate; trunks generally 2–20 m tall. 3. Blade scales setose, with few to many stiff marginal setae, usually darker in color than the scale body. 4. Scales of stipes, rachises, and costae linear, pale yellow with dark teeth (Sphaeropteris). ................................................. Sphaeropteris horrida. 4. Scales of stipes, rachises, and costae whitish with sparse, sometimes black setae or scales dark brown, subentire (Alsophila). 5. Frond axes black; larger scales lanceolate, dark brown, and more or less entire, plus small spider-like scales. ........... Alsophila salvinii. 5. Frond axes brown; larger scales whitish with black setae. 6. Costae and costules with abundant bullate scales. ...................................................................................................... Alsophila tryonorum. 6. Costae and costules with scales small, plane; bullate scales absent. ................................................................................. Alsophila firma. 3. Blade scales entire or irregularly lobed, not darkly setose (if setose, with concolorous castaneous scales)(Cyathea). 7. Costae with large flat white scales, long hairs, and pale bullate scales on costules. ............................................................ Cyathea godmannii. 7. Costae lacking large pale flat scales. 8. Rachises and costae with spines; costules with copious curved hairs to 0.5 mm; indusia absent; sori with hairs. ......................................................................................................................................................................................... Cyathea microdonta. 8. Rachises and costae lacking spines; costules with scattered hairs and scales; indusia present or absent; sori lacking hairs (except in C. myosuroides). 9. Sterile blades essentially glabrous, but fertile with curly hairs and also scales, attached proximally at the base of the sori. ...................................................................................................................................................................................... Cyathea costaricensis. 9. Sterile and fertile blades essentially the same, with scattered hairs and/or scales on costules. 10. Indument of short, stiff acicular hairs, hairs in sori, lamina grayish; stipe scales toothed, concolorous, castaneous. ................................................................................................................................................................ Cyathea myosuroides. 10. Indument of hairs and scales or only scales, lamina green; stipe scales bicolorous, brown-centered, cells near margin of different color and orientation. 11. Blades with bullate scales and hairs. 12. Indusia sphaeropteroid; abaxial segment hairs ca. 1 mm. ............................................................................. Cyathea fulva. 12. Indusia absent; abaxial segment hairs ca. 2 mm. ..................................................................................... Cyathea bicrenata. 11. Blades with sparse scales; hairs absent. 13. Scales bullate, dark brown; segment apices broadly rounded; abaxial laminar surfaces dull; pinnules sessile; indusia absent. ................................................................................................................................ Cyathea schiedeana. 13. Scales flat, castaneous, lustrous; segment apices acute; abaxial laminar surfaces lustrous; pinnules stalked; indusia globose. ............................................................................................................................................... Cyathea divergens.

1. CYATHEA BICRENATA Cyathea bicrenata Liebm., Mexic. Bregn. 289 (reprint 137). 1849. Figs. 98A–F. Alsophila bicrenata (Liebm.) E. Fourn., Mexic. Pl. 1: 134. 1872. Trichipteris bicrenata (Liebm.) R. M. Tryon, Contr. Gray Herb. 200: 44. 1970. Lectotype (chosen by Smith 1981: 237). Mexico. Oaxaca: Trapiche de la Concepcio´n to Totontepec, Liebmann s.n. [Fl. Mex. 926] (C!; isotype US!). Alsophila scabriuscula Maxon, Proc. Biol. Soc. Wash. 23: 125. 1919. Trichipteris scabriuscula (Maxon) R. M. Tryon, Contr. Gray Herb. 200: 44. 1970.

Type. Guatemala. Cubilquitz, Tu¨rckheim s.n. [Donn. Sm. 7806] (US!; isotype NY!).

Trunks 5–15 m tall, 3.5–6 cm diam.; stipes stramineous to light brown with stout, short to long spines 1–5 mm long, glabrous to densely hairy, stipe scales bicolorous, brown with very pale margins, 5–6 mm wide; fronds to 4 m long, bipinnate-pinnatifid with the segments deeply lobed; pinnae nearly sessile, stalks less than 1 cm, pinnules sessile, 2.5–3 cm wide; segments slightly ascending, bullate scales to base of segments, hairs on segment midveins abaxially; costules hairy adaxially, segment midveins gla-

236

CYATHEA

brous to only sparsely hairy adaxially; sori medial, indusia absent, paraphyses shorter than the sporangia.

Distribution. Moist forests to nearly semiarid, drier areas; 50– 1200 m. Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan. Distribution. Wet montane forests at middle elevations; 100– 1800 m. Mexico; Guat, Hond, Salv, Nic, CR, Pan. Selected Specimens Examined. Chis (Breedlove 23123, NY). Gro (Hinton 14185, NY; Wagenbreth 529, MEXU). Oax (Mickel 5106, NY). Pue (Lorea 1097, FCME; Marquez R. 755, XAL; Riba 252, MEXU). Ver (Ibarra Manriquez 918, MEXU; Rzedowski 32944, NY; Ventura A. 17255, MEXU).

Trichipteris scabriuscula has been distinguished on the basis of heavily pubescent stipes, but this character is quite variable and, we think, unreliable. The hairs and bullate scales are similar to those of C. godmanii, but the blades of C. bicrenata are slightly less dissected.

2. CYATHEA COSTARICENSIS Cyathea costaricensis (Mett. ex Kuhn) Domin, Acta Bot. Bohem. 9: 107. 1930. Figs. 99A–E. Hemitelia costaricensis Mett. ex Kuhn, Linnaea 36: 159. 1869. Trichipteris costaricensis (Mett. ex Kuhn) Barrington, Rhodora 78: 1. 1976. Lectotype (chosen by Barrington, 1978: 76). “Costa Rica et Veragua.” Warscewicz 36 (B!, frags. NY!, US!).

Trunks to 8 m tall; fronds ca. 3 m long; stipes stramineous, spines absent, scales pale, often with brown centers, hairs absent; blades bipinnate-pinnatifid, thin-textured, pinnae stalked, pinnules sessile to stalked 1–2 mm; segments dentate to shallowly lobed, slightly falcate, apices acute; rachises and costae glabrous abaxially; adaxial surfaces with costules hairy, also with whitish flat to subbullate scales, segment midveins with 1–3 hairs; sterile laminae glabrous abaxially; sori medial, subtended by 1–3 bullate scales, resembling indusia, and with sparse hairs distally, paraphyses shorter than sporangia.

Selected Specimens Examined. Chis (Breedlove 24184, DS). Gro (Hinton 10793, US). Jal (Iltis et al. 452, NY, XAL). Mich (Hinton 16262, US). Nay (Gentry et al. 19480, NY). Oax (Figueroa 89, MEXU, XAL; Mickel 5176, NY). Pue (Flores Castorena 654, ENCB). Tab (Matuda 37495, MEXU). Ver (Nee et al. 26106, NY).

Cyathea costaricensis is distinct by the smooth stramineous stipes and sori subtended by 1-3 bullate scales.

3. CYATHEA DIVERGENS VAR. TUERCKHEIMII Cyathea divergens Kunze var. tuerckheimii (Maxon) R. M. Tryon, Contr. Gray Herb. 206: 56. 1976. Figs. 99F– H, J, K. Cyathea tuerckheimii Maxon, Contr. U.S. Natl. Herb. 13: 4. 1909. Type. Guatemala. Alta Verapaz: Near Coba´n, Tu¨rckheim II 1645 (US!; isotypes F, GH, LE!, MO, NY!, P!, UC!). Cyathea juergensenii E. Fourn., Mexic. Pl. 1: 135. 1872. Type. Mexico. Oaxaca: “In cordillera Oajacensi” [Sierra San Pedro Nolasco], Ju¨rgensen 874 (P; isotype BM!, frag. US!).

Trunks to 12 m tall; fronds to 4 m long; stipes brown to tan, lightly to strongly spiny, spines dark, sharp, scales long, bicolorous, dark brown with pale erose margins, stipes very hairy with basal scurf whitish to tan; blades bipinnate-pinnatifid, pinnae stalked, 0.5–2.7 cm long, pinnules usually stalked 1–6 mm; segments obtuse, strongly falcate, crenulate, firm-textured, costae hairy adaxially, glabrous abaxially, segments glabrous on both sides but with sparse brown, lustrous, generally flat scales on segment midveins abaxially, scales most abundant in pinna axils; fertile veins simple or 1-forked; sori medial, indusia globose.

CYATHEA GODMANII

237

Distribution. Wet montane forests of Atlantic slope; 300–2500 m. Mexico; Bel, Guat, Hond, Nic.

Distribution. Wet montane forests; 800–2700(–3250) m. Mexico; Guat, Hond, Salv, Nic, CR, Pan; Col, Ven, Ec.

Selected Specimens Examined. Chis (Breedlove 25384, 29566, MEXU; Dressler 1688, MEXU, NY). Oax (Mickel 6503, NY; Rzedowski 33393, MEXU). Pue (Arreguı´n 537, NY, 540, XAL; Riba et al. 296, MEXU). Ver (Pe´rez et al. 114, MEXU; Ventura A. 9269, NY).

Selected Specimens Examined. Chis (Breedlove 34677, NY). Gro (Hinton 14274, NY; Lorea 2366, MEXU ). Hgo (Gonza´lez Quintero 1361, NY; Rzedowski 32677, XAL). Oax (Hallberg 1433, NY, US; Mickel 5604, NY). Pue (Rzedowski 23408, NY, 31843, XAL). Ver (Ventura A. 288, NY).

Cyathea divergens is distinct among Mexican species of Cyathea in its lack of costal hairs on the abaxial blade surfaces and the frequently stalked (1–6 mm) pinnules. There is often a black, tar-like fungus on the adaxial laminar surfaces. Var. divergens occurs from Costa Rica to Surinam and Peru.

Cyathea fulva is distinct from C. divergens in its laminar hairs on the abaxial costal surfaces and consistently short-stalked (1– 2 mm) pinnules.

4. CYATHEA FULVA Cyathea fulva (M. Martens & Galeotti) Fe´e, Me´m. Foug. 9: 34. 1857. Figs. 99L–P. Alsophila fulva M. Martens & Galeotti, Me´m. Foug. Mexique 78, pl. 23. 1842. Type. Mexico. Oaxaca: “Cordilliere Orientale, Talea,” Galeotti 6346 (BR!, frags. NY!, US!; isotypes BR!, P!). Cyathea delicatula Maxon, Contr. U.S. Natl. Herb. 13: 4. 1909. Type. Guatemala. Alta Verapaz: Between Tactic and Coba´n, Tu¨rckheim II. 1629 (US!; isotype GH!). See R. Tryon (1976) for more complete synonymy.

Trunks to 12 m tall, 25 cm diam.; stipes tan to stramineous, with prickles to well developed spines, scales dark brown with paler erose margins to medium brown with virtually undifferentiated margins, also with dark brown scurf of small scales and deciduous hairs; blades to 5 m long, bipinnate-pinnatifid; pinnae stalked 4–6(–11) mm, pinnules sessile except for the basal 1–3 pinnules, with stalks 1–2 mm; segments obtuse, slightly ascending, crenulate; costules with curved hairs abaxially, sparse hairs adaxially on segment midveins and abundantly hairy on costules; bullate scales pale brown on segment midveins abaxially; veins mostly 1-forked, rarely simple; sori medial, indusia globose, enclosing the sori and shattering at maturity.

5. CYATHEA GODMANII Cyathea godmanii (Hook.) Domin, Pterid. Dominica 262. 1929. Figs. 100A–F. Alsophila godmanii Hook. in Hooker & Baker, Syn. Fil. 36. 1867. Type. Guatemala. Coba´n, Salvin & Godman s.n. (K!). Alsophila mexicana Mart., Icon. Pl. Crypt. 70, t. 45. 1834, hom. illeg., non Cyathea mexicana Schldtl. & Cham., 1830. Cyathea valdecrenata Domin, Pterid. Dominica 263. 1929, nom. nov. Trichipteris mexicana (Mart.) R. M. Tryon, Contr. Gray Herb. 200: 44. 1970. Type. Mexico. Oaxaca: San Pablo de Teoxomulco, Karwinski s.n. (isotype P, not found, frag. F; L, photo P!).

Trunks to 10 m tall; stipes dark brown, muricate with small spines 1 mm or less long, densely clothed with large, pale, concolorous scales ca. 5 mm wide; blades 3–4 m long, tripinnatepinnatifid, pinnae nearly sessile, ca. 1 cm long, pinnules sessile, markedly reduced at pinna bases, broadest in the middle, ca. 2.5–3.2 cm wide, segments slightly ascending; rachises and costae with long, fine, lax hairs and conspicuous broad, pale scales, these ovate to ovate-lanceolate with finely fringed margins, scales varying to small and bullate, hairs on costules and major veins of surfaces; sori restricted to base of pinnulets (tertiary divisions), indusia absent, paraphyses longer than sporangia.

238

CYATHEA

Distribution. Wet montane forests; 1300–1900 m. Mexico; Guat, Hond, Pan. Specimens Examined. Chis (Breedlove 22648, MEXU, 29657, NY, 31957, 35251, MEXU). Gro (Lorea 1691, 4535, FCME; Martı´nez S. 4238, ENCB, XAL; Wagenbreth 530, MEXU) Oax (Karwinski s.n., P, frag. F). Unverified, Doubtful, or Mistaken Reports. Ver (reported by Mickel & Beitel, 1988, but not verified).

Distribution. Wet montane forests; 0–1700 m. Mexico; Bel, Guat, Hond, Nic, CR, Pan; Cuba, Jam, Hisp; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol. Specimens Examined. Chis (Matuda 17435, MEXU; Seler 5515, GH; Xolocotzi & Sharp X-241, US). Gro (Lorea 2861, FCME). Tab (Rovirosa 48, US). Ver (Orcutt 3195, 3194 MEXU, NY; Smith 2095, NY).

This rare species is distinct by its broad pale scales on the abaxial blade axes.

Cyathea microdonta is distinguished by its slender spines on the rachises and secondary axes, whereas in other species of Cyathea, spines, when present, are limited to the stipes.

6. CYATHEA MICRODONTA

7. CYATHEA MYOSUROIDES

Cyathea microdonta (Desv.) Domin, Pterid. Dominica 263. 1929. Figs. 100G, H, J.

Cyathea myosuroides (Liebm.) Domin, Pterid. Dominica 263. 1929. Figs. 98G, H, J, K.

Polypodium microdontum Desv., Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck. Gesammten Naturk. 5: 319. 1811. Alsophila microdonta (Desv.) Desv., Me´m. Soc. Linn. Paris 6: 319. 1827. Trichipteris microdonta (Desv.) R. M. Tryon, Contr. Gray Herb. 200: 46. 1970. Type. “America australi?” s. coll. s.n. (P!).

Alsophila myosuroides Liebm., Mexic. Bregn. 236 (reprint 84). 1849. Sphaeropteris myosuroides (Liebm.) R. M. Tryon, Contr. Gray Herb. 200: 20. 1970. Type. Mexico. Oaxaca: Chinantla, between Lacoba and Teotalcingo, Liebmann s.n. [Pl. Mex. 2081, Fl. Mex. 514, 902, 905, 906, 907, 908] (C!; isotype US!).

Trunks to 6 m tall; stipes with sharp-pointed spines, light to medium brown, scales concolorous, brown; blades 2–3 m long, bipinnate-pinnatifid, papyraceous, pinnae with stalks 0.6–0.7 cm, rachises and pinnae with sparse sharp spines; pinnules essentially sessile, stalks 1 mm, apices acute to acuminate, curved, coarsely crenate to dentate; segments densely hairy on rachises and pinna axes; adaxial surfaces densely short-hairy on pinnules, some hairs also on segment midveins; abaxial surfaces with hairs and minute bullate scales; sori medial, receptacles pilose, indusia absent; 2n⫽138 (Jam).

Trunks to 2–5 m tall, ca. 10 cm diam.; stipes dark brown, prickly, but lacking coarse spines, scales conform, castaneous, concolorous, margins with short teeth, hairs of major axes not abundant, mostly adpressed; blades 1.5–4 m long, bipinnatepinnatifid, firm to subcoriaceous; pinnae ca. 1.7 cm wide, stalks 10–14 mm long; pinnules stalked ca. 1–2 mm, lobed ca. halfway to segment midveins; veins 1-forked; costules hairy on both sides, segment hairs sparse adaxially, a few more abaxially, occasional irregularly toothed dark scales on costules abaxially; sori medial at vein forks or more commonly on acroscopic branches, receptacles pilose, indusia absent.

CYCLOPELTIS

239

Distribution. Wet montane forests; 0–1200 m. Mexico; Guat, Bel, Hond, Nic; Cuba.

Distribution. Wet montane forests; 100–1400 m. Mexico; Bel, Guat, Hond, Nic, CR, Pan; Col.

Selected Specimens Examined. Chis (Breedlove 21836, NY; Castillo C. et al. 3936, XAL). Oax (Mexia 9244, NY; Mickel 7378, MEXU, NY; Torres C. 11152, MEXU). Pue (Contreras J. 6714, MEXU). Tab (Cowan 2096, NY; Ferna´ndez N. 1053, XAL). Ver (Dorantes et al. D-2707, MEXU; Smith 2094, NY).

Selected Specimens Examined. Chis (Breedlove 21892, DS; Garcı´a F. 718, XAL). Gro (Lorea 1540, FCME; Te´llez 6160, MEXU). Oax (Conzatti 3816, MEXU; Mickel 5711, MEXU, NY). Pue (Contreras J. 6121, 6989, MEXU). Ver (Calzada 8370, XAL; Orcutt 3213, NY).

Cyathea myosuroides is distinguished by the exindusiate sori and by castaneous stipe scales with many fine, concolorous teeth. Abundant epiphyllous lichens (Aulaxina quadrangula (Stirton) R. Sant. and three sterile Asterothyriaceae; Richard Harris, pers. comm.) give a distinct pale gray color to the blades. This species and C. microdonta are the only species of tree ferns in Mexico that occur also in the West Indies.

8. CYATHEA SCHIEDEANA Cyathea schiedeana (C. Presl) Domin, Pterid. Dominica 263. 1929. Figs. 98L–P. Alsophila schiedeana C. Presl, Suppl. Tent. Pterid. 62. 1836 [based on a description of a Polypodium without binomial in Schldtl. & Cham., Linnaea 5: 609. 1830]. Trichipteris schiedeana (C. Presl) R. M. Tryon, Contr. Gray Herb. 200: 46. 1970. Type. Mexico. [Veracruz:] “Inter Huitamalco et Cuapa regionis calidae,” Schiede & Deppe 757 (B!–2 sheets, frags. GH!, US!; isotype LE!). See Barrrington (1978: 38) for additional synonyms based on non-Mexican types.

Trunks to 7 m tall, 6 cm diam.; stipes stramineous to brown, armed with slender to stout spines, scales brown to dark brown, concolorous or with narrow, pale, entire to slightly erose margins, scurfy with tiny squamules; blades 2–3 m long, bipinnatepinnatifid; rachises stramineous to brown, with occasional low spines, glabrous to scurfy; laminae firm; pinnae sessile or with stalks to 1 cm; pinnules sessile or stalked to 1 mm, 1.5–2 cm wide, crenulate to broadly dentate, segments 3–4 mm wide, rounded at tips, apices obtuse; sterile veins simple or 1-forked; costae hairy adaxially, segment midveins with 1–2 hairs near tip; costae abaxially with short hairs (variable amount) and dark brown bullate scales; sori medial on simple veins or with forks, indusia absent, paraphyses shorter than sporangia.

Unverified, Doubtful, or Mistaken Reports. Tab (Linden s.n., K, cited by Cowen, 1983, but not verified).

Among the exindusiate species of Mexican tree ferns, C. schiedeana differs by its concolorous dark brown stipe scales, dark brown bullate costal scales, and rounded segment apices. Excluded Species Cyathea marginalis (Klotzsch) Domin, Pterid. Dominica 263. 1929. Alsophila marginalis Klotzsch, Linnaea 18: 542. 1844. Sphaeropteris marginalis (Klotzsch) R. M. Tryon, Contr. Gray Herb. 200: 20. 1970. Cyathea marginalis is represented by Ju¨rgensen 915 (K), from Sierra San Pedro Nolasco, Oaxaca, but it is otherwise known only from the Guyana area. Tryon (Rhodora 74: 441–450. 1972) believed this to be an erroneous label. This number is cited by Fournier (1872) and Stolze (1974) as Cnemidaria decurrens.

33. C Y C L O P E L T I S Cyclopeltis J. Sm., Bot. Mag. 72(Compendium): 36. 1846. Type: Cyclopeltis semicordata (Sw.) J. Sm. [⬅ Polypodium semicordatum Sw.]. Terrestrial; rhizomes woody, short-creeping, scaly; fronds medium-sized, erect, monomorphic, not articulate; stipes and rachises adaxially sulcate, deciduously scaly; blades once-pinnate, reduced at base; pinnae numerous, articulate, deciduous, entire (the apical one subconform or hastate proximally), glabrous, costae rounded adaxially; veins free (to casually anastomosing); sori round, in 1–4 rows on each side of costa, indusia round, peltate, deciduous; spores bilateral, with prominently folded or winged perispore; x⫽41.

240

CYSTOPTERIS Luna 13422, GUADA, UC). Ver (Brigada Dorantes 2893, MEXU, MO, UC, XAL; Va´zquez 727, UAMIZ, XAL).

Cyclopeltis comprises six species, five in Southeast Asia, only one (widespread) in lowland wet forests of the New World tropics. Precise relationships of the genus are not entirely clear, but seem to be dryopteroid or polystichoid. Unpublished molecular work by Cranfill (pers. comm.) suggests an alliance with Lomariopsis. Cyclopeltis is sometimes placed in its own family (e.g., by Ching, 1940).

This species is easily distinguished by the round, peltate indusia and the auricles at the pinna bases overlapping the rachis.

1. CYCLOPELTIS SEMICORDATA

Cystopteris Bernh., Neues J. Bot. 1(2): 5, 26. 1806 [1805], nom. cons. Type: Cystopteris fragilis (L.) Bernh. [⬅ Polypodium fragile L.].

Cyclopeltis semicordata (Sw.) J. Sm., Bot. Mag. 72(Compendium): 36. 1846. Figs. 101A, B. Polypodium semicordatum Sw., Prodr. 132. 1788. Aspidium semicordatum (Sw.) Sw., J. Bot. (Schrader) 1800(2): 31. 1801. Dryopteris semicordata (Sw.) Kuntze, Revis. Gen. Pl. 2: 813. 1891. Polystichum semicordatum (Sw.) T. Moore, Index Fil. lxxxiv. 1857. Type. Jamaica. Swartz s.n. (S?; isotypes B-Willd. 19742, UPS).

Terrestrial; rhizomes erect to suberect; rhizome scales light brown, 8–15 ⫻ 0.8–1.5 mm, linear-attenuate, entire; fronds clumped; stipes pale brown, 14–30 cm long, 1⁄10–1⁄3 the frond length, scaly at base; blades 40–160 ⫻ 12–35 cm, 1-pinnate, lanceolate, subconform apex usually lobate at base, proximal 2–5 pinna pairs somewhat shortened and deflexed; rachises with scattered hair-like scales 3–6 mm long, abaxial surface densely puberulent with stout, minute, erect hairs 0.1 mm long; pinnae linear-lanceolate, entire or slightly undulate, 6–18 ⫻ 1–2 cm, sessile, semicordate to auriculate at the base, auricle basiscopic, overlying the brownish rachis; blades firm-membranaceous to chartaceous, glabrous; veins 3–5-forked, 1–3 proximal branches bearing sori; sori borne in 1–3 rows on each side of costa, nearer to costa than margin; indusia peltate, persistent or deciduous, 1– 2 mm diam., margin minutely ciliate; 2n⫽82 (Jam).

34. C Y ST O P T E R I S

Terrestrial or epipetric; rhizomes slender, short- to longcreeping, scaly; fronds small to medium-sized, monomorphic, not articulate, erect to spreading; stipe slender, brittle, sparsely scaly to glabrescent; blades 1–3-pinnate-pinnatifid; laminae thin, glabrous; veins free; sori abaxial, roundish, indusia hood-like, each attached on the basiscopic side and overarching the sorus, often shriveling or deciduous; paraphyses absent; spores bilateral, dark; x⫽42. Cystopteris is a temperate genus of about 10 species; C. fragilis is generally considered to be cosmopolitan, perhaps the most widespread fern species in the world, but it has not been studied critically in recent years outside of North America and Europe and is probably a complex of several taxa. There are at least three species in Mexico and probably more. Malformed spores have been found in some species, suggesting hybridization within C. fragilis s.l. Relationships of the genus are probably closest with Gymnocarpium, less so to Woodsia (Cranfill, ined.). Cystopteris is distinct by the hood-like indusia. References Blasdell, R. F. 1963. A monographic study of the fern genus Cystopteris. Mem. Torrey Bot. Club 21(4): 1–102; Pearman, R. W. 1976. A scanning electron microscopic investigation of the spores of the genus Cystopteris. Brit. Fern Gaz. 11: 221–230.

Key to the Mexican Species of Cystopteris 1. Laminae membranous, 1–2 cells thick; plants growing in the spray of waterfalls; rhizome scales fleshy, without fine hairs interspersed; veins ending in teeth. ..... 2. C. membranifolia. 1. Laminae thin, but more than 2 cells thick; plants terrestrial in mesic woods, roadbanks, streambanks; rhizome scales membranous, interspersed with fine hairs; veins usually ending in tiny emarginations. 2. Blades bipinnate to tripinnate-pinnatifid; segments oblong, 1.5–4 mm wide. .............................................. 1. C. fragilis. 2. Blades tripinnate-pinnatifid to quadripinnate; segments linear, 1 mm wide. ................................... 3. C. millefolia.

Distribution. Terrestrial along stream banks and in ravines in wet forests; 50–550 m. Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol. Selected Specimens Examined. Chis (Breedlove 58021, CAS; Gonza´lez Q. 3473, ENCB, NY; Martı´nez S. 7566, 8013, ARIZ, 15021, UC). Tab (Cowan 1720, CAS, CHAPA, ENCB, NY, 2071, CHAPA; Dı´az

1. CYSTOPTERIS FRAGILIS Cystopteris fragilis (L.) Bernh., Neues J. Bot. 1(2): 26, t. 2, f. 9. 1806 [1805]. Figs. 102A–F, 103A–E, H, J. Polypodium fragile L., Sp. Pl. 2: 1091. 1753. Type. Uncertain. See Blasdell (1963) for discussion of typification.

CYSTOPTERIS MEMBRANIFOLIA Polypodium diaphanum Bory, Voy. ˆIles Afrique 1: 328. 1804. Cystopteris fragilis (L.) Bernh. subsp. diaphana (Bory) Litard., Bull. Soc. Bot. Deux-Se`vres 23: 88. 1912. Cystopteris diaphana (Bory) Blasdell, Mem. Torrey Bot. Club 21(4): 47. 1963. Type. Re´union, Bory s.n. (P). Aspidium fragile (L.) Sw. var. fumarioides M. Martens & Galeotti, Me´m. Foug. Mexique 67. 1842. Type. Mexico. Temascal, Galeotti 6259 (BR!; isotype BR!, P!). Cystopteris acuta Fe´e, Me´m. Foug. 5: 300. 1852. Type. Mexico. [Veracruz:] Pic d’Orizaba, Galeotti 6260 (P!).

Rhizomes short-creeping, not protruding past youngest leaves; rhizome scales brown, lanceolate, lustrous, membranous, entire, mixed with fine hairs; fronds to 47 cm long, clumped; stipes 1⁄3–1⁄2 the frond length, stramineous, often reddish brown and with scales at bases, glabrous or with scattered glandular hairs; blades (6–)9–20 ⫻ (2.5–)4–8 cm, bipinnate to tripinnate-pinnatifid, ovate-lanceolate with proximal 1–2 pairs of pinnae slightly shortened; rachises glabrous to pubescent with glandular hairs; pinnae 7–15 pairs, 1–4 cm long, thin, but more than 2 cells thick; segments oblong, 1.5–4 mm wide; veins ending in small emargination or in teeth; indusia glabrous with subentire to lacerate apices; 2n⫽168 (Chis).

241

of Mexico have been (chromosomes, isozymes, etc.), a conservative treatment is followed, one that combines C. fragilis s.str. with C. diaphana. The North American taxa include diploids, allopolyploids, and various sterile hybrids, with species often having limited ranges. Cystopteris reevesiana Lellinger of the southwestern United States probably occurs in northern Mexico but is not readily distinguishable from Mexican C. fragilis and is not treated here. It is supposed to be distinguished from C. fragilis by blades bipinnate-pinnatifid to tripinnate (vs. pinnatepinnatifid to bipinnate-pinnatifid) and rhizomes usually longcreeping (vs. short-creeping). These characters do not seem to be correlated in Mexican specimens, and more work is needed to circumscribe the species. Cystopteris fragilis most closely resembles a small Dryopteris, but the hood-like indusia of the former readily distinguish it.

2. CYSTOPTERIS MEMBRANIFOLIA Cystopteris membranifolia Mickel, Amer. Fern J. 62: 93. 1972. Type. Mexico. Oaxaca: Distrito Teotitla´n, about 33 km NE of Teotitla´n del Camino on road to Huautla, Mickel 4544 (NY!; isotype US!). Figs. 102G, H, J. Rhizomes long-creeping, the tips extending beyond the stipes, sparsely clothed with dark brown, fleshy, ovate scales 2.5–3.5 ⫻ 0.3–0.8 mm; fronds 14–28 cm long, distant; stipes 4–9 cm long, ca. 1⁄3 the frond length, green to stramineous; blades pinnatepinnatifid, lanceolate or ovate-lanceolate, apices pinnatifid, membranaceous, 1–2 cells thick, glabrous; pinnae 2–3 ⫻ 1–2 cm, deltate-lanceolate, sessile; pinnules broadly oblong or elliptic, incised, apices obtuse, margins dentate; segments oblong; veins ending in teeth; sori round, indusia very small, deciduous.

Distribution. Terrestrial, often on or among rocks, on mossy ledges, fir forests, tropical deciduous forests, oak forests, montane cloud forests, pine-oak forests; (600–)1050–3400(–4050) m. Northern and western North America; Mexico; Guat, Hond, Salv, Nic, CR, Pan; Gr Ant; Col, Ven, Ec, Peru, Braz, Bol, Chile, J Fdez, Arg; Europe, Africa, Asia, Hawaii. Selected Specimens Examined. BCS (Carter et al. 2336, DS, UC). Chih (Correll & Gentry 23207, MO, UC). Chis (Breedlove 15111, DS). Coah (Wynd & Mueller 521, MO). DF (Bourgeau 251, MO). Dgo (Tenorio L. et al. 4159, MEXU). Gro (Mexia 9068, MO, UC). Gto (Ventura & Lo´pez 2758, IEB). Hgo (Arreola 5, FCME). Jal (Dı´az-Luna 6027, GUADA, UC). Me´x (Fanti 1037, ENCB, FCME, MO). Mich (Dı´azLuna 9292, GUADA, UC). Mor (Hinton 17521 bis, ENCB, MEXU). Nay (McVaugh 18888, IEB, MEXU, MICH, MO). NL (Hinton et al. 17304, MO). Oax (Mickel 5529, NY, UC). Pue (Orcutt 3983, MO). Qro (Zamudio 6894, IEB). Sin (Breedlove 44959, CAS). SLP (Palmer 176, MO, UC). Tam (Lof 705, UC). Ver (Ventura A. 17697, MEXU). Unverified, Doubtful, or Mistaken Reports. Ags (Siqueiros 4346, HUAA, and three other collections, cited by Siqueiros-Delgado & Gonza´lez-Adame, 2004, but not verified).

Until the Mexican members of the Cystopteris fragilis complex have been studied and characterized to the degree that taxa north

Distribution. Terrestrial by waterfalls on Atlantic slope, montane cloud forests; 1300–2500 m. Mexico; Guat. Selected Specimens Examined. Oax (Fraser-Jenkins s.n., 14 May 1987, UAMIZ; Mickel 6656, NY). Ver (Rhoads s.n., Mar 1899, US).

Cystopteris membranifolia is a rare species of limited distribution, a “filmy” fern found only in very humid habitats. It is very distinctive, although there are indications of a hybrid origin

242

DANAEA

in the irregular and often malformed spores. Rhizome scales are fleshy when fresh.

3. CYSTOPTERIS MILLEFOLIA Cystopteris millefolia Mickel & Tejero, Brittonia 56: 000. 2004. Type. Mexico. Me´xico: Mpio. Ocuilan, km 18, OcuilanCuernavaca, Tejero-Dı´ez 2234 (NY!; isotype IZTA!). Figs. 103F, G. Rhizomes short- to long-creeping, ca. 2 mm diam., not protruding beyond fronds; rhizome scales brown, membranous, lustrous, lanceolate, entire, ca. 1.5 mm, mixed with fine hairs; fronds 14–40 cm long, clumped; stipes 1⁄3–1⁄5 the frond length, stramineous, reddish brown and with scales to 3 mm long at the bases, mixed with hairs as on rhizomes, glabrous distally; blades lanceolate, tripinnate-pinnatifid to quadripinnate, 12–30 ⫻ 4.5– 14 cm, with proximial 2–3 pinna pairs reduced; rachises glabrous or with scattered, minute, glandular hairs in adaxial grooves; pinnae 12–16 pairs, 2–8 cm long, more than two cells thick; segments linear, 1 mm wide; veins ending in small emarginations; indusia glabrous, entire.

Terrestrial; rhizomes creeping, green, clothed with thinmargined stipules in two rows on the dorsal part of the rhizomes; roots fleshy, thick; leaf axes with swellings at pinna insertions and occasionally on the stipes; fronds dimorphic; sterile blades 1-pinnate, rarely simple (not in ours), thick-herbaceous to subcoriaceous; veins free, curving at margins into cartilaginous edge; scales on axes and costae, minute scales and trichomidia on veins and lamina; fertile fronds similar in dissection to the sterile, but often with narrower pinnae; sporangia fused into linear, multilocular (20–100 locules) synangia extending from costae to the blade margins, synangia nearly covering the abaxial surfaces; spores bilateral; x⫽40. Danaea is a genus of about 30–35 species of the American tropics, in wet montane forests at low to middle elevations. Three species occur in Mexico. The genus is not closely allied to others in the Marattiaceae and is sometimes segregated as a distinct family. Its distinguishing features include its rhizome stipules, fleshy, dimorphic fronds, jointed frond axes, and synangia extending from costa to margin. The family Marattiaceae occupies a basal position within the ferns (Hasebe et al., 1995; Pryer et al., 2001) and has a fossil record back more than 300 million years. References Lellinger, D. B. 2000. On the lectotypification of Danaea elliptica. Amer. Fern J. 90: 100–103; Underwood, L. M. 1902. American Ferns-V. A review of the genus Danaea. Bull. Torrey Bot. Club 29: 669–679; Underwood, L. M. 1909. Marattiaceae. N. Amer. Fl. 16: 17–23.

Key to the Mexican Species of Danaea

Distribution. Wet mossy rocks in forest streams, moist banks in pine forests; 2200–3000 m. Mexico. Selected Specimens Examined. Me´x (Barkley et al. 2346, LL; Barr 62-795, ARIZ; Bartholomew 2886, GH, MEXU; Beaman 2061, TEX; Flores F. 817, MEXU; Hinton 15711, TEX). Mich (Cruden 1068, GH, MICH; Hinton 13499, ARIZ, GH, LL, PH, TEX; Rzedowski 48315, MEXU). Mor (Pe´rez-Garcı´a 766, UAMIZ, cited as Cystopteris reevesiana Lellinger by Riba et al., 1996, but the cited specimen is C. millefolia).

Cystopteris millefolia is one of the most distinct of the segregates in the C. fragilis complex because of its finely divided blades.

35. DANAEA Danaea Sm., Me´m. Acad. Roy. Sci. (Turin) 5: 420. 1793, nom. cons. Type: Danaea nodosa (L.) Sm. [⬅ Acrostichum nodosum L.].

1. Sterile pinnae 1.5–2 cm wide; pinna margins strongly serrulate toward tips. ...................................................... 1. D. cuspidata. 1. Sterile pinnae 2.5–4.5 cm wide; pinna margins entire to faintly serrulate toward tips. 2. Stipes lacking nodes; pinnae 7–11 pairs, caudate, with abruptly long-attenuate undulate tip; veins 11– 15/cm. ............................................................................. 3. D. nodosa. 2. Stipes with 2–5 nodes; pinnae 3–6 pairs, acute to acuminate; veins 8–11/cm. ......................................... 2. D. elliptica.

1. DANAEA CUSPIDATA Danaea cuspidata Liebm., Mexic. Bregn. 307 (reprint 155). 1849. Type. Mexico. Oaxaca: Chinantla, between Lobani and Petlapa, Liebmann s.n. [Pl. Mex. 2184, Fl. Mex. 255, 656] (C!). Figs. 104C, D. Danaea muenchii Christ in Rosenstock, Repert. Spec. Nov. Regni. Veg. 22: 23. 1925 [“munchii”]. Type. Mexico. Chiapas: San Pablo, Mu¨nch 159 (P!; isotypes DS!, US!).

Rhizomes short-creeping, green, 7–13 mm diam.; rhizome scales dark; fronds to 120 cm tall; stipes 18–36 cm long, ca. 1⁄2 or slightly less the frond length, green-brown, with (0–)2–4 swollen nodes, with dense (especially at nodes) to remote, brown, appressed scales; blades 75–165 ⫻ 20–32 cm; pinnae 7–13 pairs, sterile pinnae 9–12(–16) ⫻ 1.5–2 cm, narrowly elliptic, bases

DANAEA NODOSA

243

obliquely cuneate or truncate, apices strongly serrulate, longacuminate; laminae dark green, with minute scales on veins and trichomidia between veins; veins free, simple or 1-forked; fertile pinnae similar in shape to those of the sterile fronds, but narrower, 4.5–6 ⫻ 0.5–0.7 cm; synangia 56–62 per pinna.

Distribution. Terrestrial in wet montane forests of Atlantic slope; 150–850 m. Mexico; Guat, Bel, Hond, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol. Distribution. Terrestrial in wet montane forests near rivers on Atlantic slope; 250–1650 m. Mexico; Guat, CR, Pan. Selected Specimens Examined. Chis (Mu¨nch 159, DS, P, US). Oax (Martı´nez C. 399, UC, US; Mickel 962, ENCB, MEXU, MICH, NY, 1552, ENCB, NY, 5653, 5888, 6456, NY, UC; Romero et al. 1007, MEXU). Ver (Lira 162, MEXU, UAMIZ, XAL).

Danaea cuspidata is distinguished by its slender, strongly serrulate pinnae. It is closely allied with or perhaps identical to D. moritziana C. Presl of South America and possibly the West Indies; the latter is the older name.

2. DANAEA ELLIPTICA Danaea elliptica Sm. in Rees, Cycl. 11: Danaea No. 2. 1808. Lectotype (chosen by Lellinger, Amer. Fern Soc. 90: 102. 2000). Jamaica. Sloane s.n. (BM-Hb. J. E. Smith cat. no. 1645.7 ex LINN). Figs. 104E–G. Rhizomes short-creeping, horizontal, green, 8–10 mm diam.; rhizome scales light or dark brown; fronds to 1 m tall; stipes 1⁄2– 2⁄3 the frond length, olive-brown, glabrous or with remote, appressed, black to brown scales, with 2–5 nodes; blades 13–35 cm wide; pinnae 3–6 pairs, sterile pinnae 9–17 ⫻ 2.5–4.5 cm, narrowly elliptic, margins subentire, bases cuneate, apices acute to acuminate; laminar scales sparse, heavier on costae; laminae chartaceous, olive-green; veins simple to 1-forked, 8–11/cm; fertile pinnae similar to sterile ones in shape, 5.5–11.5 ⫻ 1.2–2.3 cm; synangia 33–118 per pinna; 2n⫽160 (Trin).

Specimens Examined. Chis (Purpus 7247, F, NY, UC, US). Oax (Herna´ndez G. 1647, MEXU; Mickel 6407, 7203, NY, 6511, ENCB, MEXU, NY; Rzedowski 33315, MEXU; Va´zquez T. 1554, XAL). Ver (Castillo-Campos et al. 12598, XAL; Lorence & Ton 3964, MEXU).

Danaea elliptica resembles D. nodosa in the broad sterile pinnae, but the former has nodose stipes, fewer pinnae, and more widely spaced veins. There has been some confusion regarding the typification of D. elliptica, its having been lectotypified twice before (Underwood, 1902: 672; Proctor, 1985: 62), but Lellinger (2000) has carefully explained the situation.

3. DANAEA NODOSA Danaea nodosa (L.) Sm., Me´m. Acad. Roy. Sci. (Turin) 5: 420. 1793. Figs. 104A, B, H. Acrostichum nodosum L., Sp. Pl. 2: 1070. 1753. Lectotype (chosen by Proctor, Fl. L. Antill. 2: 47. 1977). Plumier, Traite´ Foug. Ame´r., pl. 108, 1705, illustrating a plant from Martinique or Haiti.

Rhizomes horizontal, short-creeping, green, 10–12 cm diam.; rhizome scales brown; fronds to 2 m tall, 27–40 cm wide; stipes ca. 1⁄2 the frond length, brown to olive-brown, with sparse, brown, appressed scales; blades pinnate, pinnae 7–11 pairs, sterile ones 14–34 ⫻ 3–4.5 cm, narrowly elliptic, slightly falcate, margins subentire or sometimes undulate, faintly serrulate near apices, bases unequally rounded or cuneate, caudate with abruptly longattenuate, undulate tips; laminae coriaceous, with minute brown blade scales, these denser along costae; veins simple or 1-forked, 11–15/cm; fertile pinnae similar in shape to sterile but narrower, to 26 ⫻ 2 cm; synangia 150–200 per pinna; 2n⫽160 (Trin).

244

DENNSTAEDTIA

appears likely that D. media is simply a juvenile of D. elata. They were probably described under different names (in the same publication, with a mention that they grow together) because D. media has nodes on the stipes (at least one of the syntypes, possibly a juvenile; see Smith, 1981), whereas D. elata does not. That juveniles often have nodes but adults more often do not is a characteristic shared with populations of D. media elsewhere in Central America. Typical D. nodosa, type from Martinique, does not have nodes on the stipe, even when juvenile, and it also differs from Mexican and Central American material in pinna shape and much bulkier rhizomes. Tuomisto and Christenhusz tentatively plan to use the name D. media for Mexican and Central American plants previously called D. nodosa. Distribution. Terrestrial in tropical lower montane and wet montane forests of Atlantic slope, evergreen rain forests, undisturbed forests near rivers; 100–600 m. Mexico; Guat, Bel, Hond, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol, Parag. Selected Specimens Examined. Chis (Breedlove 34139, DS, MEXU; Martı´nez S. et al. 25610, MEXU; Purpus 6760, MO, NY, UC, US). Oax (Hammel & Merello 15596, MO, US; Herna´ndez 1617, CHAPA, NY, 1899, CHAPA, MEXU; Mickel 6408, NY, UC, US, 6846, NY, UC; Torres C. 11136, MEXU). Ver (Dorantes 2501, MEXU, MO, XAL).

This is the largest of the Danaea species in Mexico, and, ironically, most easily distinguished by the lack of nodes on the stipe. Unplaced Names Danaea elata Liebm., Mexic. Bregn. 306 (reprint 154). 1849. Type. Mexico. Veracruz: Barranca de la Hacienda de Jovo, Liebmann s.n. [Pl. Mex. 2199, Fl. Mex. 848] (C!). Danaea media Liebm., Mexic. Bregn. 306 (reprint 154). 1849. Syntypes. Mexico. Veracruz: Barranca de la Hacienda de Jovo, Liebmann s.n. [Pl. Mex. 2185, Fl. Mex. 653, 654, 849, and 850] (C!). Tuomisto and Christenhusz (Univ. Turku, Finland) are currently studying the systematics of Danaea and have recently concluded (in litt.) that D. elata and D. media, variously treated by several authors (e.g., Underwood, 1909; Smith, 1981; Mickel & Beitel, 1988; Lellinger, 1989; Camus & Pe´rez-Garcı´a in Davidse et al, 1995), apply to the same taxon. Tuomisto and Christenhusz believe that this taxon is distinct from the widespread and polymorphic D. nodosa, a name that has been applied to Mexican material with large fronds and nodeless stipes. The types of both D. elata and D. media were collected at the same locality, and it

36 . D E N N ST A E D T I A Dennstaedtia Bernh., J. Bot. (Schrader) 1800(2): 124. 1801. Type: Dennstaedtia flaccida (G. Forst.) Bernh. [⬅ Trichomanes flaccidum G. Forst.]. Terrestrial; rhizomes creeping or less commonly ascending, usually stout, pubescent, solenostelic; fronds large, often with a bud near the base of each stipe; blades bipinnate to quadripinnate-pinnatifid, chartaceous to coriaceous, glabrous to pubescent; veins free, extending not quite to the segment margins, tips often clavate; sori marginal, without paraphyses, inner and outer indusia forming a cup- or purse-shaped sorus, indusia not greatly modified; spores tetrahedral; x⫽? Dennstaedtia is most diversified in the Old World (eastern Asia and the South Pacific), with 20 species in America. In Mexico it most closely resembles Saccoloma, from which it is distinct in the hairy rhizome and the sori at the very margin. Dennstaedtia is probably most closely related to Hypolepis, Pteridium, and Histiopteris, these four genera occupying a relatively basal position (in Dennstaedtiaceae) among the higher leptosporangiate ferns. We thank Hugo Navarrete of Pontificia Universidad Cato´lica del Ecuador, Quito, for reviewing the manuscript and offering valuable suggestions. References Navarrete, H. & B. Øllgaard. 2000. The fern genus Dennstaedtia (Dennstaedtiaceae) in Ecuador–new characters, new species and a new combination. Nordic J. Bot. 20: 319–346; Tryon, R. M. 1960. A review of the genus Dennstaedtia in America. Contr. Gray Herb. 187: 23–52.

Key to the Mexican Species of Dennstaedtia 1. Indusia 0.5–1 mm deep; hairs confined to abaxial surfaces, clear brown or tan, rarely subopaque and whitish, relatively straight and spreading; laminar axis grooves continuous, bordered by perpendicular herbaceous wings; stipe base buds absent; rhizomes very hairy, branched. 2. Basal segments of pinnules alternate and unequal in size; lamina coriaceous, lustrous on both surfaces; segments deeply and sharply toothed; indusia cylindrical to subcylindrical, 0.8 mm deep, 0.8–1 mm wide, glabrous. ................................................. 2. D. bipinnata. 2. Basal segments of pinnules opposite and equal in size; lamina herbaceous, dull on both surfaces; segments shallowly and obtusely toothed; indusia globular, 1 mm deep, 1–1.3 mm wide, slightly hairy. ............................................................................. 7. D. globulifera.

DENNSTAEDTIA BIPINNATA

245

1. Indusia 0.1–0.3 mm deep; hairs on abaxial surfaces whitish to brownish and subopaque, tortuous and spreading or straight and tightly appressed; hairs on adaxial surfaces rare to dense; laminar axis grooves interrupted at junctions; stipe buds present, sometimes forming secondary rhizomes; rhizomes glabrous or pubescent, unbranched. 3. Indusia 1–1.5 mm wide; vein ends clavate; hairs on abaxial surfaces 0.1–0.3 mm long and tightly appressed; adaxial laminae glabrous or nearly so, hairs on adaxial surfaces rare to scattered, 0.5 mm long, each with enlarged base; roots glabrous. 4. Sori contiguous along segment margins. ........................................................................................................................................... 5. D. coronata. 4. Sori distant along segment margins. 5. Fertile blades bipinnate to subtripinnate. ..................................................................................................................................1. D. auriculata. 5. Fertile blades tripinnate-pinnatifid to quadripinnate. ................................................................................................................ 4. D. cornuta. 3. Indusia 0.5–0.8 mm wide; vein ends slender; hairs on abaxial surfaces 0.8–1 mm long and spreading, straight or tortuous; laminae pubescent, hairs on adaxial surfaces scattered to dense, 0.3–0.5 mm long, slender, or hairs limited to wings of costae and major veins; roots densely pubescent or nearly glabrous in D. spinosa. 6. Frond axes spiny. ..................................................................................................................................................................................... 8. D. spinosa. 6. Frond axes not armed. 7. Pinnae stalked, alternate, not reduced proximally; hairs abundant abaxially, two types mixed on axes, somewhat tortuous hairs 0.8–1 mm long, and shorter, straight, acicular hairs 0.1–0.3 mm; major axes tan to reddish brown. ...... 3. D. cicutaria. 7. Pinnae sessile, opposite or subopposite, reduced at pinna base; hairs sparse to abundant abaxially, long (0.8–1 mm), somewhat tortuous hairs on axes grading to short (0.1 mm) clavate hairs on lamina; major axes usually stramineous. ...................................................................................................................................................................................... 6. D. distenta.

1. DENNSTAEDTIA AURICULATA

Specimens Examined. Chis (Breedlove 33693, NY, TEX, 56477, CAS; Mu¨nch 60, DS, US).

Dennstaedtia auriculata H. Navarrete & B. Øllgaard, Nordic J. Bot. 20: 337. 2000. Type. Ecuador. Prov. Pichincha, Km 21 de la carretera Quito-Sto. Domingo, Navarrete 449 (QCA; isotypes AAU, QCNE). Figs. 107D, E.

Dennstaedtia auriculata closely resembles D. arborescens (Willd.) Ekman ex Maxon of the West Indies except the former has lobate fertile segments, lacks laminar buds, and has reflexed basal pinnules, whereas the latter has entire fertile segments, laminar buds, and basal pinnules patent to ascending.

Rhizomes short-creeping, unbranched, ascending, ca. 3 cm diam., glabrous; fronds 1.5–3 m long, to 1.5 cm distant; stipes ca. 1 ⁄2 the frond length, erect to scandent, each with one bud at base; blades arching or scandent, ovate-lanceolate, bipinnate to subtripinnate; proximal pinnae usually subsessile, with basal pinnules reduced and reflexed; axes of penultimate segments without perpendicular herbaceous wings adaxially, or with them, but wing on basiscopic side not decurrent onto axis of next order; sterile vein tips clavate to punctate, ending well short of glabrous margins; blade axes glabrate to usually pubescent abaxially, hairs brownish, subappressed, ascending, somewhat curled; laminar tissue glabrous, dark green, dull, thick-herbaceous; fertile segments lobate; indusia purse-shaped to cup-shaped.

Distribution. Montane rain forests; 300–1200 m. Mexico; Guat, CR, Pan; Gr Ant; Col, Ven, Ec, Peru, Bol.

2. DENNSTAEDTIA BIPINNATA Dennstaedtia bipinnata (Cav.) Maxon, Proc. Biol. Soc. Wash. 51: 39. 1938. Figs. 105K–Q. Dicksonia bipinnata Cav., Descr. Pl., 174. 1801. Dicksonia adiantoides Humb. & Bonpl. ex Willd., Sp. Pl., ed. 4, 5(1): 488. 1810, nom. superfl. Dennstaedtia adiantoides (Humb. & Bonpl. ex Willd.) T. Moore, Index Fil. xcvii. 1857. Type. Puerto Rico. Ventenat s.n. (MA; isotype B-Willd. 20165-1!, frag. US!).

Rhizomes long-creeping, branched, 5–10 mm diam., hairy; fronds 2–3 m long, clumped; stipes 1⁄2 the blade length, lacking buds at the base; blades deltate, 1–1.5 m wide; fronds 3–4-pinnate; texture coriaceous to firm herbaceous; proximal pinnules stalked, not especially reduced, basal segments of pinnules of central pinnae alternate, unequal in size, axes of penultimate segments with wings on adaxial surfaces; sterile vein tips slender; laminae glabrous adaxially, hairs scattered abaxially, mostly at axis intersections, 1 mm long, clear brown, spreading and straight; indusia cylindrical to subcylindrical, 0.8 mm deep, 0.8–1 mm wide; 2n⫽188 (Jam).

246

DENNSTAEDTIA

Distribution. Wet montane forests; 450–1300 m. USA (Fla); Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Gr Ant; Col, Ven, Trin, Ec, Peru, Bol.

Distribution. Wet montane forests; (200–)750–1200 m. Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Gr Ant; Col, Ven, Ec, Peru, Braz, Bol, Arg.

Selected Specimens Examined. Chis (Breedlove 23292, DS; Ghiesbreght 356, NY). Hgo (Kenoyer 710, US). Oax (Campos V. 1037, NY; Mickel 6373, NY). Pue (Arreguı´n 549, XAL). Qro (Rzedowski 45205, ENCB). SLP (Pringle 3407, NY). Ver (Ventura A. 5047, 9355, NY). Yuc (Butterwick 243, LL, TEX).

Selected Specimens Examined. Chis (Breedlove 22270, 25072, NY). Hgo (Mickel 621, NY). Mich (Hinton 16267, NY). Oax (Hellwig 453, NY; Mickel 5076, NY). Pue (Boege 1219, MEXU). Qro (Rzedowski 42570, IEB, MEXU, TEX). SLP (Pringle 3824, NY). Ver (Purpus 2936, NY).

Unverified, Doubtful, or Mistaken Reports. Gro (Vela´zquez 282, FCME, cited by Lorea-Herna´ndez & Vela´zquez M., 1998, but not verified).

The lustrous, coriaceous blades and cylindrical indusia distinguish this species.

Unverified, Doubtful, or Mistaken Reports. Gro (Lorea 2923, cited by Lorea-Herna´ndez & Vela´zquez M., 1998, but not verified). Tam (reported by Lof, 1980, but not verified).

Dennstaedtia cicutaria resembles D. distenta in the pubescent blades, slender vein ends, and small sori, but is distinct in having stalked alternate pinnae reduced at pinna bases, major axes reddish brown, and mixed hair types on axes.

3. DENNSTAEDTIA CICUTARIA 4. DENNSTAEDTIA CORNUTA Dennstaedtia cicutaria (Sw.) T. Moore, Index Fil. xcvii. 1857. Figs. 106F–H, J. Dicksonia cicutaria Sw., J. Bot. (Schrader) 1800(2): 91. 1801. Type. Jamaica. Swartz s.n. (S-frag. and photo US!; isotype B-Willd. 20156!, photo GH). Dicksonia rubiginosa Kaulf., Enum. Filic. 226. 1824. Dennstaedtia rubiginosa (Kaulf.) T. Moore, Index Fil. xcvii. 1857. Type. Brazil. Rio de Janeiro: Herb. Mertens (isotype LE!, photo GH). Dicksonia umbrosa Liebm., Mexic. Bregn. 262 (reprint 110). 1849. Type. Mexico. Veracruz: “Inter Colipa et Misantla,” Liebmann s.n. [Pl. Mex. 2663, Fl. Mex. 804] (C!; isotype US!).

Rhizomes short-creeping, unbranched, 5–10 mm diam.; fronds 1–4 m long; stipes ca. 1⁄2 the frond length, erect, with 1–2 buds at bases; blades deltate, tripinnate to quadripinnate- pinnatifid; proximal pinnae stalked and alternate, basal pinnules not much reduced; rachises tan to reddish brown; axes pubescent with dimorphic hairs, 1-celled, acicular hairs 0.1–0.3 mm long, and and multicellular, lax to somewhat twisted hairs 0.8–1 mm long; sterile vein tips slender; vertical wings absent adaxially; sori mostly in sinuses, saucer- or purse-shaped, 0.1–0.3 mm deep, 0.5–0.8 mm wide; 2n⫽188 (Jam).

Dennstaedtia cornuta (Kaulf.) Mett., Ann. Sci. Nat. Bot., se´r. 5, 2: 260. 1864. Figs. 105A–J. Dicksonia cornuta Kaulf., Enum. Filic. 227. 1824. Type. Brazil. “Ex Spreng.” [Sello s.n., B may be the collection described (R. Tryon, Contr. Gray Herb. 194: 24. 1964)].

Rhizomes short-creeping, unbranched, 1–4 cm diam.; fronds 1.5–3.5 m long, to 4 cm distant; stipes ca. 1⁄2 the frond length, stramineous, each with 1(–2) buds at base; blades 3–4-pinnate, deltate; proximal pinnae opposite, subsessile, buds frequently in axils of pinnae, basal pinnules reduced; apical segments separate to pinna tips; laminae glabrous to pubescent abaxially, especially on axes, hairs whitish, subappressed, ascending and more or less curled; sterile vein ends clavate; vertical wings absent adaxially; sori distant along segment margins; indusia purse- to cup-shaped, much wider than deep, 1–1.3 mm wide, 0.3 mm deep; 2n⫽92 (Oax), 94 (Chis).

DENNSTAEDTIA DISTENTA

Distribution. Wet montane forests; (220–)900–2450 m. Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Col, Ven, Ec, Peru, Bol. Selected Specimens Examined. Chis (Breedlove 32210, 33443, 33686, NY; Ghiesbreght 368, NY). Gro (Lorea 2347, MEXU). Oax (Hallberg 1751, NY; Mickel 1519, 5353, 6565, 6748, NY). Ver (Copeland herb. 71, UC).

Dennstaedtia cornuta is distinguished by its clavate vein ends and broad indusia. This species has previously been treated as D. dissecta (Sw.) T. Moore, which Navarette and Øllgaard (2000) restricted to the Antilles, Costa Rica, Panama, and the Andes. They distinguished D. cornuta from D. dissecta by the latter having rhizomes creeping or subterranean and roots on the stipe bases (vs. rhizomes ascending and roots absent from the stipe bases in D. cornuta). Unfortunately, rhizomes and stipe bases are rarely present on herbarium specimens, so the characters are of limited practical value.

5. DENNSTAEDTIA CORONATA Dennstaedtia coronata (Sodiro) C. Chr., Index Filic. 216. 1905. Figs. 107F, G. Dicksonia adiantoides Humb. & Bonpl. ex Willd. var. coronata Sodiro, Recens. Crypt. Vasc. Quit. 23. 1883. Dicksonia coronata (Sodiro) Sodiro, Crypt. Vasc. Quit. 48. 1893. Lectotype (chosen by Sodiro, Anales Univ. Quito 7(50): 105 (reprint 49). 1892). Ecuador. [Pichincha:] Corazo´n, Sodiro s.n. (K, AAU photo ex Q, Q, CA photo, US!).

Rhizomes short-creeping, unbranched, ascending, 1–2.5 cm diam.; fronds 1.5–3.5 m, 0.5–1 cm distant; stipes 1⁄3–1⁄2 frond length, stramineous, each with one bud at base; blades arching, rarely scandent, ovate-lanceolate to deltate, bipinnate-pinnatifid, often with axillary buds on rachises and costae; pinnae opposite or subopposite, vertical wings absent adaxially; laminae sparsely pubescent, the hairs 0.2 mm, appressed; sterile veins clavate at tips; sori contiguous on segment margins, indusia cup- or purseshaped, rounded, 1–1.3 mm diam., 0.3–0.5 mm deep.

247

Distribution. Montane rain forests; 300 m. Mexico; Guat, CR, Pan; Gr Ant; Col, Ven, Ec, Peru, Bol. Specimens Examined. Chis (Mu¨nch 60, DS, US).

This species has previously been treated as a synonym of D. arborescens (Willd.) Ekman ex Maxon, which Navarette and Øllgaard (2000) considered distinct and confined to the Antilles.

6. DENNSTAEDTIA DISTENTA Dennstaedtia distenta (Kunze) T. Moore, Index Fil. 306. 1861. Figs. 106A–E. Dicksonia distenta Kunze, Analecta Pteridogr. 39. 1837. Type. Mexico. Veracruz: Jalapa, May 1829, Schiede s.n. (LZ, destroyed!; isotypes B!, NY!).

Rhizomes long-creeping, unbranched, ca. 5 mm diam., bearing brown hairs; fronds 2–4 m long, clumped; stipes brown proximally, stramineous distally, glabrous, each with one bud at the base; blades deltate, 3–4-pinnate-pinnatifid; rachises stramineous; pinnae opposite or subopposite, sessile, with 1 pair of reduced pinnules at pinna bases; sterile vein tips not enlarged, ending well before the margins; abaxial surfaces with hairs 0.8–1 mm long, somewhat tortuous hairs on axes grading to clavate hairs 0.1 mm on laminae; vertical wings absent adaxially; indusia shallow, 0.1– 0.3 mm deep, 0.5–0.8 mm wide; 2n⫽ca. 88 (Chis).

248

DENNSTAEDTIA

Distribution. Wooded banks, often near streams in wet montane forests; 1200–2300 m. Mexico; Guat, Salv, CR, Pan; Jam, Hisp. Selected Specimens Examined. Chih (Nesom & Vorobik 5598, MEXU, TEX). Chis (Breedlove 34619, NY). Dgo (Roberts & Keil 10239, NY). Gro (Hinton 14046, NY). Hgo (Rzedowski 32478, ENCB). Jal (Iltis & Guzma´n M. 3174a, NY). Me´x (Hinton 3748, NY). Mich (Rzedowski 42879, IEB). Mor (Hinton 17065, NY). Oax (Mickel 5011, NY). Pue (Herrera & Riba 3, MEXU). Qro (Rzedowski 46561, IEB). Sin (Gentry 7300, NY). Son (Gentry 1395, UC, US). Ver (Ventura A. 5186, NY).

Dennstaedtia distenta is distinct from D. cicutaria by having sessile, nearly opposite, unreduced, proximal pinnae not reduced at pinna base, stramineous major axes, and monomorphic hairs on axes. “Dennstaedtia mexicana Rosenstock” has never been effectively published but has been used for this species by some authors, e.g., Matuda, Anales Inst. Biol. Univ. Nac. Me´xico 27: 66. 1956. See note by Morton, Amer. Fern J. 48: 124. 1958.

7. DENNSTAEDTIA GLOBULIFERA Dennstaedtia globulifera (Poir.) Hieron., Bot. Jahrb. Syst. 34: 455. 1904. Figs. 105R–U. Polypodium globuliferum Poir. in Lam., Encycl. 5: 554. 1804. Dicksonia globulifera (Poir.) Kuntze, Revis. Gen. Pl. 3: 378. 1898. Type. Plumier, Traite´ Foug. Ame´r., pl. 30, 1705, illustrating a plant from Hispaniola. Dennstaedtia gracilis A. Rojas & Tejero, Revista Biol. Trop. 50(3⁄4): 1008. 2002 [2003]. Type. Mexico. Hidalgo: Mpio. de Molango, km 118 de la Carretera Federal 105, entre Zacualtipa´n y Molango, Rojas et al. 5376 (MEXU; isotypes CR, F, MO, NY!, UAMIZ, UC, US).

Rhizomes long-creeping, 4–10 mm diam., hairy; fronds 1–3 m long; stipes stramineous, lacking buds at bases; blades deltate, tripinnate to subquadripinnate, 1–1.5 m wide; proximal pinnae stalked, basal pinnae opposite, basal pinnules not especially reduced; axes of penultimate segments winged adaxially; sterile vein tips slender, clavate; laminae pubescent on abaxial surfaces, hairs 1 mm long, clear brown, stiff, spreading; indusia globular, (0.5–)1 mm deep, (0.6–)1–1.3 mm wide; 2n⫽ca. 94 (Jam, Parag).

Distribution. Wet montane forests; 600–2100 m. Mexico; Guat, Hond, Nic, CR, Pan; Gr Ant; Col, Ven, Trin, Ec, Peru, Braz, Bol, Parag, Arg, Uru. Specimens Examined. Chis (Breedlove 31351, NY). Gro (Tenorio L. 424, ENCB). Hgo (Clark 7375, NY). Jal (Machuca Nun˜ez 1269, NY). Mich (Torres y Ramı´rez 13771, XAL). Mor (Lyonnet 2801, MEXU; Rose 11058, NY). Oax (Mickel 6145, NY). NL (Rodrı´quez L. 12, NY). Qro (Rzedowski 46579, XAL). SLP (Mickel 598, NY). Tam (Sharp 5077, US). Ver (Purpus 6192, NY).

The large, globular indusia and opposite basal pinnules distinguish this from other species of Dennstaedtia. The small sori of D. gracilis seem to be only a variant of D. globulifera.

8. DENNSTAEDTIA SPINOSA Dennstaedtia spinosa Mickel, Amer. Fern J. 58: 90. 1968. Type. Costa Rica. Osa Peninsula, southwest of Rinco´n de Osa, in rain forest behind Tropical Science Center, Mickel 2730 (US!; isotypes ISC!, LP!, NY!, USJ!). Figs. 107A–C. Rhizomes creeping, unbranched, clothed with black hairs; fronds nearly 2 m long; axes stramineous, spiny and pubescent, the spines to 3 mm long, straight or slightly antrorse; stipes ca. 1 ⁄2 the frond length, buds at bases not seen, but expected; blades ca. approx. 1 ⫻ 1 m, deltate, quadripinnate-pinnnatifid, lacy; pinnae alternate, short-petiolulate to 12 mm; basal pinnules opposite, not or only slightly reduced, the larger 10–13 ⫻ 2.5–3 cm; laminae hirsute abaxially with straight, hyaline hairs, lustrous adaxially and nearly glabrous except on costal wings and major veins; laminar grooves not continuous, with vertical wings adaxially; vein ends slender, clavate; indusia ca. 0.6 mm deep, 0.5– 0.8 mm wide, entire to irregularly dentate, low, broader than high, more or less bivalvate.

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valve fitting into the outer; sporangia with oblique annuli; spores tetrahedral; x⫽65. Dicksonia is a genus of about 20 species, mostly of the temperate Australasian areas, with three species thought to occur in tropical America, generally at high elevations. There is one species in Mexico. Dicksonia is allied to Cibotium, differing from that genus by its erect trunk and nonglaucous abaxial laminae. Another ally is Culcita, differing from Dicksonia by the distinct trunks and the blades 3-pinnate or less divided (vs. trunkless and the blades 4-pinnate). Reference Maxon, W. R. 1913. Studies of tropical American ferns. No. 4. Contr. U.S. Natl. Herb. 17: 133–179.

Distribution. Terrestrial in lower montane rainforests; 450 m. Mexico; Guat, Hond, Nic CR, Pan. Specimens Examined. Chis (Breedlove 57862, CAS, ENCB).

This species closely resembles D. cicutaria s.l., but can be distinguished readily by the spiny axes and finer blade dissection. Both species have dicyclic solenostelic rhizomes, but the steles of D. spinosa are terete and uncorrugated. The spores of D. cicutaria are verrucate, whereas those of D. spinosa have low, irregularly anastomosing ridges. Furthermore, the indusia of D. spinosa are more regularly dentate than those of D. cicutaria. Although D. cicutaria occasionally has a few prickles on its axes, and a few specimens described as D. decomposita Christ are as dissected as D. spinosa, no specimens have the regular and consistent spininess or the anatomical and spore characteristics of D. spinosa. Excluded Species Dennstaedtia obtusifolia (Humb. & Bonpl. ex Willd.) T. Moore, Index Fil. 306. 1861. Dicksonia obtusifolia Humb. & Bonpl. ex Willd., Sp. Pl., ed. 4, 5(1): 483. 1810. Type. Venezuela. Caracas, Bredemeyer s.n. (B-Willd. 20163!; frag. LE!). Specimens previously considered as this species are here treated under D. cornuta, the true D. obtusifolia being limited to Venezuela, Colombia, Ecuador, and Peru and having rounded (vs. elongate) sori.

1. DICKSONIA SELLOWIANA Dicksonia sellowiana Hook., Sp. Fil. 1: 67. 1844. Lectotype (chosen by R. Tryon & Stolze, 1989: 105). Brazil, Sellow s.n. [no. 9] (K!, frag. NY!; isolectotype HBG, photos GH, US). Figs. 108A–C. Dicksonia gigantea H. Karst., Fl. Columb. 5: 177, t. 193. 1869. Type. Colombia. Andes of Bogota, “montis Guadeloupe,” Karsten s.n. (B!). Dicksonia ghiesbreghtii Maxon, Contr. U.S. Natl. Herb. 17: 155. 1913. Type. Mexico. Chiapas: Ghiesbreght 353 (US!; P!, frag. US; isotypes BM!, K!).

Stems to 5 m tall, 12–15 cm diam., apices clothed with dark golden-brown hairs; fronds 2–3 m long; stipes short, dark at bases, brown to stramineous distally, spines absent, bearing golden brown hairs to 4–5 cm long; blades tripinnate to tripinnatepinnatifid, coriaceous, primary pinnae to 70 ⫻ 20 cm, smaller toward blade bases, essentially sessile, stalks less than 1 cm; hairs scattered on axes abaxially, 2–3 mm long, short hairs on adaxial costules; sori marginal, bivalvate, 1 mm long, paraphyses numerous, 1 mm long.

3 7. DI C K S O N I A Dicksonia L’He´r., Sert. Angl. 30. 1788 [1789]. Lectotype (chosen by Presl, Suppl. Tent. Pterid. 135. 1836.): Dicksonia arborescens L’He´r. Balantium Kaulf., Enum. Filic. 228. 1824. Lectotype (chosen by Maxon, J. Wash. Acad. Sci. 12: 455. 1922): Balantium auricomum Kaulf. [⫽ Dicksonia arborescens L’He´r.].

Terrestrial; stems forming erect trunks, apices densely clothed with stiff golden brown to brown hairs; fronds large, not articulate, monomorphic (fertile laminae slightly contracted); stipes hairy; blades bipinnate-pinnatifid to tripinnate, coriaceous, with sparse hairs; veins free; sori marginal, indusia bivalvate, the inner

Distribution. Montane rain forests, cloud forests; 1450–2450 m. Mexico; Guat, Hond, Salv, Nic, CR, Pan; Col, Ven, Ec, Peru, Braz, Bol, Uru. Selected Specimens Examined. Chis (Breedlove 32654, DS, MEXU, NY; Purpus 6716, BM, MO). Gro (Lorea 2390, ENCB, FCME, IEB; Rzedowski & McVaugh 21, CAS, ENCB, IEB, MEXU, MICH, NY).

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Hgo (Gimate 977, ENCB, MEXU). Oax (Gonza´lez & Conzatti 724, US; Mickel 5348, ENCB, MEXU, NY, US). Pue (Riba et al. 297, MEXU, NY, US; Ventura A. 8091, ENCB, MEXU, NY). Ver (Ventura A. 287, XAL).

Several names have been applied to this species in different parts of its range, but there seems to be only one widespread species of Dicksonia in America and regional variation does not seem to be taxonomically useful. Mexican material has previously been referred to D. gigantea.

3 8 . D I C R A N O PT E R I S Dicranopteris Bernh., Neues J. Bot. 1(2): 38. 1806 [1805], nom. nov. for Mertensia Willd., Kongl. Vetensk. Acad. Nya Handl. 25: 165. 1804, hom. illeg., non Roth, 1797. Lectotype (chosen by Bernhardi, Neues J. Bot. 1(2): 38. 1806 [1805]): Mertensia dichotoma (Thunb. ex Murray) Willd. [⬅ Polypodium dichotomum Thunb. ex Murray] ⬅ Dicranopteris dichotoma (Thunb. ex Murray) Bernh. Hicriopteris C. Presl, Epimel. Bot. 26. 1849 [1851]. Type: Hicriopteris speciosa C. Presl [⬅ Dicranopteris speciosa (C. Presl) Holttum]. For further synonymy, see Pichi Sermolli (Webbia 26: 491–536. 1972).

Terrestrial; rhizomes long-creeping, cord-like, with stiff, dark, bristle-like hairs; fronds monomorphic, erect or scandent, to several meters long; blades pseudodichotomous, with pinnae consistently and repeatedly equally branching, i.e., both branches of a fork approximately the same size, bearing accessory branches at the base of some of the forks (especially proximally), axes terete; dormant buds covered by stiff, reddish brown, multicellular hairs, these often basally branched, also with a pair of foliaceous stipule-like outgrowths in some buds, especially in the proximal forks; penultimate divisions pectinate, bearing narrowly lanceate to linear segments; indument absent or of branched hairs on the blade and axes; veinlets free, 2–3(–4)-forked; sori abaxial, round, of ca. 6–15 sporangia; paraphyses absent; sporangia each with oblique annulus; indusia absent; spores whitish, tetrahedral, surfaces smooth, pitted, or granulate; x⫽39. Dicranopteris is a pantropical genus of about 10 species, four in tropical America. The single species in Mexico is widespread and common in the Neotropics. The genus is allied to Gleichenella and Sticherus, and is a member of an ancient and isolated lineage of basal leptosporangiate ferns. It occurs at low to middle elevations (some South American species at relatively high elevations), on roadsides and in clearings, in weedy, disturbed areas, sometimes forming thickets. From Gleichenella, it is distinct by the equally forking pseudodichotomies, presence of accessory pinnae subtending all but the penultimate divisions, terete axes lacking lateral ridges, and by the tetrahedral spores. Dicranopteris is distinct from Sticherus by the veins forking 2–3 times and the sori of 6–15 or more sporangia. See Gleichenella for a discussion of family level relationships.

References Ching, R. C. 1940. On the genus Gleichenia. Sunyatsenia 5: 269–288; Holttum, R. E. 1957. Morphology, growth-habit, and classification in the family Gleicheniaceae. Phytomorphology 7: 168–184; Maxon, W. R. 1909. Gleicheniaceae. N. Amer. Fl. 16: 53–63; Nakai, T. 1950. New classification of Gleicheniales, etc. Bull. Natl. Sci. Mus., Tokyo 29: 1–71; Østergaard Andersen, E. & B. Øllgaard. 1996. A note on some morphological terms of the leaf in the Gleicheniaceae. Amer. Fern J. 86: 52–57; Østergaard Andersen, E. & B. Øllgaard. 2001. 10. Gleicheniaceae. In: G. Harling & L. Andersson (eds.), Flora of Ecuador 66: 105– 170.

1. DICRANOPTERIS FLEXUOSA Dicranopteris flexuosa (Schrad.) Underw. Bull. Torrey Bot. Club 34: 254. 1907. Figs. 109A–D. Mertensia flexuosa Schrad., Go¨tt. Gel. Anz. 1824: 863. 1824. Gleichenia flexuosa (Schrad.) Mett., Ann. Mus. Bot. Lugduno-Batavum 1: 50. 1863. Type. Brazil. Espirito Santo: Wied-Neuwied s.n. (BR!, photo BM!).

Rhizomes 2–4 mm diam., with stiff, red-brown hairs that abrade to leave a scabrous surface; stipes very brittle, sharpsplintering, tan to brown, glabrous, dull; frond apices dormant or often continuing, pinnae 0–2 times forked; ultimate branches mostly 10–30 ⫻ 2–5 cm, basally often wider due to enlarged, sometimes pinnatifid basiscopic basal segments; ultimate segments linear, mostly 10–25 ⫻ 2–5 mm, dilated at base, subcoriaceous, often emarginate at the tip, usually glabrous or glabrescent, green to glaucous abaxially, margins often somewhat revolute, especially in fertile segments, veins immersed; sori medial between midvein and margin; 2n⫽ca. 156 (Jam).

Distribution. On dry slopes in open areas of lower Atlantic slopes, sandstones bluffs with oaks; 100–1400 m. USA (Ala, Fla); Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol, Parag. Selected Specimens Examined. Chis (Breedlove 21967, DS, NY, 45960, CAS). Gro (Kruse 940, MEXU; Lorea 4802, FCME, IEB). Oax (King 831, ENCB, MICH, US; Mickel 7379, ENCB, MEXU, NY, UC, US). Tab (Hill & Cowan 1533, CAS, CSAT). Ver (Orcutt 3205, BM, DS, K, MEXU, MO, NY, US; Palacios-Rios 3138, NY, XAL; Smith 2107, MEXU, NY, UC, US).

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This species is exceedingly close to D. linearis (Burm. f.) Underw. from the Paleotropics, and the two species may prove to be conspecific. In Mexico, Dicranopteris flexuosa is most similar to Gleichenella pectinata, which see, but much less common.

3 9. D I DYM O C H L A E N A Didymochlaena Desv., Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck. Gesammten Naturk. 5: 303. 1811. Type: Didymochlaena sinuosa Desv. [⫽ Didymochlaena truncatula (Sw.) J. Sm.]. Terrestrial; rhizomes massive, suberect to erect, scaly; fronds clumped, monomorphic, bipinnate; pinnules articulate, oblong, subdimidiate; veins free, forked, not reaching margin; hydathodes present adaxially; costae channelled, lateral ridges with numerous awns 0.5–0.8 mm long, these clustered at base of pinnules; sori terminal on veins, elongate; indusia elliptical, rounded at distal end, cordate or sagittate at base, fixed to vein along medial line; spores bilateral, perispore saccate or folded; x⫽41. Didymochlaena is a monotypic genus, with the single species pantropical and occurring at relatively low elevations. Precise relationships of the genus are unclear, but are probably dryopteroid/polystichoid. Unpublished molecular work by Cranfill (pers. comm.), using especially the rps4 gene, supports a hypothesis that Didymochlaena may lie near the base of the dryopteroid lineage, but also that it is a rather isolated element within the Dryopteridaceae s.l. Didymochlaena is distinguished by its oblong, entire, subdimidiate segments (pinnules) and elliptical sori, each sorus on both sides of a vein and continuous over the vein at the acroscopic end.

1. DIDYMOCHLAENA TRUNCATULA Didymochlaena truncatula (Sw.) J. Sm., J. Bot. (Hooker) 4: 196. 1841. Figs. 110A–E. Adiantum lunulatum Houtt., Nat. Hist. 2(14): 209, t. 100, f. 1. 1783, hom. illeg., non Burm.f., 1768. Aspidium truncatulum Sw., J. Bot. (Schrader) 1800(2): 36. 1801. Type. Java. s.coll. s.n. (S).

Rhizome scales tan to brownish, concolorous, 10–20 ⫻ 1–2 mm, linear-lanceolate; stipes stramineous, 40–45 cm long, 1⁄3 the frond length, with scattered scales like those of rhizome, especially at base, and a dense undercovering of matted, tortuous, woolly hairs; blades 70–90 ⫻ 30–45 cm, rachises similar to stipes; proximal pinnae somewhat shortened; pinnae numerous, 19–30 ⫻ 3–4 cm; costae with scales and lax, matted hairs and with clusters of 2–10 awns (0.6–0.8 mm long) at pinnule attachments; pinnules oblong, 1–2.5 ⫻ 0.5–1 cm, entire rounded or crenate at apex, basal ones of each pinna somewhat smaller; blades dark green, glabrous on both surfaces, chartaceous; sori 3–5 per pinnule, along acroscopic side or 1–2 at tip of pinnules; indusia tan, 1–2.5 ⫻ 1–1.5 mm, entire; 2n⫽82 (CR).

Distribution. Terrestrial in wet lowland and montane forests; 200–1700 m. Mexico; Guat, Bel, Hond, Nic, CR, Pan; Cuba, Hisp; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Bol, Braz, Parag, n Arg; Africa, Asia. Selected Specimens Examined. Chis (Breedlove 48436, 50459, CAS; Dressler 1607, MEXU, US; Ton 6742, MEXU, UC). Oax (Hallberg 1746, NY, UC; Herna´ndez G. 1969, CHAPA, NY; Wendt et al. 3722, CHAPA, NY, UC). Pue (Campos V. 411, IEB). Ver (Fink 95, US; Ventura 8119, ENCB, NY).

Didymochlaena truncatula is easily distinguished by the unique horseshoe-shaped sori and indusia, the large bipinnate blades with entire articulate pinnules, and the awns borne along the ridges of the costae adaxially.

40. DIPLAZIUM Diplazium Sw., J. Bot. (Schrader) 1800(2): 61. 1801. Lectotype (chosen by J. Smith, Hist. Fil. 325. 1875): Diplazium plantaginifolium (L.) Urb. [⬅ Asplenium plantagineum L.]. Terrestrial; roots stout and somewhat fleshy; rhizomes usually suberect to erect (caudices sometimes trunk-like), scaly; fronds medium-sized to often large, monomorphic, not articulate; stipes usually glabrous, scaly at bases, with two crescent-shaped vascular bundles in cross section; blades thin-herbaceous to chartaceous or subcoriaceous, usually pinnate to 2–3-pinnatepinnatifid, sometimes 4⫹-pinnate, rarely simple, some species producing buds in axils of blade bases, pinnae, or pinnules, these sometimes developing into plantlets; veins generally free, rarely casually to abundantly anastomosing, pinnate in the ultimate segments or sometimes strongly curved toward blade margins and appearing parallel; indument abaxially of brownish to tan scales on rachises, costae, and costules, sometimes also of hairs, these often septate, adaxially the blades glabrous or with short papillae or hairs ca. 0.1 mm (or less) in rachial, costal, and costular grooves; sori abaxial, linear or oblong on the veins, usually with at least some of them (sometimes only the basal ones) backto-back (diplazioid) on a single vein; indusia generally covering each sorus, rarely absent, paraphyses absent; sporangia shortstalked, stalks less than half the length of capsules; spores bilateral, with winged perispores; x⫽41.

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Diplazium comprises 350–400 species, with probably more than 150 in the New World; it is generally a genus of wet tropical forests of low to middle (rarely high) elevations. Twenty-five species are known from Mexico. Chromosome counts for species range from diploid, 2n⫽82, to octaploid, 2n⫽328. Judging by the number of intermediate specimens that show malformed spores and a few chromosome counts, hybridization is apparently common in the genus, at least in the Neotropics. Putative hybrids have been suggested between species in Ecuador (Stolze et al., 1994) and Mexico (Mickel & Beitel, 1988; Smith, 1981). Monographic studies are needed for a better understanding of the species and their relationships. Diplazium is distinguished by its commonly indusiate sori (among Mexican species, indusia are absent, or nearly so, only in D. lindbergii) that are elongate and at least sometimes back-to-back along the veins, relatively thick roots, suberect to erect rhizomes, two ribbon-like vascular

bundles in the petioles, and a chromosome base number of x⫽41. At least half of the Mexican species have distinctive papillae 0.05–0.1 mm long in the rachial, costal, and costular grooves adaxially. Relationships of the genus are with the athyrioid ferns, especially Athyrium and a number of Old World genera usually now segregated from Diplazium and Athyrium, e.g., Callipteris, Cornopteris, and Deparia. Diplazium has more distant relationships with Onocleopsis and the onocleoid ferns, and, beyond that, with the blechnoid ferns (Cranfill, unpubl.). References Adams, C. D. 1992. Eleven new species, a new variety, and a new varietal combination in the fern genera Asplenium and Diplazium in Central America. Novon 2: 290–298; Lellinger, D. B. 1978. Rı´o Palenque fern notes. Selbyana 2: 283–285; Stolze, R. G., L. Pacheco & B. Øllgaard. 1994. Polypodiaceae–Dryopteridoideae– Physematieae. In: G. Harling & L. Andersson (eds.), Flora of Ecuador 49: 1–108.

Key to the Mexican Species of Diplazium 1. Blades simple. ...................................................................................................................................................................................... 17. D. plantaginifolium. 1. Blades compound, 1-pinnate or more divided. 2. Blades pinnate to pinnate-pinnatifid, if 2-pinnate, then only proximal pinnae with separate basal pinnules. 3. Blades ternate, having 3 pinnae (two lateral plus a terminal). ................................................................................................. 21. D. ternatum. 3. Blades pinnate with more than 3 pinnae. 4. Blades pinnatifid in the distal half to one-third of their length, 6–12 cm wide. ........................................................... 23. D. vera-pax. 4. Blades pinnate almost to apices, more than 15 cm wide. 5. Blade apices with a Ⳳ conform terminal pinna similar to the lateral ones; bud(s) borne at pinna bases; pinnae entire, or nearly so. ....................................................................................................................................................... 16. D. obscurum. 5. Blade apices lacking a distinct terminal pinna similar to the lateral ones, the apices pinnatifid, confluent; bud(s) present or absent at pinna bases; pinnae entire to crenate to shallowly or deeply lobed. 6. Pinna bases of median pinnae equilateral, truncate; rachial and costal grooves with dense erect papillae to 0.1 mm long. 7. Pinnae entire to lobed 1⁄3 or less of the distance to costae. 8. Pinnae Ⳳ entire in basal half; ultimate veins running in Ⳳ parallel fashion to margins; sori commonly borne on 1–2 veins per vein group; buds absent. ................................................. 10. D. grandifolium. 8. Pinnae shallowly lobed to one-third the distance to costae in basal half; veins pinnate, with 3–4 pairs of veinlets running to pinna margins; sori commonly borne on 2–6 veins per vein group; bud(s) sometimes present at pinna bases. 9. Largest pinnae more than 4 cm wide. ..................................................................................... 15. D. neglectum. 9. Largest pinnae to 3.5 cm wide. ............................................................................................. 22. D. urticifolium. 7. Pinnae lobed 2⁄3 or more of the distance to costae. 10. Laminar tissue between veins glabrous abaxially; costae and rachises abaxially with scattered, brownish, lax, septate hairs or linear-lanceolate scales 0.5–5 mm long on costae and costules; indusia absent or caducous, ⬍0.2 mm wide. .................................................................................. 13. D. lindbergii. 10. Laminar tissue between veins, costae, and rachises abaxially with scattered, erect, hyaline hairs 0.1–0.3 mm long; indusia conspicuous and persistent, 0.3–0.6 mm wide. 11. Proximal pinnae pinnatifid, basal segments adnate and confluent; pinnae 7–14(–16) ⫻ 1.8–3 cm; rhizome scale cell walls straight. ................................................................................... 19. D. striatastrum. 11. Proximal pinnae fully pinnate at their bases, with 1–2 free pinnules; pinnae 7–30 ⫻ (3–)3.5– 9 cm; rhizome scale cell walls sinuous. .................................................................................... 20. D. striatum. 6. Pinna bases of median pinnae inequilateral, basiscopically excavate; rachial and costal grooves with sparse papillae ⬍0.1 mm long. 12. Proximal pinnae with free or nearly free basal acroscopic pinnules. 13. Pinnae 18–20(–25) ⫻ (3.5–)4.5–10 cm, segments strongly oblique and falcate; indusia 0.3–0.5 mm wide, subentire. .................................................................................................................... 5. D. drepanolobium. 13. Pinnae (2–)8–16 ⫻ 2–4(–6) cm, segments slightly oblique and straight; indusia 0.5–1 mm wide, erose. .............................................................................................................................................. 14. D. lonchophyllum. 12. Proximal pinnae undulate to deeply lobed, without free basal acroscopic pinnules. 14. Pinnae deeply lobed (1⁄2 to 3⁄4 of distance to costae); indusia subentire, 0.8–1 mm wide. .........2. D. cristatum. 14. Pinnae undulate to shallowly lobed (1⁄4 of distance to costae); indusia erose, 0.3–0.5 mm wide.

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15. Pinnae 20–28 pairs per blade; blades each bearing a bud or plantlet in the axil of a distal pinna. .................................................................................................................................................. 6. D. errans. 15. Pinnae usually 5–10 pairs per blade; blades lacking buds or plantlets in axils of distal pinnae. .................................................................................................................................... 25. D. werckleanum. 2. Blades 2–4-pinnate. 16. Pinnae and/or pinnules inequilateral, basiscopically excavate and acroscopically slightly auriculate. 17. Rachial and costal grooves puberulent adaxially with minute hairs 0.1 mm long; rachises, costae, and costules abaxially with short, erect hairs. .................................................................................................................................... 18. D. puberulentum. 17. Rachial and costal grooves glabrous adaxially, or nearly so; costae and rachises with scattered hair-like scales 0.5–2 mm long, or glabrous. 18. Blades 2-pinnate-pinnatifid, pinnules lobed (1⁄3–)1⁄2–3⁄4 of distance to costules; indusia 0.5–0.8 mm wide, entire to erose; stipes and rachises glabrous. ........................................................................................................................... 8. D. franconis. 18. Blades 3-pinnate, at least proximally; indusia 0.1–0.5 mm wide, entire to subentire. 19. Blades 3-pinnate to 3-pinnate-pinnatifid proximally; costae, costules, and more minor axes not prominently winged adaxially, or the wings stramineous to tan; indusia 2–4 mm long; stipes and rachises with numerous brownish hair-like, lax scales 0.8–2 mm long. ............................................................... 4. D. donnell-smithii. 19. Blades 4-pinnate proximally; costae, costules, and more minor axes winged and deeply grooved adaxially, wings green and perpendicular to plane of blades; indusia 1–2.5 mm long; stipes and rachises scaleless. ................................................................................................................................................................................... 24. D. wendtii.

16. Pinnae and pinnules equilateral, truncate. 20. Blades 3-pinnate or more divided. 21. Blades 3-pinnate to 3-pinnate-pinnatifid; pinnules opposite to subopposite, nearly equilateral at their bases. ...................................................................................................................................................................................... 9. D. gomezianum.

21. Blades 3-pinnate-pinnatifid to 4-pinnate; pinnules alternate, strongly inequilateral at their bases. .................... 24. D. wendtii. 20. Blades 2-pinnate to 2-pinnate-pinnatifid. 22. Rhizome scales solid, horizontal walls intact, dark brown with denticulate margins; rachial and costal grooves glabrous. 23. Uncut portions of indusia (not including the cilia) 0.5–0.8 mm wide; buds present at the bases of some pinnae and/or pinnules. ....................................................................................................................................... 1. D. altissimum. 23. Uncut portions of indusia (not including the cilia) 0.1–0.3 mm wide; buds absent. .............................. 3. D. diplazioides. 22. Rhizome scales retiform, surface walls gone, leaving only lateral wall latticework, margins frayed; rachial and costal grooves densely and minutely puberulent. 24. Largest pinnules incised at bases 1⁄3 or less their width with 1 or 2 pairs of lobes, elsewhere crenate to serrulate. .................................................................................................................................................................. 20. D. striatum. 24. Largest pinnules incised at bases 1⁄2 or more toward costules with 5 or more pairs of lobes. 25. Laminar tissue between veins abaxially with numerous hairs; scales absent on costae or costules (rare scales at pinnule bases); indusia erose, 0.3–0.5 mm wide. ...................................................................... 7. D. expansum. 25. Laminar tissue between veins abaxially lacking hairs; narrow scales on costules, often also on costae and rachises; indusia erose or laciniate-ciliate, 0.1–0.5 mm wide. 26. Indusia laciniate-ciliate, 0.1–0.3 mm wide (not including cilia); rachises and costae with numerous, narrow scales. ..................................................................................................................... 11. D. hellwigii. 26. Indusia erose, 0.3–0.5 mm wide; rachises and costae glabrous or with relatively sparse scales, scales along costules. ................................................................................................................................. 12. D. hians.

1. DIPLAZIUM ALTISSIMUM Diplazium altissimum (Jenman) C. Chr., Index Filic. 227. 1905. Figs. 112K, L. Asplenium altissimum Jenman, J. Bot. 17: 259. 1879. Type. Jamaica. Jenman 64 (K; isotype US!). Diplazium entecnum Mickel & Beitel, Mem. New York Bot. Gard. 46: 154. 1988. Type. Mexico. Oaxaca: Distrito Ixtla´n, 65 km N of Ixtla´n, 44 km S of Valle Nacional, Mickel 5581 (NY!; isotype UC!).

Differs from D. diplazioides by the blades bearing buds at bases of some pinnules (along costae), and sori with laciniate-ciliate indusia 1–2 mm long, uncut portion 0.5–0.8 mm wide. Distribution. Terrestrial in wet forests; 1150–2000 m. Mexico; Cuba, Jam, Hisp.

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Specimens Examined. Chis (Breedlove 15402, DS, US). Oax (Mickel 4281, 4302, 5567, 5641, 5712, 7100, NY, UC, 1551, US; Rzedowski 33822, ENCB, NY, 34082, IEB, 34044, 34083, ENCB, IEB).

See discussion under D. diplazioides, the closest relative. Many Jamaican specimens of D. altissimum also have buds at bases of the pinnules, e.g., Maxon 9992 (US). The sole collection from Chiapas was cited by Adams (in Davidse et al., 1995) as D. diplazioides, but this specimen bears buds at the bases of the pinnules, and so we refer it to D. altissimum.

2. DIPLAZIUM CRISTATUM Diplazium cristatum (Desr.) Alston, J. Bot. 74: 173. 1936. Figs. 112M, N. Meniscium cristatum Desr. in Lamarck, Encycl. 4: 94. 1797. Type. Martinique. Martin s.n. (P-Lam.). Asplenium arboreum Willd., Sp. Pl., ed. 4, 5(1): 320. 1810. Diplazium arboreum (Willd.) C. Presl, Suppl. Tent. Pterid. 114. 1836. Type. Venezuela. Caracas, Bredemeyer s.n. (B-Willd. 19892-1!). Asplenium denticulosum Desv., Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck. Gesammten Naturk. 5: 323. 1811. Diplazium denticulosum (Desv.) C. Chr., Index Filic. 231. 1905. Type. “In America calidiore,” Dombey s.n. (P-JU!; isotype P!). Asplenium shepherdii Spreng., Nova Acta Phys.-Med. Acad. Caes. Leop.Carol. Nat. Cur. 10: 231, t. 17, f. 5, 6. 1821. Type. Jamaica. Shepherd s.n. (LZ, destroyed; isotype PH).

Similar to D. werckleanum except: rhizome scales 5–7 ⫻ 1–1.5 mm; stipes 30–55 cm long; blades pinnate-pinnatifid, 30–45(–100) cm long; pinnae (5–)12–13 ⫻ (1.5–)2.5–3.5(–4.5) cm, deeply lobed 1⁄2–3⁄4 of the distance to costae, lobes serrate, basal acroscopic lobe not free; sori with subentire indusia 4–9 ⫻ 0.8–1 mm; 2n⫽ca. 164 (Jam, Trin).

Distribution. Terrestrial in montane forests of the Isthmus and along the Pacific slope; 1300–2700 m. Mexico; Guat, Bel, Hond, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, s Braz, Bol, Parag, n Arg. Also cited from El Salvador by Adams (in Davidse et al., 1995: 233), but the specimen (Standley 21742, MO!) appears to us to be D. lonchophyllum. Selected Specimens Examined. Chis (Breedlove 7374, 9295, 22652, 29210, DS). Gro (Correll 14406, US; Velazquez 799, FCME). Oax

(Campos V. 1900, MEXU, UC; Mickel 5100, 6046a, NY; Torres C. 11894, UC).

Citation of this species from Nayarit by Mickel and Beitel (1988) is the result of misidentification of a specimen of D. lonchophyllum. Several other specimens cited as D. cristatum from Oaxaca (Mickel & Beitel, 1988), from the Sierra de Miahuatla´n, are probably also D. lonchophyllum (Mickel 5026, NY, UC, 6010, NY, juvenile, 6019, NY), but both species apparently occur in that area. Two of the four specimens cited by Palacios Rios (1992) as D. cristatum are D. lonchophyllum: Conzatti 802, MEXU; Palacios Rios & Herna´ndez M. 610, XAL; we have not seen the other two (Castillo & Medina 4288, XAL, and Nee 22582, F, XAL), but their identity needs verification). See discussion under D. werckleanum for a discussion of differences and hybrids with that species.

3. DIPLAZIUM DIPLAZIOIDES Diplazium diplazioides (Klotzsch & H. Karst. ex Klotzsch) Alston, J. Bot. 74: 174. 1936. Fig. 114E. Lotzea diplazioides Klotzsch & H. Karst. ex Klotzsch, Linnaea 20: 358. 1847. Asplenium klotzschii Mett., Fil. Lips. 79. 1856, non Asplenium diplazioides Hook. & Arn., 1832. Diplazium klotzschii (Mett.) T. Moore, Index Fil. lv. 1857, hom. illeg. Type. Colombia. Karsten 23(II) (B). Diplazium laciniatum Mickel & Beitel, Mem. New York Bot. Gard. 46: 156. 1988. Type. Mexico. Oaxaca: Distrito Ixtla´n, 79 km N of Ixtla´n de Jua´rez on Rte 175, 1 km S of Campamento Vista Hermosa, ca. 1 hour along trail towards Tarabundı´, Mickel 5713 (NY!; isotype UC!).

Rhizomes erect; rhizome scales concolorous, dark brown, 5–10 ⫻ 1.5–2.5 mm, not retiform, margins denticulate; fronds 100–200

cm long; stipes stramineous, darkened at bases, 45–90 cm ⫻ 5– 10 mm, 1⁄3–1⁄2 the frond length, with abundant denticulate scales at bases, otherwise glabrous; blades herbaceous to chartaceous, 2-pinnate-pinnatifid to nearly 3-pinnate, to 50 ⫻ 45 cm, deltate to ovate, gradually tapering to pinnatifid apices, lacking buds; rachises with scattered tan lanceolate scales on both sides, lacking papillae in the adaxial grooves; pinnae 10–12 pairs, stalked to 25 mm, 15–32 ⫻ 7–18 cm, equilateral or nearly so, pinnules sessile to adnate distally, or largest ones on proximal pinnae stalked to 6 mm, equilateral, cut 3⁄4–9⁄10 the distance to costules, costae and costules adaxially with stramineous to tan, interrupted wings perpendicular to the plane of the blades; veins free, pinnate, 4– 7 pairs per segment, often 1-forked; indument adaxially absent in rachial and costal grooves (minute papillae absent), abaxially of sparse to numerous brownish to tan hair-like to lanceolate denticulate scales 1–3 mm long on costae, costules, and veins, tissue between veins glabrous; sori pinnately arranged in ultimate segments, with indusia 1–3 mm long, deeply laciniate-ciliate, uncut portions 0.1–0.3 mm wide, cilia 0.5–0.8 mm long.

DIPLAZIUM DREPANOLOBIUM

255

adnate-decurrent toward pinna tips, acute at apices, the larger pinnules with 1–4 free pinnulets at the bases; indument absent except for minute papillae 0.05 mm long in costal and costular grooves adaxially and scattered to rare, dark reddish brown, hairlike scales 0.5–2 mm long on costae and costules abaxially; indusia 2–4 ⫻ 0.4–0.5 mm, entire to subentire.

Distribution. Terrestrial in wet forests; 800–1900 m. Mexico; Guat, Nic, CR; Col, Ven, Trin (Stolze et al., 1994), Ec, Bol. Also cited from Honduras by Adams (in Davidse et al., 1995). Selected Specimens Examined. Oax (Croat 47938, MO; Mickel 1453, US, 5674, 5675, 5688, 6541, 6605, 6692, UC; Rzedowski 32988, ENCB).

Diplazium laciniatum, recently described from Oaxaca, is identical to specimens of D. diplazioides from Central America and Venezuela. Another possible synonym is D. croatianum C. D. Adams, type from Panama. We regard Breedlove 15402, cited as D. diplazioides by Adams (in Davidse et al., 1995), as D. altissimum. Diplazium diplazioides and D. altissimum are distinguishable from the other 2-pinnate-pinnatifid taxa in Mexico by the combination of large, concolorous, dark brown, non-retiform stipe base scales with denticulate margins, the lack of hairs or papillae in the rachial, costal, and costular grooves adaxially, and pinnae that range from equilateral to having the basiscopic basal pinnule without matching acroscopic pinnule. Diplazium diplazioides differs from D. altissimum in having wider, erose indusia (0.8–1 mm) and scales with denticulate margins on the rachises, costae, and costules; similar scales are also found in the rachial and costal grooves adaxially. Another closely related species with toothed rhizome scales and deeply laciniate-ciliate indusia is D. l’herminieri Fe´e of Guadeloupe. The characters distinguishing D. diplazioides from D. altissimum as given in the key are minor, but consistent; however, biosystematic study is necessary to clarify the relationships of these taxa vis-a`-vis other 2-pinnatepinnatifid species.

4. DIPLAZIUM DONNELL-SMITHII Diplazium donnell-smithii Christ, Bull. Herb. Boissier, se´r. 2, 7: 270. 1907. Type. Honduras. San Pedro Sula, Thieme s.n. [Donn. Sm. 5680] (P?; isotype US!, frag. NY!). Figs. 113C, D. Differs from D. franconis by: stipes greenish stramineous, with sparse to scattered hair-like, lax, twisted scales 0.8–2 mm long; blades herbaceous to chartaceous, 3-pinnate, at least proximally, deltate; pinnae with pinnules stalked 1–4 mm, 10–15 free pairs per pinna, the largest 5–10 ⫻ 1.5–2.5 cm, inequilaterally deltate,

Distribution. Terrestrial in wet montane forests; (150–)1300– 1650 m (in Guat). Mexico; Guat, Hond, Salv, Nic. Also cited from Costa Rica by Adams (in Davidse et al., 1995), citing Go´mez. Specimens Examined. Oax (Hellwig 457, NY, US). Ver (Barrington 445, MEXU; Conant 737, MEXU; Fink 32, US; Herna´ndez M. 1381a, NY; Palacios-Rios 3323, XAL; Ventura A. 3271, ENCB, MEXU).

Other than having more divided blades (truly 3-pinnate, at least proximally), the Oaxacan specimen of D. donnell-smithii is distinguishable from D. franconis by the numerous hair-like scales on the rachises, costae, and costules abaxially. The isotype at US (Donnell-Smtih 5680) differs from the Oaxacan specimen in having rare, hair-like, clathrate scales. Diplazium hahnii (E. Fourn.) C. Chr. (Asplenium hahnii E. Fourn., Mexic. Pl. 1: 109. 1872; type from Mexico, Veracruz, Mt. Escamela, Hahn s.n., P!, photos NY!, UC, US) may be conspecific with D. donnell-smithii, but the holotype is somewhat teratological. If D. hahnii is conspecific with D. donnell-smithii, the former has priority. See under D. franconis for further comparison and discussion of putative hybrids with that species. Adams (in Davidse et al., 1995) cited Ghiesbreght 361 (BM, not seen) as this species from Guatemala, but we are unaware that Ghiesbreght collected in Guatemala; most likely the specimen is from Chiapas. Ghiesbreght 361 (K) is from Chiapas, and is D. drepanolobium (Smith, 1981), a species quite different from D. donnell-smithii.

5. DIPLAZIUM DREPANOLOBIUM Diplazium drepanolobium A. R. Sm., Amer. Fern J. 70: 21. 1980. Type. Mexico. Chiapas: 10 km above Rayon, Breedlove 26122 (DS!; isotype MO!). Fig. 112J.

256

DIPLAZIUM

Differs from D. lonchophyllum by: rhizome scales 5–7 ⫻ 2–3 mm; stipes 30–65 cm long; blades 50–70 ⫻ 20–40 cm; pinnae 18– 20(–25) ⫻ (3.5–)4.5–10 cm, proximal pair somewhat reduced, stalked 6–8 mm, segments strongly oblique and falcate, margins often serrate or doubly serrate; sori with subentire indusia 5–7 ⫻ 0.3–0.5 mm.

0.4–0.7 cm, elliptic or rhombic, bases cuneate basiscopically, slightly auriculate acroscopically, apices caudate-acuminate, margins shallowly lobed to undulate and with lobes faintly serrulate; veins free, main lateral veins from costae 4–6-branched (2–3 pairs), branches subabruptly curved and running toward pinna margins; indument absent on both sides of blades except for minute papillae ⬍ 0.1 mm in adaxial costal grooves; sori along 1–3 acroscopic and also along 0–2 basiscopic veins of a vein group, with indusia 2.5–9.5 ⫻ 0.2–0.4 mm, entire.

Distribution. Terrestrial in lowland and montane rain forests; 100–1200(–1700) m. Mexico; Guat. Selected Specimens Examined. Chis (Croat 43895a, UC; Ishiki et al. 2031, NY). Oax (Herna´ndez G. 280, 1968, CAS, CHAPA, NY, 1686, CAS, CHAPA, NY, UAMIZ; Mickel 7165, UC; Wendt et al. 2761, CAS, CHAPA, ENCB, NY). Tab (Cowan 1725, CAS, NY, 3131, CAS, ENCB, NY). Ver (Dorantes 3551, IEB; Ibarra M. 1986, CHAPA; Martı´nez C. 2184, MO, NY; Sinaca C. 909, MO). Unverified, Doubtful, or Mistaken Reports. Me´x (Tejero-Dı´ez 1986, IZTA, cited by Tejero-Dı´ez & Arreguı´n-Sa´nchez, 2004, but not verified).

See discussion under D. lonchophyllum, the closest relative, for differences between D. drepanolobium and a discussion of putative hybrids. Mickel 5222 (NY), determined as this species by Mickel & Beitel (1988), is D. franconis.

6. DIPLAZIUM ERRANS Diplazium errans Lorea-Hern. & A. R. Sm., Amer. Fern J. 89: 181. 1999. Type. Mexico. Guerrero: Mpio. Petatla´n, 10 km NNE of El Mameyal, dirt road Papanoa–Corrales, Campos 1531 (XAL; isotypes NY!, UC!). Figs. 116A, B. Rhizomes ascending to erect; rhizome scales concolorous, dark brown, lustrous, 0.5–1 ⫻ 0.3–0.5 mm, lanceolate, entire; fronds ca. 100 cm long; stipes pale gray-green or dark green, 34–40 cm ⫻ 2–3 mm, 1⁄3–2⁄5 the frond length, glabrous except for brown, dull scales at bases; blades thick-herbaceous to subcoriaceous, 1pinnate, 52–66 ⫻ 17–22 cm, lanceate, apices pinnatifid, one or two buds developing in pinna axils on distal 1⁄4 of blades; rachises lacking hairs and scales, but with papillae ⬍0.1 mm long in adaxially grooves; pinnae stalked 5–7 mm, falcate, inequilateral, 20–28 pairs per blade, largest pinnae (usually second or third pair) 11–15 ⫻ 1.7–2.5 cm, lanceolate, smallest pinnae 0.7–1.3 ⫻

Distribution. Terrestrial in montane rain forests; 1000 m. Mexico. Known only from the type collection. This species is mostly closely related to D. werckleanum, which agrees in having 1-pinnate blades, shallowly lobed pinnae, and similar venation and soriation. However, D. werckleanum has at most 10 (usually 5–10) free pinna pairs per blade and lacks buds, while D. errans has more than 20 free pinna pairs per blade and bears buds in the axils of distal pinnae. Another related gemmiferous species in Mexico is D. vera-pax, but the latter has only 1–3 free pinna pairs. It seems unlikely that D. errans is a hybrid involving either D. werckleanum or D. vera-pax, because no other 1-pinnate Diplazium with inequilateral pinna bases, more than 20 pinna pairs, glabrous blades, and proliferous buds is known to occur in Mexico or Mesoamerica. Diplazium errans is known only from the type collection; however, the species was not rare at the type locality, according to observations by the collector. Pine-oak forests at ca. 1000 meters on the western slopes of the Sierra Madre, where D. errans grows, are not common in Guerrero, having been mostly cut in farming and logging operations.

7. DIPLAZIUM EXPANSUM Diplazium expansum Willd., Sp. Pl., ed. 4, 5(1): 354. 1810. Figs. 114A, B. Asplenium expansum (Willd.) C. Presl, Reliq. Haenk. 1(1): 46. 1825. Athyrium expansum (Willd.) Milde, Bot. Zeitung (Berlin) 28: 353. 1870, hom. illeg., non T. Moore, 1860. Type. Venezuela. Caracas, Bredemeyer s.n. (B-Willd. 19948!).

Rhizomes erect; rhizome scales brown to black, clathrate with horizontal walls gone, leaving side-wall latticework (scales reti-

DIPLAZIUM FRANCONIS

form), 5–7(–15) ⫻ 2–3 mm; fronds 100–200(–300) cm long; stipes tan except for darkened bases, 55–100 cm long, 1⁄3–1⁄2 the frond length, scaly at bases, glabrous or glabrescent distally; blades thick-herbaceous to chartaceous, 2-pinnate-pinnatifid, to 100 cm long, deltate to ovate, apices pinnatifid, lacking buds; rachises with often numerous hairs 0.1–0.4 mm, lacking papillae in adaxial grooves; pinnae stalked 1–4.5 cm, equilateral, (17–)25–35 ⫻ 8–13 cm, pinnules sessile to stalked 1–2 mm, equilateral, cut 1⁄2– 3⁄4 of distance to costules, lobes obtuse, crenulate to subentire, costae and costules adaxially winged perpendicular to the plane of the blades; indument adaxially of hairs 0.1 mm long in rachial, costal, and costular grooves, abaxially with scattered tan linearlanceolate scales along costae and costules, scales 0.5–2 mm long, the blades also with usually moderate to dense, often septate hairs 0.1–0.3 mm long on costae, costules, veins, and laminae between veins; veins free, pinnate, 4–8 pairs in ultimate segments, simple or 1-forked; sori pinnate on ultimate segments, with erose indusia 3–5 ⫻ 0.3–0.5 mm; 2n⫽ca. 328 (Jam).

Distribution. Terrestrial in montane rain forests; (100–)800– 1650(–2600) m. Mexico; Guat, CR; Gr Ant; Col, Ven, Ec, Peru, s Braz, Bol. Adams (in Davidse et al., 1995) also cited a collection from Nicaragua. Proctor (1977) reported the species from St. Vincent, but this may be based on a misidentification of D. striatum (e.g., Morton 5469, UC). Selected Specimens Examined. Gro (Lorea 3007, FCME, NY). Hgo (Herna´ndez M. 3897, UC; Kenoyer s.n., ARIZ). Oax (Hallberg 1450, NY; Mickel 973, NY, UC). Pue (Riba et al. 300, NY; Ventura A. 21620, ENCB, UAMIZ). Ver (Pringle 7886, MEXU, MO, US; Ventura A. 4927, CHAPA, ENCB, NY, 14372, ARIZ, ENCB).

Among the 2-pinnate-pinnatifid taxa, the combination of retiform rhizome scales, minute hairs in the rachial and costal grooves, and equilateral pinnae distinguish D. expansum, D. hellwigii, and D. hians from the other taxa. The presence of hairs on the rachises, costae, veins and laminae between veins, the paucity of rachial and costal scales, and erose indusia 0.3–0.5 mm wide distinguish D. expansum from D. hellwigii, which has numerous rachial and costal scales, no hairs on rachises, costae, veins, or tissue between veins abaxially, and narrower, laciniateciliate indusia (uncut portion 0.1–0.3 mm wide). Occasional specimens of D. expansum have more numerous costal scales and rather sparse laminar hairs. Diplazium hians also lacks hairs on

257

the rachises, costae, veins, and tissue between veins abaxially, but has erose indusia 0.3–0.5 mm wide and narrow scales on the costae and costules abaxially. Diplazium expansum is similar to D. ingens Christ of Costa Rica, which is glabrous except for a few minute hairs on the costules and hairs in the rachial and costal grooves adaxially; that species also has narrower, ciliate indusia (uncut portion 0.1–0.3 mm wide).

8. DIPLAZIUM FRANCONIS Diplazium franconis Liebm., Mexic. Bregn. 256 (reprint 104). 1849. Figs. 113E, F. Asplenium franconis (Liebm.) Mett., Abh. Senckenberg. Naturf. Ges. 3: 166, t. 5, f. 30. 1859. Lectotype (chosen by Smith, 1981: 95). Mexico. Oaxaca: Between Tonaguia and Roayaga, Liebmann s.n. [Pl. Mex. 2664, Fl. Mex. 790] (C!–photo US!; isolectotypes B!, K!, P!, frag. NY!, US!). Asplenium aspidiiforme Fe´e, Me´m. Foug. 5: 199. 1852. Type. Mexico. [Oaxaca:] Tepitongo, Galeotti 6483 (P!–2 sheets, photo US!; isotypes BR!–2 sheets, photo US!, LE!, P!, frag. NY!). Asplenium distans Fe´e, Me´m Foug. 5: 198. 1852, hom. illeg., non D. Don, 1825. Type. Mexico. Veracruz: Cerro San Martı´n, Galeotti 6579 (BR!–photo US!; isotypes P!–photo US!, RB). Morton’s discussion (Contr. U.S. Natl. Herb. 38: 221. 1973) of the typification of this name is incorrect; the type is Galeotti 6579, as stated in the protologue and notebooks at BR, and not Galeotti 6529, from Villa Alta, Oaxaca (as presumed by Morton). Both Galeotti 6579 and 6529 are Diplazium franconis. Diplazium camptocarpon Fe´e, Me´m. Foug. 8: 84. 1857. Type. Mexico. Veracruz: Co´rdoba, Nieto s.n. (Schaffner 69) (RB; not found at P). Asplenium franconis (Liebm.) Mett. var. gracile E. Fourn., Mexic. Pl. 1: 109. 1872. Type. Mexico. Chiquihuite, Bourgeau 2149bis (P!–2 sheets; isotype K!, frag. NY!). Asplenium shepherdii Spreng. var. bipinnatum Christ, Bot. Gaz. 20: 545. 1895. Type. Guatemala. Santa Rosa: Volca´n Tecuamburro, Heyde & Lux s.n. [Donn. Sm. 4680] (isotypes NY!, US!).

Rhizomes erect; rhizome scales dark brown, linear-lanceolate, dull to lustrous, 5–9 ⫻ 1–1.5 mm, entire; fronds (50–)70–170 cm long; stipes green to stramineous or brownish, darker at bases, 25–80 cm ⫻ (2–)3–6 mm, ca. 1⁄2 the frond length, often with dark brown scales at bases, glabrous distally; blades chartaceous, 2-pinnate to 2-pinnate-pinnatifid, 30–100 ⫻ 25–70 cm, ovatelanceolate, apices pinnatifid, lacking buds; rachises glabrous or occasionally with papillae ca. 0.05 mm long in adaxial grooves; pinnae stalked 5–7 mm toward their bases, sessile or adnate distally, to ca. 6 free pairs, (10–)15–26 ⫻ 3–9 cm, inequilateral, pinnules sessile to adnate, infrequently stalked 1–2 mm, also inequilateral, 2–7 ⫻ 0.8–2 cm, cut 1⁄3–3⁄4 of distance to costules, narrowly to broadly cuneate proximally, more deeply lobed acroscopically than basiscopically, segment margins dentate; indument essentially absent on both surfaces, or adaxially of sparse minute papillae 0.05 mm in costal grooves, abaxially sometimes with sparse brown, septate hairs to ca. 1 mm long; veins free, mostly 5–8 pairs per segment, forked; sori pinnate, on the anterior veinlet of a forked pair, with entire to erose indusia mostly 2–4 ⫻ 0.5–0.8 mm; 2n⫽164 (Oax).

258

DIPLAZIUM

and, sometimes, a single acroscopic shallow lobe; also, there are fewer free pinnules on the proximal pinnae.

9. DIPLAZIUM GOMEZIANUM Diplazium gomezianum C. D. Adams, Novon 2: 294. 1992. Type. Costa Rica. Puntarenas: Road to Puerto Jime´nez, Osa, 40 km W of IA route 2, Go´mez 19533 (MO; isotype UC!). Figs. 115E, F.

Distribution. Terrestrial in wet forests; 150–2450 m. Mexico; Guat, Bel (Davidse 36869, UC), Hond, Salv, Nic, CR, Pan. Selected Specimens Examined. Chis (Breedlove 22671, NY). Gro (Lorea 1949, FCME, IEB). Hgo (Rzedowski 27653, IEB, NY). Me´x (Hinton 2768, US). Mich (Rzedowski 46052, ENCB). Mor (Lyonnet 550800002, MEXU). Oax (Mickel 4114, ENCB, UC). Pue (Arreguı´n 541, ARIZ, CHAPA, NY). Qro (Carranza 3851, IEB, UC; Rzedowski 46583, IEB). SLP (Pringle 3958, DS, NY, UC, US). Ver (Copeland herb. 44, UC, US).

Diplazium franconis is extremely variable and more or less intermediate in blade division between D. lonchophyllum (barely 2-pinnate proximally, with usually a single free acroscopic pinnule on lower pinnae) and D. donnell-smithii (3-pinnate). Diplazium franconis is distinguished from D. donnell-smithii (q.v.) by the less dissected blades and absence of numerous hair-like scales on the rachises and costae abaxially. Small fronds of D. franconis can be mistaken for unusually divided forms of D. lonchophyllum. Hybridization has probably occurred between D. franconis and D. drepanolobium in Chiapas (Smith, 1981) producing putative hybrids with misshapen spores (e.g., Breedlove 22293, 22484, DS, NY). Mickel 5368 (NY), from Oaxaca, also has malformed spores and was reported as “hexaploid” with 90 pairs and 93 univalents (Smith & Mickel, 1977), but this number is well beyond hexaploid, closer to heptaploid. The specimen is intermediate between D. franconis and D. donnell-smithii and may represent the hybrid between those two species. Mickel 1619 (NY, US), also from Oaxaca, was reported by Mickel and Beitel (1988) as D. drepanolobium ⫻ lonchophyllum, but we now see it more likely as D. franconis ⫻ lonchophyllum. This collection was reported as triploid, ca. 41II ⫹ 75I in meiosis (Mickel et al., 1966; Mickel & Beitel, 1988), but this number more closely approximates what would be expected of a tetraploid in Diplazium (2n⫽164). It also suggests that D. franconis and D. lonchophyllum have a genome in common. Breedlove 36649 (DS, CAS), from Chiapas, has malformed spores and might also be D. franconis ⫻ lonchophyllum. The parentage of these and other putative hybrids in this group cannot be resolved satisfactorily until further cytological and probably also isozymic studies are done. Specimens seen from the states of Quere´tero, San Luis Potosı´, and Hidalgo are less dissected than those in southern Mexico. Pinnules are often rounded at the tips, with crenulate margins

Rhizomes short-creeping; rhizome scales brown, lustrous, linear, 15 ⫻ 0.5–1.5 mm, subentire, remotely denticulate; fronds 150– 300⫹ cm long; stipes darkened proximally, tan to stramineous distally, 50–120 cm ⫻ 10–18 mm, ca. 1⁄2 the frond length, scaly at bases; blades herbaceous, 3-pinnate-pinnatifid proximally, ca. 100–150 ⫻ 75–100 cm, ovate, apices pinnate-pinnatifid and ultimately pinnatifid, gradually reduced, lacking buds; rachises glabrous; pinnae stalked to 3 cm, equilateral, 30–60 ⫻ 12–30 cm, the proximal pair slightly shortened, pinnules stalked 1–3 mm or sessile, to 15 ⫻ 4 cm, 14–20 pairs per pinna, costae and costules broadly green-winged in a plane perpendicular to the blade, wings 0.3–0.5 mm wide; veins free, pinnate, 3–5 pairs per segment, simple or 1-forked in each ultimate segment; indument adaxially of sparse to moderately dense hairs ⬍ 0.1 mm long in costal and costular grooves, abaxially with scattered similar hairs on the costae and costules and also with longer brownish septate hairs or linear scales to 1 mm long; sori pinnate in ultimate segments, to 7 pairs per segment, with tan, entire indusia to 4 ⫻ 0.3–0.5 mm.

Distribution. Terrestrial in montane rain forests; ca. 1000 m. Mexico; CR, Pan. Specimen Examined. Chis (Purpus 7112, UC).

Diplazium gomezianum differs from congeners in Mexico by having fully 3-pinnate to 3-pinnate-pinnatifid blades proximally and nearly equilateral pinnules. Specimens of this species were previously identified as D. herbaceum Fe´e (Smith, 1981: 96), a name that should be restricted to a Brazilian species, according to Adams (1992).

DIPLAZIUM HIANS

10. DIPLAZIUM GRANDIFOLIUM Diplazium grandifolium (Sw.) Sw., J. Bot. (Schrader) 1800(2): 62. 1801. Figs. 116C–E. Asplenium grandifolium Sw., Prodr. 130. 1788. Type. Jamaica. Swartz s.n. (S– photo US; isotype B-Willd. 19946, microfiche UC!).

Rhizomes erect, stout, trunk-like, caudices 1.5–2.5 cm diam.; rhizome scales dark brown or blackish, lanceolate, mostly 3–7 ⫻ 0.5–1.5 mm, entire; fronds to 120 cm long; stipes purplish to brownish, 35–60 cm ⫻ 2–4 mm, 1⁄3 to nearly 1⁄2 the frond length, scaly at bases, otherwise glabrous; blades chartaceous, 1-pinnate, 30–70 ⫻ 15–30 cm, apices pinnatifid, buds absent; rachises abaxially with scattered minute hairs 0.1–0.2 mm long; pinnae stalked to 5 mm, equilateral, 9–12(–15) pairs per blade, oblong to lanceolate, 11–15 ⫻ 2–4 cm, bases rounded to truncate, apices acuminate, the margins subentire to broadly crenate, or serrate towards apices; veins free, 2–3(–4)-forked, all branches strongly ascending and becoming Ⳳ parallel as they approach the pinna margins; indument adaxially of dense hairs 0.1 mm long in rachial and costal grooves, abaxially with scattered brownish linear hair-like scales and septate hairs to ca. 1 mm long; sori 8–15 ⫻ 0.5 mm, those on first acroscopic veinlet usually diplazioid, those on first basiscopic veinlet asplenioid, sori often produced only on acroscopic veins, and thus appearing Ⳳ parallel along the length of the pinnae, or on one acroscopic and one basiscopic vein of a vein group, indusia brown with paler, delicate, minutely erose to subentire margins; 2n⫽ca. 328 (Jam, Trin).

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lack of buds in axils of distal pinnae, and 2–3-forked veins running in Ⳳ parallel fashion to pinna margins.

11. DIPLAZIUM HELLWIGII Diplazium hellwigii Mickel & Beitel, Mem. New York Bot. Gard. 46: 155. 1988. Type. Mexico. Oaxaca: Distrito Ixtla´n, Km 110 on road to Valle Nacional, Hellwig 404 (NY!; isotype UC!). Figs. 114C, D. Similar to D. hians except: indument abaxially of scattered brown, entire scales 0.5–1.5 ⫻ 0.1–0.5 mm on rachises and costae, adaxially glabrous except for minute hairs 0.1 mm long in rachial, costal, and costular grooves; sori with deeply laciniateciliate indusia 0.1–0.3 mm wide.

Distribution. Terrestrial in wet montane forests; 1200–2000 m. Mexico. Specimens Examined. Gro (Lorea 4573, FCME, IEB; Lozano V. 728, FCME). Oax (Hellwig 403, NY, UC).

Diplazium hellwigii possesses characters of both D. expansum (pubescence in costal grooves and retiform scales) and D. diplazioides (numerous scales, lack of laminar hairs, and narrow, laciniate-ciliate indusia); this could indicate a hybrid origin for this taxon. See further discussion under D. expansum. The Lorea collection cited from Guerrero was identified as D. hians by Lorea-Herna´ndez & Vela´zquez M. (1998). Distribution. Terrestrial in montane rain forests; 150 m. Mexico; Guat, Bel, Hond, Nic, CR, Pan; Gr Ant; Col, Ven, Trin, Tob, Ec, Peru, Braz, Bol. Specimen Examined. Chis (Martı´nez S. 25876, MEXU).

Our specimen is referable to var. grandifolium; var. andicola Stolze occurs in Andean South America (Stolze et al., 1994), and that entity may be sufficiently distinct to warrant recognition at species rank. Diplazium grandifolium is distinguished from congeners in Mexico by the 1-pinnate blades ending in pinnatifid or confluent triangular apices, entire or faintly crenate pinnae,

12. DIPLAZIUM HIANS Diplazium hians Kunze ex Klotzsch, Linnaea 20: 361. 1847. Type. Venezuela. Me´rida, Moritz 289 p.p. (B, frag. NY!, US; isotypes BM, K!–2 sheets, frag. NY!). Fig. 112H. Diplazium amplum Liebm., Mexic. Bregn. 256 (reprint 104). 1849. Lectotype (chosen by Smith, 1981: 96). Mexico. Veracruz: Mirador, Liebmann s.n. [Pl. Mex. 2384, Fl. Mex. 799] (C!; isolectotypes K!, US!, frag. NY!).

Differs from D. expansum by: pinnae with pinnules stalked 4–5 mm; indument abaxially of sparse, brown, linear to lanceo-

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late or narrowly ovate, entire or erose scales 0.5–5 mm long on costae and costules, adaxially the blades glabrous except for minute hairs 0.1 mm long in rachial, costal, and costular grooves; sori with erose indusia 0.3–0.5 mm wide; 2n⫽328 (Jam).

apices, buds absent; rachises glabrescent or with scattered hairlike scales; pinnae stalked 4–10 mm, 20–40 ⫻ 3.5–4.5(–6.5) cm, equilateral, 12–20 pairs per blade, linear-lanceolate, cut usually 3 ⁄5–4⁄5 the way to the costae, bases truncate, apices acuminate, segments 1.8–2(–3) ⫻ 0.5–1 cm, to ca. 40 pairs per pinna, 1 or 2 proximal pairs shortened, all segments broadly adnate and confluent with adjacent segments, obtuse and dentate at the tips; veins free, 10–15 pairs per segment, simple or the basal ones in each segment 1-forked; indument adaxially of dense hairs 0.1 mm long in rachial and costal grooves, abaxially the costae and segment midveins with lax, brown septate hairs and hair-like or linear-lanceolate scales 0.5–2 mm long; sori pinnate, 10–15 pairs per segment, with very narrow (⬍0.2 mm), caducous, erose indusia 1–4 ⫻ 0.1–0.2 mm, indusia often hidden by sporangia or seemingly absent in some specimens.

Distribution. Wet montane forests; 1050–2150 m. Mexico; Guat, Hond, CR, Pan; Jam; Col, Ven, Ec, Peru, Bol, s Braz. Adams (in Davidse et al., 1995) also cited Hispaniola in the range. Selected Specimens Examined. Chis (Breedlove 23133, DS, MO, NY, 34613, DS, MO). Gro (Lorea 2337, NY, 2337A, FCME, IEB). Oax (Hallberg 1802, UC; Geith 95-M107, NY). Ver (Mu¨ller 445, NY; Palacios-Rios 3369, XAL; Ventura A. 10056, ASU, CAS, ENCB, XAL).

Stolze et al. (1994) restricted the application of this name to plants from South America, but specimens from Mesoamerica seem conspecific. Specimens determined by Stolze (1981) from Guatemala and elsewhere as D. diplazioides seem mostly to be D. hians. Fink 120 (UC), cited by Palacios-Rios (1992) as this species, is D. expansum. For differences with D. expansum, see discussion under that species.

13. DIPLAZIUM LINDBERGII Diplazium lindbergii (Mett.) Christ in Pittier, Prim. Fl. Costaric. 3(1): 27. 1901. Figs. 114J, K. Asplenium lindbergii Mett., Ann. Sci. Nat. Bot., se´r. 5, 2: 236. 1864. Lectotype (chosen by Lellinger, Proc. Biol. Soc. Wash. 89: 707. 1977). Brazil. Lindberg 543 (B). Asplenium induratum Christ, Bull. Soc. Roy. Bot. Belgique 35(1): 201. 1896, hom. illeg., non Hook., 1861. Diplazium induratum Diels, Nat. Pflanzenfam. 1(4): 226. 1899. Lectotype (chosen by Lellinger, Proc. Biol. Soc. Wash. 89: 706. 1977). Costa Rica. Limo´n: Cordillera de Talamanca, Tsaˆki, Tonduz s.n. (Herb. inst. physico-geogr. nat. costaric. 9438) (US!; isolectotype BR).

Rhizomes erect; rhizome scales dark brown, lustrous, linearlanceolate, twisted, 10–20 ⫻ 2–4 mm; fronds 100–200 cm long; stipes atropurpureous throughout, 40–70(–100) cm ⫻ 5–10 mm, ca. 1⁄3 the frond length, bases persistently scaly for at least a third of their length, with abundant, erect hairs ca. 0.1 mm long in the adaxial grooves; blades thick-herbaceous to subcoriaceous, deeply pinnate-pinnatifid proximally (but never 2-pinnate), 60– 100 ⫻ 40–60(–80) cm, gradually tapering distally to pinnatifid

Distribution. Terrestrial in wet montane forests; 1200–2300 m. Mexico; Guat, Hond, Salv (Lo¨tschert 414, US), CR, Pan; Jam, Hisp; Col, Ven, Ec, Peru, s Braz, Bol. Adams (in Davidse et al., 1995) also cited Nicaragua and Uruguay in the range. Selected Specimens Examined. Chis (Breedlove 32390, DS, NY; Croat 46680, MO). Gro (Lorea 2904, FCME, NY, 3003A, FCME, IEB). Oax (Mickel 5077, 6049, NY, UC; Torres C. 11900, MO, US). Pue (Ventura A. 383, ARIZ, NY). Ver (Copeland herb. 45, UC, US; de la Sota 4052, US; Riba et al. 387, MEXU).

The differences between D. lindbergii and D. induratum (the name used by Mickel & Beitel, 1988, for this species) are minor, the latter having larger indusia and larger blade scales. Also closely related to D. lindbergii, and agreeing with that in having very narrow indusia, is D. tussacii Fe´e, from Cuba and Hispaniola. The complex needs more study. Stolze (1981) has misdetermined some plants of this species as D. striatum (e.g., Tu¨rckheim 1011, US).

14. DIPLAZIUM LONCHOPHYLLUM Diplazium lonchophyllum Kunze, Linnaea 13: 141. 1839. Type. Mexico. Veracruz: Jalapa, Schiede s.n. (isotype BM, photos MO!, NY!, UC!, US!). Figs. 111A–C. Diplazium inaequilaterum Liebm., Mexic. Bregn. 255 (reprint 103). 1849. Syntypes. Mexico. Veracruz: all collected by Liebmann s.n., “Mirador,” 1⁄43 [Pl.

DIPLAZIUM NEGLECTUM Mex. 2379, Fl. Mex. s.n.], “circa Mirador,” 3⁄42 [Pl. Mex. 2379, Fl. Mex. 355], “Mirador,” 7-11⁄41 [Pl. Mex. 2379, Fl. Mex. s.n.], “Baranca de Jovo,” 5⁄41 [Pl. Mex. 2380, Fl. Mex. 354, s.n.] (C, photos NY!, UC!, US!, frag. NY!; isosyntypes K!–2 sheets). Diplazium acutale Fe´e, Me´m. Foug. 5: 215. 1852. Type. Mexico. [Veracruz: Mirador] Galeotti 6289 (P!; isotypes BR!, K! RB).

Rhizomes erect; rhizome scales 4–6 ⫻ 1–1.5 mm, black, sclerotic, linear-lanceolate, entire; fronds (15–)45–100 cm; stipes green to stramineous, dark at bases, (3–)14–22(–40) cm ⫻ 2–3 mm, 1⁄4–2⁄5 the frond length, bases occasionally with black scales; blades chartaceous, mostly pinnate-pinnatifid, 2-pinnate only at bases, (8–)25–50 ⫻ (4–)12–20 cm, deltate to ovate-lanceolate, apices pinnatifid, buds absent; rachises glabrous; pinnae stalked (1–)3–8 mm, inequilateral, (8–)15–20 pairs per blade, (2–)8–16 ⫻ (1–)2–4(–6) cm, cut 4⁄5 to the costae or more, bases cuneate to subtruncate, basiscopic segments shorter and more oblique than acroscopic ones, an acroscopic lobe free or nearly so on proximal 1–4 pinna pairs, apices long-acuminate, lobes usually ascending, narrowly triangular to deltate, acute and serrulate at tips; veins free, pinnate, 6–9 pairs per segment, mostly 1-forked; indument adaxially absent or of sparse, papillae less than 0.1 mm long in rachial and costal grooves, blades abaxially glabrous except for brownish hair-like scales 1–2 mm long on costae; sori pinnate in segments, 6–10 pairs per segment (sometimes on both veins of a forking veinlet), with tan to brownish Ⳳ entire to erose indusia 4–6 ⫻ 0.5–1 mm; 2n⫽82 (Ver), 164 (Chis, Oax).

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study. It differs from those two species by the proximal pinnae each having a distinct, free or nearly free, basal acroscopic pinnule. Diplazium drepanolobium has longer and wider pinnae that are strongly oblique and falcate, longer rhizome scales, and narrower, subentire indusia. Intermediate specimens (Mickel 1619, 6184, NY) have malformed spores, and Mickel 1619 is tetraploid, with 41II and 75I; Mickel et al., 1966); these represent putative hybrids involving D. lonchophyllum as one parent. In Chiapas, Breedlove 21624A, 22293, 22484, and 27480 (DS) are all more or less intermediate between D. drepanolobium and D. lonchophyllum. Breedlove 33658 (DS), cited as D. prominulum Maxon by Smith (1981) from Chiapas, is D. lonchophyllum. Diplazium lonchophyllum was reported from Nuevo Leo´n by Aguirre-Clavera´n and Arreguı´n-Sa´nchez (1988), and a possible specimen is Sa´nchez 603, MEXU. This specimen varies in the direction of D. franconis.

15. DIPLAZIUM NEGLECTUM Diplazium neglectum (H. Karst.) C. Chr., Index Filic. 236. 1905. Figs. 115C, D. Asplenium neglectum H. Karst., Fl. Columb. 1: 106, pl. 52. 1860. Type. Colombia. “Cordillera Bogotensis,” 2000 m, presumably Karsten s.n. (B?).

Similar to D. obscurum except: blades with apices pinnatifid or lobed proximally; pinnae 4–5⫹ cm wide, broadest at the truncate bases, crenulate to shallowly lobed to 5 mm, lobes rounded to truncate, ca. 10 mm wide; veins pinnate from the main lateral veins, vein groups comprising 4–5 pairs of curved and ascending veinlets, each of these veinlets potentially fertile.

Distribution. Terrestrial on slopes and along streams; 750– 1550(–2450) m. Mexico; Guat, Hond, Salv, Nic, CR, Pan; Ec. Adams (in Davidse et al., 1995) also cited this species from Colombia. Selected Specimens Examined. Chis (Breedlove 33658, DS). Col (Sanders 10468, CAS, UC). Gro (Hinton 14048, 14210, MO, NY, US). Hgo (Sa´nchez M. 624, US). Jal (McVaugh 26165, IEB, NY). Me´x (Dı´az M. 214, ENCB, NY). Mich (Hinton 16258, NY, US). Mor (Pringle 10260, MO, NY, UC). Nay (McVaugh 18980, CAS, IEB, MEXU, NY). NL (Chase 7810, MO). Oax (Herna´ndez G. 301, NY). Qro (Rzedowski 46583, XAL). Tab (Cowan 3162, CAS). Ver (Seaton 305, NY). Unverified, Doubtful, or Mistaken Reports. SLP (reported by Smith, 1981, and Mickel & Beitel, 1988, but not verified).

Diplazium lonchophyllum is quite variable and, like D. cristatum and D. drepanolobium (q.v.), in need of monographic

Distribution. Terrestrial in montane rain forests, 1550–1800 m. Mexico; CR, Pan; Col. Stolze (1981) and Adams (in Davidse et al., 1995) also cited this species from Guatemala and Venezuela, respectively. Specimens Examined. Chis (Martı´nez S. 19817, MEXU; Matuda 3985, MEXU, NY, US; Purpus 6861, MO, UC, US).

Aside from D. obscurum, which has entire lateral pinnae and a Ⳳ conform terminal pinna, this species most resembles D. urticifolium, which also has shallowly lobed pinnae and buds in

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the axils of distal pinnae. Adams (in Davidse et al., 1995) cited D. neglectum from Belize on the basis of Schipp 535 (BM, K). This collection in UC appears to be D. urticifolium vel aff., not D. neglectum. Diplazium neglectum differs from D. urticifolium principally in size, the former having pinnae greater than 4 cm wide, the latter with pinnae up to 3.5 cm wide.

16. DIPLAZIUM OBSCURUM Diplazium obscurum Christ, Bull. Herb. Boissier, se´r. 2, 7: 269. 1907. Lectotype (redesignated here; Lellinger, Proc. Biol. Soc. Wash. 89: 711. 1977, selected a specimen at US as lectotype). Costa Rica. Tablazo, Biolley s.n. (P!; isolectotype US!). Biolley 60 (US) was cited as lectotype by Lellinger, but the specimen at US is without number, and the Christ herbarium is at P, not US. Figs. 115A, B. Diplazium flavescens (Mett.) Christ var. proliferum Christ, Bull. Herb. Boissier, se´r. 2, 5: 729. 1905. Type. Mexico. Chiapas: San Pablo, Mu¨nch 166 (P!; isotypes DS!, US!, frag. NY!).

Rhizomes erect or suberect, caudices 2–3 cm diam., with thick prop-like roots 2–3 mm diam.; fronds mostly 1–1.6 m long; stipes dark purplish brown, 30–45 cm ⫻ 4–7 mm, 1⁄3–1⁄2 the blade length, toward bases persistently scaly, scales brownish, to ca. 10 ⫻ 2 mm; blades subcoriaceous, pinnate with a Ⳳ conform terminal pinna that is like the lateral ones, but sometimes a little broader at the base and with 1 or 2 basal lobes and then shallowly crenate for a few cm, 45–110 ⫻ 25–50 cm, each blade with a bud in the axil of a distal pinna; rachises purplish, nearly glabrous except for minute hairs ⬍ 0.1 mm in adaxial grooves; pinnae stalked to 6 mm, equilateral, mostly 9–12 pairs per blade, 13–23 ⫻ 3–4.5 cm, oblong-elliptic, bases rounded to broadly cuneate, apices acuminate, commonly entire, but crenulate or serrulate near apices; veins free, forking 3–4 times, all veins strongly abruptly curved and running toward pinna margins, ultimately Ⳳ parallel; indument adaxially of sparse minute hairs in costal grooves, abaxially with brownish fibrils or narrowly lanceolate scales 1–3 mm long and sparse minute hairs ⬍0.1 mm long on costae; sori confined mostly to proximal half of first acroscopic branch of each vein group, mostly back-to-back, with brownish, entire indusia ca. 5–13 ⫻ 0.7 mm.

Distribution. Terrestrial in montane rain forest; 800–2000 m. Mexico; Guat, Hond, Nic (Stevens 5980, UC), CR, Pan; Col, Ven, Ec (Neill 5982, UC). Selected Specimens Examined. Chis (Breedlove 29817, DS, NY, 49319, CAS; Croat & Hannon 65231, MO, US). Gro (Lorea 3002, FCME, IEB). Oax (Ingela 95-M039, NY). Pue (Campos V. 397, IEB). Ver (Ventura A. 11616, ENCB, IEB, MEXU, XAL).

Diplazium obscurum differs from congeners in Mexico by the combination of entire pinnae (except at tips), Ⳳ conform terminal pinna, and a bud at the base of a distal pinna.

17. DIPLAZIUM PLANTAGINIFOLIUM Diplazium plantaginifolium (L.) Urb., Symb. Antill. 4: 31. 1903. Figs. 113J, K. Asplenium plantaginifolium L., Syst. Nat., ed. 10, 2: 1323. 1759. Asplenium plantagineum L., Sp. Pl., ed. 2, 2: 1537. 1763, nom. superfl. Type. Jamaica. Manchester: “Mayday-hills,” Brown s.n.; the specimen, formerly in the Linnaean Herbarium, is no longer there and is presumed lost. Neotype (designated by Proctor, 1985: 394). Jamaica. St. Catherine: Mt. Diablo, vic. of Hollymount, Maxon 1949 (US!; isoneotypes BM, NY!–2 sheets).

Rhizomes ascending to erect; rhizome scales concolorous black, lustrous, deltate, 0.8–1 ⫻ 0.5–0.8 mm; fronds (20–)30–60(–70) cm; stipes green, (10–)20–35 cm ⫻ 1–2.5 mm, 1⁄3–2⁄3 the frond length, bases with sparse black scales 2–3 mm long, otherwise glabrous; blades simple (lacking pinnae), 15–22(–30) ⫻ 3.5–6 (–8) cm, lanceolate to narrowly elliptic, bases truncate to broadly cuneate, apices abruptly narrowed to attenuate tips, margins crenulate or bicrenulate, occasionally serrulate toward apices, occasionally with a bud at blade bases; veins free, 3–5-forked, branches abruptly curving toward margins and running nearly parallel to each other; indument absent on all parts of the blades, or midribs abaxially with scattered brownish hair-like fibrils to 1 mm long; sori mostly single on veins, often along only acroscopic vein branch, occasionally diplazioid, appearing Ⳳ parallel, with entire indusia 4–30 ⫻ 0.3–0.5 mm.

DIPLAZIUM STRIATASTRUM

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lobes oblong to deltate, toothed; veins free, pinnate, 3–5 pairs per segment; indument adaxially of erect hairs less than 0.1 mm long in rachial and costal grooves, abaxially the costae, costules, and veins with erect hairs 0.1 mm long, tissue between veins glabrous; sori pinnate in the ultimate segments, with subentire indusia 3–5 ⫻ 0.5–0.8 mm.

Distribution. Terrestrial in lowland and lower montane rain forests; 150–1600 m. Mexico; Guat, Bel, Hond, Nic, CR, Pan; Gr & L Ant; Ven, Trin, Peru, s Braz, Bol, Parag. Adams (in Davidse et al., 1995) also included Colombia and Ecuador in the range, but Stolze et al. (1994) did not cite the species from Ecuador and we have seen no specimens. Selected Specimens Examined. Chis (Breedlove 67033, 70863, CAS; Dressler 1620, MEXU, NY, US). Oax (Herna´ndez G. 1006, 1645, 1626, 2553, CHAPA, NY). Pue (Pacheco 3226, UAMIZ). Ver (Purpus 1970, 1970a, UC; Ventura 8213, ARIZ, CAS, IEB, MO, NY).

Diplazium plantaginifolium is the only Mexican Diplazium with simple, undivided blades. It hybridizes with D. ternatum, forming plants with ternate blades, entire to slightly undulate margins, anterior vein branch forming areoles, and malformed spores: Hallberg 1615, NY, US, from Oaxaca, 150–250 m. This hybrid occurs at a relatively low elevation for D. ternatum. Another putative hybrid between the same two parents is Ventura A. 20222 (ENCB), from Puebla. This specimen has the terminal pinna 3–4 times longer than the two lateral pinnae and malformed spores. Other hybrids involving D. plantaginifolium have been seen from Hispaniola (Mickel 8990, NY, UC).

18. DIPLAZIUM PUBERULENTUM Diplazium puberulentum Mickel & Beitel, Mem. New York Bot. Gard. 46: 157. 1988. Type. Mexico. Oaxaca: Distrito Villa Alta, summit of ridge between Yetzelalag and Lovani, Hallberg 1543 (NY!; isotype UC!). Figs. 112D, E. Rhizomes erect; rhizome scales brown, concolorous, 1–3 ⫻ 1 mm, entire; fronds to ca. 80 cm long; stipes stramineous to tan, 27–38 cm ⫻ 3–4 mm, ca. 1⁄2 the frond length, densely puberulent on all surfaces, including adaxial grooves, the hairs ca. 0.1 mm long; blades herbaceous, 2-pinnate-pinnatifid proximally, ovate, 35–40 ⫻ 20–25 cm, apices pinnatifid, buds absent; rachises puberulent on all surfaces, hairs 0.1 mm long; pinnae stalked to 5 mm, Ⳳ inequilateral (proximal pinnae with basiscopic pinnules longer than acroscopic pinnules) to equilateral, ca. 10–12 pairs per blade, 11–15 ⫻ 6–9 cm, the largest with 8–10 pairs of inequilateral pinnules, these lobed nearly to costules at their bases,

Distribution. Terrestrial in wet forest; 1600–1750 m. Mexico. Specimens Examined. Oax (only the type). Ver (Nee & Taylor 26270, NY).

The absence of scales on the stipes, rachises, and costae and the dense minute hairs on the stipes, rachises, costae, and veins abaxially distinguish this species from congeners in Mexico. Probably, it is most closely related to D. franconis, which it resembles in having inequilateral pinnules.

19. DIPLAZIUM STRIATASTRUM Diplazium striatastrum Lellinger, Selbyana 2: 283. 1978. Type. Costa Rica. Tuı´s, near Turrialba, Pittier s.n. (Herb. inst. physicogeogr. nat. costaric. 11248) (US!, photos NY!, UC!; isotypes BM, K, MO!). Figs. 114G, H. Similar to D. striatum except: rhizome scales with straight cell walls; stipes 20–50 cm long; blades pinnate-pinnatifid proximally, 35–60(–100) ⫻ (14–)20–30(–40) cm; pinnae 7–14(–16) ⫻ 1.8–3 cm.

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Distribution. Wet montane forests; 400–2000 m. Mexico; Guat, Nic, CR, Pan; Col, Ven, Ec. Also cited from Honduras by Lellinger (1978: 284). Selected Specimens Examined. Chis (Breedlove 25063, 42626, DS). Hgo (Barkley 17M064, US; Clark 7376, MO, NY). Oax (Martı´nez C. 762, CAS; Mickel 3954, NY, UC, 4228, ENCB, NY, UC). Qro (Rzedowski 43300, IEB). Ver (Palacios-Rios 2287, UC, XAL; Purpus 1983, NY, UC, US, 4353, UC, US).

distally, 30–65 cm ⫻ 5–9 mm, 1⁄2–3⁄5 the frond length, with abundant, minute, erect, pale hairs ca. 0.1 mm long in the adaxial grooves; blades herbaceous, pinnate-pinnatifid proximally, or sometimes barely 2-pinnate, 35–100 ⫻ 25–50 cm, apices pinnatifid, buds absent; rachises glabrescent or moderately hairy; pinnae stalked 2–7 mm, equilateral, ca. 15–22 pairs per blade, 7–30 ⫻ 3–9 cm, oblong-lanceolate, proximal several pinna pairs sometimes with a few free or nearly free pinnules at bases, oblonglanceolate, cut usually 3⁄5–4⁄5(–9⁄10) the way to the costae, bases truncate, apices acuminate, pinnules or lobes 1.5–4 ⫻ 0.7–1.5 cm, slightly larger on sterile fronds, ca. 12–18 pairs per pinna, 1 or 2 proximal ones shortened, largest proximal ones broadly to narrowly adnate, distal ones broadly adnate and usually confluent, the largest pinnules lobed to 1⁄3 their width at their bases, otherwise merely crenate, spreading, obtuse at the tips; veins free, pinnate, 8–12 pairs per lobe, 1(–2)-forked; indument adaxially of abundant hairs in rachial and costal grooves, abaxially with a few scales on rachises and costae, also with hairs especially abundant on costae and costules but scattered to often dense between veins as well, hairs erect, 0.1–0.3 mm long, the longest ones septate; sori pinnate, 3–8 pairs per segment, with subentire indusia 2–8 ⫻ 0.3–0.6 mm; 2n⫽328 (Jam).

This differs from D. striatum by the traits given in the diagnosis. Riba et al. 387 (MEXU), cited as this species from Veracruz by Palacios-Rios (1992), is probably D. lindbergii; Mickel 743 (NY), also cited by Palacios-Rios as D. striatastrum, is D. striatum. Many of the specimens cited by Palacios-Rios as D. striatum from Veracruz appear to be D. striatastrum, e.g., Finck 26, 148, UC; Purpus 1983, 4353, and 15735 (UC).

20. DIPLAZIUM STRIATUM Diplazium striatum (L.) C. Presl, Suppl. Tent. Pterid. 114. 1836. Figs. 112F, G. Asplenium striatum L., Sp. Pl. 2: 1082. 1753. Type. Petiver, Pter. Amer., t. 3, f. 3, 4. 1712, which was based on Plumier, Traite´ Foug. Ame´r., pl. 18, 19, 1705, based on a collection from Martinique. Diplazium crenulatum Liebm., Mexic. Bregn. 254 (reprint 102). 1849. Lectotype (first chosen by Lellinger, Proc. Biol. Soc. Wash. 89: 710, 1977; rechosen by Mickel & Beitel, 1988: 158, a choice accepted here). Mexico. Veracruz: between Colipa and Misantla, Liebmann s.n. [Pl. Mex. 2383, Fl. Mex. 360] (C!, photo US!; isolectotypes K! frag. NY!). An unnumbered Liebmann specimen at US, from “between Colipa and Misantla,” was chosen as lectotype by Lellinger, but this specimen probably represents D. striatastrum; however, the proximal part of the blade, which might prove definitive, is absent. One of the syntypes in Copenhagen (Liebmann s.n. [Pl. Mex.] 2665, C, photos MO!, NY!) is also D. striatastrum and may be from the same gathering as Lellinger’s choice of lectotype. Diplazium elongatum Fe´e, Me´m. Foug. 5: 215. 1852. Type. Mexico. Oaxaca: [Dto. Choapan] Petlapa, Galeotti 6471 (P! photos UC!, US!; isotypes BR!–4 sheets). Diplazium matudae C. D. Adams, Novon 2: 295, f. 7. 1992. Type. Mexico. Chiapas: Escuintla, near Col. Zintalapa, 160 m, Matuda 18164 (MEXU; isotypes DS!, MO!, US!).

Rhizomes erect; rhizome scales dark brown, lustrous, somewhat retiform, linear-lanceolate, 10–15 ⫻ 2–3 mm, cell walls sinuous; fronds 100–160 cm long; stipes dark brown at bases, stramineous

Distribution. Wet montane forests; 150–2000 m. Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Ec, Peru, s Braz, Bol. Selected Specimens Examined. Chis (Breedlove 31343, DS, NY; Purpus 6736, UC, US). Gro (Croat 45808, 45808A, MO). Oax (Herna´ndez G. 1546, CHAPA; Mickel 4750, ENCB, NY, UC, 5232, NY, UC). Pue (Riba et al. 319, NY; Vera Santos 3326, ENCB). Ver (Smith 2005, UC; Ventura A. 1096, ENCB, NY). Tab (Linden 1491, FI–photo US!, K). Unverified, Doubtful, or Mistaken Reports. Hgo (reported by Mickel & Beitel, 1988, and Palacios-Rios, 1992, but not verified). Qro (Rzedowski 43300, IEB, cited by Arreguı´n et al., 2001, but this specimen is D. striatastrum).

Diplazium striatum differs from D. urticifolium in having hairs on the laminae and rachises abaxially, minute hairs in the grooves of costae and rachises adaxially, and pinnae cut 3⁄4 or more of the distance to costae. Occasional specimens of D. striatum appear glabrous abaxially, e.g., Palacios-Rios 3324 (XAL). Diplazium striatastrum (q.v.), with laminar and rachis hairs like D. striatum, was segregated from D. striatum on the basis of

DIPLAZIUM URTICIFOLIUM

several characters. In D. striatum, the proximal pinnae are fully pinnate at their bases with 1–2 free pinnules, whereas in D. striatastrum the proximal pinnae resemble the middle pinnae, being pinnatifid with adnate basal segments. The size differences given in the key also serve to separate the two species, but there is some overlap. More work is needed to determine the degree of relationship of D. striatastrum to D. striatum. It is tempting to speculate that D. striatum may have arisen following hybridization of D. striatastrum and D. expansum, because of its somewhat intermediate morphology. Diplazium matudae was compared to D. biolleyi Christ, from Costa Rica, by Adams (1992), but that species has ovate costal scales and fully 2-pinnate-pinnatifid blades. Diplazium biolleyi seems not closely related to D. striatum, but may be the same as D. marattiifolium Christ. We find no significant differences between type and paratype material of D. matudae and typical D. striatum.

21. DIPLAZIUM TERNATUM Diplazium ternatum Liebm., Mexic. Bregn. 252 (reprint 100). 1849. Figs. 113G, H. Asplenium ternatum (Liebm.) Hook., Sp. Fil. 3: 265. 1860, hom. illeg., non C. Presl, 1825. Lectotype (chosen by Smith, 1981: 98). Mexico. Oaxaca: Villa Alta, between Tonaguia and Roayaga, Liebmann s.n. [Pl. Mex. 2385; Fl. Mex. 364] (C!; isolectotypes K!, P!, US!, photo NY!).

Rhizomes short-creeping to erect; rhizome scales concolorous, dark brown to black, deltate, 0.8–1 ⫻ 0.5–0.8 mm, entire; fronds 20–55 cm long; stipes green to brownish green, 24–36 cm ⫻ 1–2 mm, 1⁄2–2⁄3 the frond length, lacking scales or hairs except at the very base; blades thick-herbaceous, pinnately tripinnate, 9–20 cm long, each blade often with a proliferous bud at the base of a lateral pinna; rachises brownish green, with a bud in the axil of one of the lateral pinnae; pinnae sessile to short-stalked 1–2 mm, 2 lateral pinnae plus a terminal pinna per blade, longacuminate, lanceolate, falcate, terminal pinna Ⳳ conform, or often slightly larger than lateral pinnae, stalked 1–2.5 cm, all 3 pinnae coarsely and doubly serrate, especially toward pinna tips; veins usually free except sometimes the anterior branch in a vein group (the primary, soriferous veinlet), which forks and usually reunites to form a narrowly elliptic areole, 2–3 pairs of veins per vein group, all curving abruptly toward pinna margins and running Ⳳ parallel to each other; indument absent on all parts of the blades; sori diplazioid or single on a given vein, often restricted to the acroscopic vein branch of a vein group, with entire indusia 5–20 ⫻ 0.5–0.7 mm wide; 2n⫽164 (Oax, Hond).

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Distribution. Terrestrial or on wet rocks in montane rain forests, mostly on Atlantic slope; 500–2200 m. Mexico; Guat, Hond, Salv, Nic. Selected Specimens Examined. Chis (Breedlove 32387, DS, NY, 34378, DS). Gro (Lorea 3025, FCME, IEB). Hgo (Herna´ndez M. 5354, US; Estrada s.n., 2 Nov 1974, NY). Oax (Herna´ndez G. 1642, CHAPA, NY; Wendt et al. 5163, CHAPA, MEXU, MO, NY). Pue (Rzedowski 32436, ENCB; Ventura A. 394, ARIZ, ENCB, NY). Ver (Ventura A. 5117, ARIZ, 15226, ENCB, IEB, NY).

This is most closely related to D. plantaginifolium, with which it agrees in the buds at the bases of blades, similar venation, and undivided pinnae. See D. plantaginifolium for a discussion of a putative hybrid.

22. DIPLAZIUM URTICIFOLIUM Diplazium urticifolium Christ in Pittier, Prim. Fl. Costaric. 3(1): 29. 1901. Type. Costa Rica. “Vallee de Tuis,” Pittier s.n. (Herb. no. 11293) (P?; isotype US!, frag. NY!). Fig. 114F. Rhizomes erect, stout, trunk-like, to 30 cm tall; rhizome scales dark brown, ovate-lanceolate to linear-lanceolate, 4–5(–10) ⫻ 1.5–3 mm, entire; fronds 65–130(–150) cm long; stipes greenish to stramineous, 10–30 cm ⫻ 4–6 mm, 1⁄3–2⁄5 the frond length, scaly at very bases, otherwise glabrous; blades thick-herbaceous, 1pinnate, 40–80 ⫻ 20–32 cm, narrowly oblong, apices pinnatifid, buds absent (ours) or, in most extraterritorial specimens, each blade bearing a bud in the axil of a distal pinna; rachises glabrous abaxially, with erect hairs 0.1–0.2 mm in rachial grooves adaxially; pinnae 7–20 pairs per blade, stalked 1.5–4(–10) mm, 12–18 ⫻ 2.5–3.5 cm, equilateral, bases truncate, apices long-acuminate, margins crenate or shallowly lobed 1⁄3 (or less) the distance to costae, lobes truncate or rounded, broader than long; veins free, pinnate, 3–4(–6) pairs per segment; indument adaxially of erect hairs 0.1–0.2 mm long in costal grooves, abaxially the blades glabrous; sori pinnate, 1–3(–4) pairs per lobe, with brown, Ⳳ entire indusia mostly 4–10 ⫻ 0.3–0.5 mm.

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DIPLAZIUM

Distribution. Terrestrial in montane rain forests; 1000–1750 m. Mexico; Guat, Bel, Hond, Nic, CR, Pan. Adams (in Davidse et al., 1995) also cited this species from Colombia, Venezuela, and Ecuador, but Stolze et al. (1994) did not treat this species for Ecuador, and we have seen no specimens from Venezuela (Smith, 1985; Smith in Steyermark et al., 1995). Selected Specimens Examined. Chis (Breedlove 32184, DS, 32629, DS, NY; Croat 46695, MO). Oax (Hallberg 1558, NY, UC, US; Mickel 972, ISC, US, 5654, NY, UC; Rzedowski 34055, ARIZ, MO). Pue (Ventura A. 409, ENCB, NY). Ver (Lyonnet 550500002, MEXU; Riba et al. 385, MEXU, NY; Ventura A. 14374, ENCB, IEB).

Diplazium urticifolium differs from congeners in Mexico by the combination of pinnatifid blade apices, shallowly lobed, equilateral pinnae, and 2–4 pairs of rather long, curving sori on regularly pinnate vein groups. Most specimens in Mesoamerica bear a bud in the axil of a distal pinna, but nearly all Mexican species appear to lack rachis buds.

23. DIPLAZIUM VERA-PAX Diplazium vera-pax (Donn. Sm.) Hieron., Hedwigia 59: 322. 1917. Figs. 115G, H. Asplenium vera-pax Donn. Sm., Bot. Gaz. 13: 77, t. 2. 1888. Type. Guatemala. Alta Verapaz: Pansamala, Tu¨rckheim s.n. [Donn. Sm. 850] (US!; isotypes K!–2 sheets, P!–2 sheets, NY!–2 sheets). Asplenium cordovense Baker, Ann. Bot. (Oxford) 8: 125. 1894. Diplazium cordovense (Baker) C. Chr., Index Filic. 230. 1905. Type. Mexico. Veracruz: near Co´rdoba, Finck [143] (K!, photo US!, drawing NY!; isotypes UC!, US!). See Maxon, Proc. Biol. Soc. Wash. 46: 142 (1933), for discussion of similarity of D. cordovense and D. vera-pax.

Similar to D. werckleanum except: blades pinnatifid to shallowly lobed in distal 1⁄3–1⁄2 of blades, 6–12 mm wide; pinnae 1– 3 free pairs per blade.

Distribution. Wet montane forests; 800–1300 m. Mexico; Guat, Bel, Nic, CR (Brener s.n., NY), Pan. Also cited from El Salvador by Seiler (1980: 50). Specimens Examined. Chis (Breedlove 33110, DS, 67989, CAS). Ver (Finck 143, K).

Diplazium vera-pax has the appearance of a hybrid between a simple-bladed species, quite likely D. plantaginifolium, and a 1-pinnate species, perhaps D. werckleanum. Mickel and Beitel (1988) and Adams (in Davidse et al., 1995) used the name D. riedelianum (Bong. ex Kuhn) Kuhn ex C. Chr., based on a Brazilian type, for this taxon, but D. riedelianum has the proximal pinna pair slightly lobed, with sori on several veins of a vein group, rather than on only the anterior vein of a vein group. It seems a distinct possibility that D. riedelianum and D. vera-pax are not conspecific and, if both hybrids, are of different parentage, since D. werckleanum is unknown from South America. Comparative isozymic or DNA sequence studies might prove illuminating.

24. DIPLAZIUM WENDTII Diplazium wendtii Mickel & A. R. Sm., Brittonia 52: 234, f. 1A–C. 2000. Type. Mexico. Oaxaca: Mpio. Sta. Marı´a Chimalapa, Can˜on del Rı´o Blanco, ca. 4.5 km S of its mouth to Rı´o del Corte, ca. 13.5 km ESE of Sta. Marı´a, Wendt et al. 5580 (NY!; isotypes MEXU, TEX, UC!). Figs. 112A–C. Rhizomes erect; rhizome scales concolorous, brown-black, lustrous, lanceolate to ovate-lanceolate, to 4 ⫻ 1.5 mm, entire; fronds 90–100 cm; stipes tan to stramineous, to ca. 40 cm ⫻ 4 mm, 2⁄5 the frond length, with dark brown scales at very bases, otherwise glabrous; blades chartaceous, 3-pinnate-pinnatifid to 4-pinnate, to ca. 60 ⫻ 48 cm, ovate, apices pinnatifid, lacking buds; rachises stramineous, glabrous; pinnae 25⫹ pairs per blade, stalked 5–7 mm, 20 ⫻ 25 cm long, inequilateral, perpendicular or ascending ca. 80⬚ from rachises, pinnules stalked ca. 2 mm, alternate, inequilateral, costules adaxially with green wings perpendicular to the plane of the blades forming deep grooves, pinnulets and some ultimate segments stalked to 1 mm, inequilateral, bases cuneate, margins toothed; veins free, pinnate in

DIPLAZIUM WERCKLEANUM

ultimate lobes, 2–4 pairs per segment; indument absent on all parts of the blades; sori pinnate in ultimate segments, with entire indusia 1.5–2.5 ⫻ 0.3 mm.

Distribution. Terrestrial in montane rain forests; 450 m. Mexico. A sterile specimen from Panama, Folsom 2662 (UC), looks suspiciously like D. wendtii but differs in having fewer and more widely spaced pinnae; if the same species, it would be a remarkable disjunction. Known only from the type specimen. Diplazium wendtii is distinguished by its highly dissected blades, pinnae perpendicular or nearly so with respect to the rachises, petiolulate inequilateral pinnae, pinnules and pinnulets, and very small ultimate segments. Of Mexican diplaziums, only D. gomezianum approaches D. wendtii in blade dissection, both species being at least 3-pinnate to 3-pinnate-pinnatifid; all other Mexican diplaziums have less dissected blades. Diplazium gomezianum also differs in having opposite to subopposite pinnules that are nearly equilateral at their bases, as well as at least some hairs or scales on the laminae abaxially, at least at the segment bases.

25. DIPLAZIUM WERCKLEANUM Diplazium werckleanum Christ, Bull. Herb. Boissier, se´r. 2, 4: 969. 1904. Type. Costa Rica. Werckle´ 81 (P?; frag. US!). Figs. 113A, B. Rhizomes erect, stout; rhizome scales black, lustrous, heavily indurate, ovate-lanceolate, 2–4 ⫻ 1–1.5 mm, entire; fronds 30– 75(–100) cm long; stipes dark at bases, light brown or stramineous distally, 10–18 cm ⫻ 1.5–3(–4) mm, 1⁄2–1⁄3 the frond length, glabrous except for blackish scales at bases; blades thickherbaceous to subcoriaceous, 1-pinnate, 15–40 ⫻ 8–20 cm, lanceolate to narrowly deltate, apices pinnatifid, lacking buds or occasionally each blade with a bud in the axil of a distal pinna; rachises stramineous to brownish, glabrous; pinnae stalked 2–4 mm, inequilateral, mostly 5–10 pairs per blade, 9–12 ⫻ 1.5–2.5(– 3.5) cm, lanceate, bases cuneate or excised basiscopically, truncate or slightly auriculate acroscopically, apices acuminate, margins undulate and serrulate to shallowly lobed 1⁄4 the distance to costae; veins free, pinnate, 3–6 branches per main lateral vein, all

267

curving abruptly toward pinna margins; indument adaxially of sparse, papillate hairs less than 0.1 mm long in rachial and costal grooves, abaxially the blades glabrous; sori often confined to first acroscopic branch of a vein group, with indusia 3–11 ⫻ 0.3–0.5 mm, erose.

Distribution. Terrestrial in montane rain forests; 550–2700 m. Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Col. Selected Specimens Examined. Chis (Breedlove 21151, 26757, DS, 48266, CAS, ENCB, NY; Purpus 7252, NY, UC, US). Gro (Anderson & Laskowski 4361, ENCB; Lorea 6454, ENCB, MEXU). Oax (Hellwig 1395, NY; Mickel 7048, NY; Wendt et al. 5025, CHAPA, MO, NY). Ver (Fink 27, UC; Purpus 15742, UC).

The D. werckleanum group (including D. cristatum and D. lonchophyllum) is taxonomically problematic in southern Mexico and Guatemala. These three species differ from each other in the degree of cutting of the proximal pinnae: in D. werckleanum the proximal pinnae are undulate to shallowly lobed 1⁄4 the distance to costae; in D. cristatum the proximal pinnae are deeply lobed without distinct pinnules and cut 1⁄2–3⁄4 the distance to costae; in D. lonchophyllum and D. drepanolobium (q.v.) the proximal pinnae have free basal pinnules that are merely serrate. There appear to be slight differences in indusial width and margins as well as rhizome scale width and length, but monographic work is necessary to determine if these differences are significant. Diplazium werckleanum may hybridize with D. lonchophyllum in Chiapas and perhaps other areas of sympatry; one such putative hybrid, having malformed spores, is Breedlove 22332 (MO). Unplaced Names Diplazium anthraxacolepis Fe´e, Me´m. Foug. 8: 84. 1857. Syntypes. Mexico. Veracruz: Huatusco, Schaffner 267b, 627a (RB). Diplazium feei Schaffner ex Fe´e, Me´m. Foug. 8: 85. 1857. Type. Mexico. Veracruz: barrancas de San Martı´n et de San Francisco, 2500 m, 1854, Schaffner 265 (RB, as cited by Windisch, Amer. Fern J. 72: 58. 1982). Schaffner s.n. (P!), bearing this name, is 2-pinnate and a little more, but less divided than D. expansum and glabrous abaxially. Excluded Species Diplazium falcatum Liebm., Mexic. Bregn. 253 (reprint 101). 1849, hom. illeg., non D. Don, 1825. Type. Mexico. Oaxaca: Teo-

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DIPLOPTERYGIUM

talcingo, Chinantla, Liebmann s.n. [Pl. Mex. 2375, Fl. Mex. 361] (C!). Maxon, in 1929, annotated this specimen as “young Hemitelia sp.” We agree that it is Cyatheaceae, but cannot say what species it might be. Diplazium pectinatum (Fe´e) C. Chr. in Urban, Symb. Antill. 9: 324. 1925. Hypochlamys pectinata Fe´e, Me´m. Foug. 5: 200, t. 17C, f. 2. 1852. Type. Hispaniola. de Tussac s.n. (P?). Diplazium pectinatum (synonym Allantodia costalis Desv.) was included in the index for pteridophytes from Oaxaca (Mickel & Beitel, 1988: 551), but there is no mention of this species on the page cited (p. 159). It is closely related to D. lindbergii, but differs by the more deeply incised pinnae, with segments cut nearly to the costae, thinner blade texture, and segments spreading at a right angle to the costae. We find no other mention of the species having been recorded from Mexico or Central America. Diplazium pectinatum is confined to Jamaica and Hispaniola (Proctor, 1985), as generally construed. Diplazium prominulum Maxon, Contr. U.S. Natl. Herb. 13: 15. 1909. Type. Guatemala. Baja Verapaz. Between Purulha´ and Panza´l, Tu¨rckheim II.1683 (US; isotype P!). Diplazium prominulum was cited from Chiapas (Smith, 1981; Smith in Breedlove 1986) on the basis of Breedlove 33658 (DS). Adams (in Davidse et al., 1995) redetermined this specimen as D. lonchophyllum, and we agree. The distribution of the species as currently known is Guat, Nic, CR, Pan; Ec (Stolze et al., 1994). Diplazium prominulum is similar to D. cristatum and D. lonchophyllum except: rhizomes erect, apices lacking scales; rachises yellowish, flexuous; blades subcoriaceous; segments rounded at tips, soriferous throughout their length; and veins free, prominent abaxially. Diplazium villosum C. Presl, Suppl. Tent. Pterid. 114. 1836. The type was said by Presl to be from Mexico (Haenke s.n., probable isotypes BM!, K!, frag. NY!). This is not a Diplazium, but a juvenile specimen of a species of Cyatheaceae, probably Sphaeropteris, possibly S. integra (J. Sm.) R. M. Tryon, from the Philippines.

mant buds with stipule-like outgrowths (pseudostipules); pinnae determinate, not forking, 2 to many opposite pairs, large, to several meters long, pinnate-pinnatisect to bipinnate, with alternating pinnate or pinnatisect pinnules, costae and costules sometimes bearing lanceolate or variously dissected scales and hairs, blade tissue sometimes glaucous; veins free, 1-forked; sori round, with or without paraphyses, of 2–4(–5) globose sporangia; indusia absent; spores tetrahedral, surfaces smooth or nearly so; x⫽56. Diplopterygium comprises about 10 species, all in the Paleotropics (India to Japan and Malesia), except D. bancroftii. The genus is distinct from other genera of Gleicheniaceae by the large, bipinnate or pinnate-pinnatisect pinnae. In this character it resembles the Old World genus Gleichenia, which has much smaller, often bead-like or pouch-like ultimate segments. The penultimate divisions of Sticherus, Dicranopteris, and Gleichenella are all pinnatifid or pinnatisect. Diplopterygium also differs from both Dicranopteris and Gleichenella by the dormant buds with scales, rather than hairs, and by the 1-forked (vs. 2–4forked) veins in the ultimate segments. Affinities of Diplopterygium are apparently not close to any of the other genera of Gleicheniaceae. For a discussion of the family relationships, see comments under Gleichenella. References Ching, R. C. 1940. On the genus Gleichenia. Sunyatsenia 5: 269–288; Holttum, R. E. 1957. Morphology, growth-habit, and classification in the family Gleicheniaceae. Phytomorphology 7: 168–184; Maxon, W. R. 1909. Gleicheniaceae. N. Amer. Fl. 16: 53–63; Nakai, T. 1950. New classification of Gleicheniales, etc. Bull. Natl. Sci. Mus., Tokyo 29: 1–71; Østergaard Andersen, E. & B. Øllgaard. 1996. A note on some morphological terms of the leaf in the Gleicheniaceae. Amer. Fern J. 86: 52–57; Østergaard Andersen, E. & B. Øllgaard. 2001. 10. Gleicheniaceae. In: G. Harling & L. Andersson (eds.), Flora of Ecuador 66: 105– 170.

1. DIPLOPTERYGIUM BANCROFTII

41 . DI PL O PT E R YG I UM

Diplopterygium bancroftii (Hook.) A. R. Sm., Amer. Fern J. 70: 26. 1980. Figs. 117A–F.

Diplopterygium (Diels) Nakai, Bull. Natl. Sci. Mus. 29: 47. 1950. Gleichenia subg. Mertensia sect. Diplopterygium Diels in Engler & Prantl, Nat. Pflanzenfam. 1(4): 353, f. 188A. 1900. Gleichenia subg. Diplopterygium (Diels) Holttum, Reinwardtia 4: 261. 1957. Lectotype (chosen by Christensen, Index Filic. LIV. 1906): Gleichenia glauca (Thunb. ex Houtt.) Hook. [⬅ Polypodium glaucum Thunb. ex Houtt.] ⬅ Diplopterygium glaucum (Thunb. ex Houtt.) Nakai.

Gleichenia bancroftii Hook., Sp. Fil. 1: 5, t. 4A. 1844. Mertensia bancroftii (Hook.) Kunze, Linnaea 18: 307. 1844. Dicranopteris bancroftii (Hook.) Underw., Bull. Torrey Bot. Club 34: 252. 1907. Hicriopteris bancroftii (Hook.) Ching, Sunyatsenia 5: 278. 1940. Lectotype (chosen here). Jamaica. Bancroft s.n. (K!; isolectotype BM!). Mertensia bancroftii (Hook.) Kunze var. vitellina Kunze, Linnaea 18: 308. 1844. Syntypes. Mexico, Leibold 108; Venezuela, Caracas, Moritz 11 (LZ destroyed; isosyntype of Moritz 11 at B). The type was incorrectly stated by Lellinger (1989: 231) to be Moritz 11.

Terrestrial, commonly scandent; rhizomes long-creeping, cord-like, protostelic, bearing deciduous, lanceolate scales; fronds monomorphic, erect or scandent; blades pseudodichotomous at top of stipes, with an arrested scaly bud at the tip, this surrounded by scales, or this bud developed and the rachis Ⳳ indeterminate, continuing to produce two opposite pinnae over time, rachis eventually ending in an arrested bud; accessory branches (borne basiscopically at the base of a fork) absent; dor-

Rhizomes long-creeping, 2–5 mm diam.; rhizome scales appressed, castaneous, linear-lanceolate, 3–4(–8) ⫻ 0.3–0.5 mm, margins entire or with minute bumps, deciduous leaving a smooth surface; fronds to more than 5 m long, erect to scandent; stipes stout, 1.5–2.5 m long, 3–5 mm diam., stramineous to light brown, glabrescent, smooth to slightly rough from fallen scales; blades with apex dormant, with only two large, bipinnate pinnae or rachis continuing and producing additional pairs of opposite

DORYOPTERIS

pinnate; dormant buds large with abundant, entire, white to yellowish or tan, ovate to lanceolate, entire scales with cordate base, 6–10 ⫻ 1.5–2.5 mm; pinnae 100–250 ⫻ 15–30 cm, with lateral wings connecting the ultimate branches (pinnules); pinnules 5– 15 ⫻ 1.5–3(–4) cm; segments linear, distant, 7–20 ⫻ 2–3 mm, decurrent, chartaceous to subcoriaceous, revolute, scaly when young, glabrescent with rare, short, clavate hairs 0.1 mm long; sori inframedial between segment margins and midveins, sporangia 3–5 per sorus, mixed with a few golden hairs 0.5 mm long; 2n⫽112 (Oax, Jam).

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42 . D O R Y O P T E R I S Doryopteris J. Sm., J. Bot. (Hooker) 4: 162. 1841. Lectotype (chosen by Morton, Amer. Fern J. 34: 26. 1944): Doryopteris palmata (Willd.) J. Sm. [⬅ Pteris palmata Willd.]. Terrestrial; rhizomes compact, erect, scaly; rhizome scales bicolorous with narrow, dark brown centers and wide, light brown margins; fronds small, monomorphic to subdimorphic, not articulate; stipes atropurpureous to black; blades usually pedate, pentagonal or deltate, laminae chartaceous to coriaceous, glabrous; veins netted or free, usually obscure in thick lamina, usually with a marginal commissural vein; sori marginal, mostly continuous, the margins reflexed as false indusia, lacking paraphyses; spores tetrahedral-globose, crested; x⫽30. Doryopteris is a genus of about 30 species, in dry, rocky, tropical regions, mostly of the Planalto of southern Brazil, with five endemic to Madagascar, and one pantropical. There are only two species in Mexico, both rare. Doryopteris is distinct by the deeply palmately lobed blade with marginal sori. Molecular studies (Gastony & Rollo, 1995, 1998) have shown Doryopteris to be most closely allied to Trachypteris of South America and Pellaea calomelanos of Africa.

Distribution. Open lands or light woods at middle to upper elevations, pine-fir forests, alder and oak forests; 1050–2200 (–2600) m. Mexico; Guat, Hond, Salv, CR; Jam, Hisp, L Ant; Col, Ven, Ec, Peru, Bol.

Reference Tryon, R. M. 1942. A revision of the genus Doryopteris. Contr. Gray Herb. 143: 1-80.

Key to the Mexican Species of Doryopteris Selected Specimens Examined. Chis (Breedlove 32393, DS, MEXU; Purpus 6719, NY, UC, US). Gro (Martı´nez S. 4218, ENCB, MEXU; Rzedowski 18619, ENCB, NY). Oax (Mickel 1105, ENCB, MEXU, NY; Rzedowski 33798, ENCB, MEXU). Pue (Lorea 1070, FCME; Riba et al. 261, MEXU, NY). Ver (Pringle 6076, MEXU; Ventura A. 9598, UAMIZ).

Nakai (1950) applied the name Diplopterygium farinosum (Kaulf.) Nakai to this species and treated Gleichenia bancroftii Hook. as a synonym, but the basionym of the former, Mertensia farinosa Kaulf. (1827), has been more recently applied to a species now regarded as belonging to the genus Sticherus, S. farinosus (Kaulf.) Ching (see Proctor, 1977), endemic to the Lesser Antilles. Neither the collector nor locality were specified in the protologue of Mertensia farinosa according to Proctor. Østergaard Andersen and Øllgaard (2001) noted that in Ecuador, the dormant rachis buds seem to break dormancy at least annually in favorable habitats, adding a new pinna pair and dormant bud each year. This allows the species to persist in regenerating vegetation, many years after disturbance, and to re-orient the new pinnae so as to assume optimum exposure and support for the scandent fronds.

1. Venation free; proliferous buds absent; stipes glabrous, except for scattered scales at bases. ................................. 1. D. concolor. 1. Venation netted; proliferous buds present at bases of blades; stipes densely pubescent with short, brown, clavate hairs 0.1 mm or less. ............................................ 2. D. palmata.

1. DORYOPTERIS CONCOLOR Doryopteris concolor (Langsd. & Fisch.) Kuhn in v. d. Decken, Reisen Ost-Afrika 3(3): 19. 1879. Figs. 118D, E. Pteris concolor Langsd. & Fisch., Pl. Voy. Russes Monde 1: 19, t. 21. 1810. Cheilanthes concolor (Langsd. & Fisch.) R. M. Tryon & A. F. Tryon, Rhodora 83: 133. 1981. Type. South Pacific. “Archipelagi Marquesas dicti insula Nucahiva,” Langsdorff s.n. (LE?; isotypes BM!, B-Willd. 19961!).

Rhizomes short, compact; rhizome scales 3–4 ⫻ 0.3 mm; fronds clumped, to 34 cm tall; stipes castaneous, flattened to grooved adaxially, glabrous, lustrous, with scattered scales at bases; blades pedate, bipinnatifid to quadripinnatifid at bases, lacking proliferous buds, texture thin to firm; veins free, visible with transmitted light; sterile margins entire to irregularly crenulate; fertile margins extending 0.1 mm beyond bases of indusia; sori marginal, continuous (breaking at sinuses and near tips of segments); indusia entire, 0.3 mm wide; spores tan; 2n⫽60 (Parag), 2n⫽120 (Taiwan).

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DRYOPTERIS

Distribution. Lowland wet forests; 150 m. Mexico; Guat, Salv, Hond; Jam, L Ant; Col, Ven, Ec, Peru, Braz, Bol, Parag, n Arg, Uru; Africa, s Asia, Malaysia, Australia, s Pacific.

Distribution. Dry, rocky slopes, pine forests, oak woods; 850– 1850 m. Mexico; Guat, Nic, CR, Pan; Col, Ven, Ec, Galapagos, Peru, Bol.

Specimens Examined. Chis (Pe´rez Farrera 442, UAMIZ). Oax (Mickel 5973, NY). Ver (Borjas 72, XAL; Hale 19387, US).

Selected Specimens Examined. Gro (Hinton 14723, TEX, US; Martı´nez S. 5544, MEXU). Hgo (Pe´rez S. 639, ENCB). Oax (Maya J. 2054, MEXU, 3806, MEXU). Ver (Fink 16, UC; Ventura A. 9788, XAL, 11748, MEXU, MICH, XAL, 16982, XAL).

Mexican material belongs to var. concolor. Tryon (1942) was uncertain of the relationships of this species because of its free venation. Tryon and Tryon (1982) transferred this species to Cheilanthes, but molecular data (Gastony & Rollo, 1995) have shown its proper place is in Doryopteris. Doryopteris concolor differs from D. palmata by its free veins and glabrous stipes.

2. DORYOPTERIS PALMATA Doryopteris palmata (Willd.) J. Sm., J. Bot. (Hooker) 4: 163. 1841. Figs. 118A–C. Pteris palmata Willd., Sp. Pl., ed. 4, 5(1): 357. 1810. Doryopteris pedata (L.) Fe´e var. palmata (Willd.) Hicken, Revista Mus. La Plata 15: 253. 1908. Type. Venezuela. Caracas, Bredemeyer s.n. (B-Willd. 19957!; photo US).

Rhizomes short-creeping, compact, stout; rhizome scales 4–5 ⫻ 0.8 mm, bicolorous, tan with atropurpureous central streak; fronds to 45 cm long, clumped; stipes 3⁄5–3⁄4 the frond length, stout, terete, atropurpureous, lustrous, with minute brown clavate hairs 0.1 mm long or less, scaly only near bases; blades pedate, to 17 cm across, at bases with 2 buds having bicolorous scales, bipinnatifid, firm to subcoriaceous; veins netted, obscure except for dark primary veins, visible with transmitted light; sterile margins irregularly crenulate; fertile margins entire; sori discontinuous at segment tips, indusia entire, 0.5 mm wide; spores tan; 2n⫽120 (Galapagos), 2n⫽60 (Jam).

Unverified, Doubtful, or Mistaken Reports. Chis (Rovirosa 1084, cited by Rovirosa, 1909, but not verified).

Doryopteris palmata is distinguished from D. concolor by the netted veins, puberulent stipe, and buds at the base of the blade. Doryopteris pedata, under which D. palmata has often been treated as a variety (e.g., by Tryon, 1942) is limited to the Greater and Lesser Antilles and differs in lacking proliferous buds.

43. D R Y O P T E R I S Dryopteris Adans., Fam. Pl. 2: 20. 1763, nom. cons. Type: Dryopteris filix-mas (L.) Schott [⬅ Polypodium filix-mas L.]. Terrestrial (rarely epiphytic); rhizomes usually stout, erect or ascending, or sometimes short-creeping, scaly; fronds generally medium-sized, erect or arching, generally clumped, usually monomorphic (ours) to subdimorphic, not articulate; stipes stramineous to reddish brown, in cross section with numerous vascular bundles arranged in a U-shape, the two adaxial bundles the largest, stipes scaly at bases and often distally; blades thin to coriaceous, pinnate-pinnatifid to 3-pinnate-pinnatifid (1-pinnate in a few extraterritorial species), deltate to ovate or lanceolate, broadest at base or above the base, anadromous at bases, main axes decurrent onto major ones, grooved adaxially and with the grooves continuous from one axis to the next; segments entire to toothed or spinulose, abaxially glabrous, glandular, or scaly; veins free, ending short of margins; sori abaxial, round, indusia reniform or round-reniform, each attached at a sinus, Ⳳ entire, glabrous or glandular, paraphyses absent; spores bilateral, with a variously winged perispore; x⫽41. Dryopteris comprises about 225 species (Fraser-Jenkins, 1986), largely of north-temperate regions; relatively few species occur at high elevations in the tropics. Most species of Dryopteris are

DRYOPTERIS

from southern and eastern Asia and the Himalayan region, but about 30 occur in the New World. Species have been grouped into four subgenera and 16 sections by Fraser-Jenk. (1986), but only subg. Dryopteris (Mexican species represented in four sections) and subg. Nephrocystis (H. Itoˆ) Fraser-Jenkins (D. futura) occur in Mexico. Dryopteris nubigena was left unplaced by Fraser-Jenkins, but it is closely related to D. futura; Dryopteris rossii and D. knoblochii were also unplaced but belong in sect. Cinnamomeae Fraser-Jenk. Unlike many Asian species, the American ones do not seem predisposed to apogamy. Two especially problematic species complexes occur in Mexico: one at high elevations (including D. wallichiana, D. pseudofilix-mas; in sect. Fibrillosae Ching and sect. Dryopteris, respectively, in the classification of Fraser-Jenkins, 1986), the other at low to middle elevations (including D. patula, D. cinnamomea, and relatives; sect. Cinnamomeae Fraser-Jenk.). Both complexes need monographic study. Dryopteris is closely allied to Arachniodes, but is distinct in its less divided blades and segments lacking mucronate tips. There is also a close relationship to Polystichum and Phanerophlebia,

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from which Dryopteris differs in the round-reniform indusia (vs. peltate) and generally more dissected blades. An intergeneric hybrid is known between Dryopteris goldiana (Hook. ex Goldie) A. Gray and Polystichum lonchitis (L.) Roth (⫽ ⫻Dryostichum singulare W. H. Wagner) from Ontario, Canada (Wagner et al., 1992). Hybridization is a well known phenomenon in Dryopteris of temperate regions, and may also be common between sympatric species in tropical and subtropical areas. At least two interspecific hybrid combinations are known in Mexico, involving D. pseudofilix-mas ⫻ wallichiana and D. cinnamomea ⫻ knoblochii, and others are likely. References Fraser-Jenkins, C. R. 1986. A classification of the genus Dryopteris (Pteridophyta: Dryopteridaceae). Bull. Brit. Mus. (Nat. Hist.), Bot. 14(3): 183–218; FraserJenkins, C. R. 1989. A monograph of Dryopteris (Pteridophyta: Dryopteridaceae) in the Indian subcontinent. Bull. Brit. Mus. (Nat. Hist.), Bot. 18: 323–477; Smith, A. R. & C. R. Fraser-Jenkins. 1982. Dryopteris paleacea is a synonym of D. wallichiana. Taxon 31: 326–329; Wagner Jr., W. H., F. S. Wagner, A. A. Reznicek & C. R. Werth. 1992. ⫻ Dryostichum singulare (Dryopteridaceae), a new fern nothogenus from Ontario. Canad. J. Bot. 70: 245–253.

Key to the Mexican Species of Dryopteris 1. Blades basically pinnate-pinnatifid or, at the bases, sometimes 2-pinnate-pinnatifid; stipes and rachises densely to sparsely scaly; blades firm to coriaceous; (1500–)2200–3950 m. 2. Segments truncate at tips; proximal pinnae with entire to serrate pinnules, these not different from segments above; rachises densely scaly, the scales linear-lanceolate (more than 10 times longer than wide), dark brown. ............................................... 13. D. wallichiana. 2. Segments rounded to acute, not truncate at tips; proximal pinnae with pinnatifid pinnules, these more divided than segments above; rachises sparsely to moderately scaly, the scales ovate-lanceolate (3–6 times longer than wide), light brown or tan. 3. Median pinnae fully pinnate, the pinnules crenate to lobed about halfway to costules; blades drying yellowish green. ...... 3. D. filix-mas. 3. Median pinnae merely deeply pinnatifid to pinnatisect, the pinnules entire, or toothed at the tips; blades drying dark green. ............................................................................................................................................................................................ 11. D. pseudofilix-mas. 1. Blades 2-pinnate-pinnatifid or more divided; rachises without scales or scales sparsely scattered; blade texture thin; 50–3000 m. 4. Blade surfaces (both sides) and indusia with conspicuous and abundant, yellowish to whitish clavate or cylindrical glands; 200– 1300 m. .................................................................................................................................................................................................. 5. D. karwinskyana. 4. Blade surfaces and indusia glabrous or with relatively inconspicuous, stipitate, hyaline to light yellowish glands; 50–3000 m. 5. Blades deltate, 3–4-pinnate-pinnatifid proximally; proximal pinnae much more developed basiscopically than acroscopically; 1750–3000 m. 6. Stipes stramineous at bases; pinnae strongly ascending; sori Ⳳ medial, at bases of ultimate segments. .............................. 4. D. futura. 6. Stipes castaneous at bases; pinnae Ⳳ perpendicular to rachis; sori submarginal, often confined to apices of segments. ...............................................................................................................................................................................................9. D. nubigena. 5. Blades lanceolate to deltate, 2–3-pinnate-pinnatifid proximally; proximal pinnae equilateral or inequilateral, but only slightly more developed basiscopically; 50–2600 m. 7. Stipe scales bicolorous, pale brown with blackish tips; rhizome scales dark brown, 2.5–5 mm long; indusia 0.5–0.8 mm diam. ..................................................................................................................................................................................................... 12. D. rossii. 7. Stipe scales concolorous, pale brown to brown, lacking blackish tips; rhizome scales brown, mostly 5–8 mm long; indusia 0.5–1.5 mm diam. 8. Indusia vaulted, firm, glabrous; blades essentially glandless; stipe bases stramineous. ................................................... 7. D. maxonii. 8. Indusia flattish or only slightly convex, often glandular; blades glandless to rather densely stipitate-glandular; stipe bases stramineous to often darkened. 9. Proximal part of blades 2-pinnate, with pinnules shallowly pinnatifid or lobed, lanceolate, equilateral or nearly so; stipes (distally) and rachises with persistent scales. 10. Basal basiscopic pinnule of proximal pinnae slightly shorter than the next pinnule; stipes and rachises persistently and obviously scaly; proximal pair of pinnae slightly or not at all reduced, the next pair the longest; segments strongly spinulose; Baja California. ............................................................................................. 1. D. arguta. 10. Basal basiscopic pinnule of proximal pinnae slightly longer that the next pinnule; stipes and rachises sparsely scaly; proximal several pairs of pinnae slightly reduced (the lowermost 8 cm, then the next 5 pairs gradually lengthened to 13.5 cm); segment margins weakly spinulose; Coahuila. ............................................. 3. D. filix-mas. 9. Proximal part of blades deeply 2-pinnate-pinnatifid, 3-pinnate, or more divided, deltate-lanceolate, inequilateral

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DRYOPTERIS (more strongly developed on basiscopic side); stipes (distally) and rachises glabrescent or with only a few persistent scales. 11. Stipe bases and rhizome scales bicolorous, tan with irregular, blackish streaks; blades nearly 3-pinnate at bases, the proximal pinnae deltate, about twice as long as wide, basiscopically more developed; blades eglandular, drying dark green. .................................................................................................................................. 8. D. muenchii. 11. Stipe bases and rhizome scales concolorous, tan to reddish brown; blades 2-pinnate-pinnatifid to 3-pinnatepinnatifid, the proximal pinnae lanceate to deltate, generally at least 2.5 times longer than wide, basiscopically more developed or not; blades glandular to eglandular, drying green to yellowish green. 12. Rhizome scales large (9–25 mm long), with strongly toothed and crisped margins and long, twisted, hair-like tips; stipes stramineous or darkened only at bases. .......................................................................... 10. D. patula. 12. Rhizome scales small (3–7 mm long) to long (5–18 mm long), with entire or subentire margins and long, twisted, hair-like or acute tips; stipes stramineous or reddish brown. 13. Rhizome scales generally 10–18 mm long, orangish, lustrous; proximal pinnae isodromous or nearly so; pinnae spreading, the lowermost often slightly reduced; indusia often 0.8 mm diam. or more. ....................................................................................................................................................... 2. D. cinnamomea. 13. Rhizome scales usually ca. 5 mm long or less, relatively sparse; proximal pinnae decidedly anadromous; pinnae ascending, the lowermost the longest; indusia less than 0.8 mm diam. ...... 6. D. knoblochii.

1. DRYOPTERIS ARGUTA Dryopteris arguta (Kaulf.) Maxon, Amer. Fern J. 11: 3. 1921. Figs. 120C, D. Aspidium argutum Kaulf., Enum. Filic. 242. 1824. Type. U.S.A. California: Chamisso s.n. (LE?)

Rhizomes short-creeping to suberect; rhizome scales concolorous, orange-brown, lustrous, to 20 ⫻ 5 mm, ovate-lanceolate, entire, sometimes twisted; fronds 25–100 cm long; stipes tan, to 35 cm ⫻ 6 mm, 1⁄4–1⁄3 the frond length, densely scaly at bases and often distally, scales orange-brown to tan, ovate to lanceolate, lustrous, entire; blades herbaceous, green to yellow-green, ovate-lanceolate, pinnate-pinnatifid to 2-pinnate at bases, to 60 ⫻ 8–30 cm, pinnae spreading to slightly ascending ca. 70⬚ from rachises, to 32 pairs; rachises stramineous to tan, with numerous persistent tan scales to ca. 6 ⫻ 1 mm, eglandular; proximal pinnae equilateral or the basiscopic segments slightly longer than the acroscopic ones, to ca. 14 ⫻ 3 cm, Ⳳ the same length as adjacent several pinna pairs or only slightly reduced, largest pinnules to ca. 20 ⫻ 6 mm; distal pinnae sessile or stalked to 3 mm, equilateral or nearly so, to ca. 14 ⫻ 3.5 cm, pinnule margins serrate with spinulose teeth; indument abaxially of scattered lanceolate scales on costae and costules, veins and leaf tissue between veins glabrous, lacking glands, adaxially the surfaces essentially glabrous; sori medial, indusia brown, ca. 1–1.5 mm diam., somewhat vaulted, glabrous; 2n⫽82 (USA).

Distribution. Terrestrial in oak woodlands, rocky banks and canyons, and in sheltered caves; 50–400 m. Can (Brit Col), USA (Ariz, Calif, Oreg, Wash); Mexico. Selected Specimens Examined. BCN (Harvey 534, DS, US; Hevly 2198, ARIZ; Moran 21716, MEXU; Pinkava 14294, NY, 14295, ARIZ, ASU; Thomas 117, DS; Thorne 62504, ENCB; Wiggins 4249, CAS, DS, LL, NY, UC, US; Wiggins & Gillespie 3908, CAS, DS, MEXU, NY, US; Wiggins & Thomas 347, DS, ENCB, NY).

Dryopteris arguta is a species of the western United States, barely ranging into northern Baja California Norte. Its closest affinities appear to be with D. pallida (Bory) C. Chr. ex Maire & Petitm., from Mediterranean Europe (see Fraser-Jenkins, 1986: 193). Of Mexican species, it is similar to D. filix-mas and differs from that primarily in the scalier stipes and rachises, less reduced proximal pinnae, and more strongly spinulose segments. There is also similarity to, and probably relationship with, D. maxonii, but the latter differs in having less scaly stipes and rachises, tan stipe scales, and ultimate segments not so distinctly toothed or spinulose at the tips. The combining authority for D. arguta is often given as Watt (Canad. Nat., II, 3: 159. 1866), but at this place Watt clearly chose to recognize Dryopteris as a subgenus (or subgroup) within Aspidium, and so Watt should not be credited with the species combination in Dryopteris.

DRYOPTERIS CINNAMOMEA

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2. DRYOPTERIS CINNAMOMEA Dryopteris cinnamomea (Cav.) C. Chr., Amer. Fern J. 1: 95. 1911. Figs. 122A–G, L, 123E, F. Tectaria cinnamomea Cav., Descr. Pl. 252. 1801. Type. Mexico. [Me´xico:] Chalma, Ne´e s.n. (MA, frag. S). Nephrodium mexicanum C. Presl, Reliq. Haenk. 1(1): 38. 1825. Dryopteris mexicana (C. Presl) C. Chr., Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk Math. Afd., ser. 7, 10: 16. 1913. Type. Mexico. Haenke s.n. (PRC; photo NY!). Aspidium athyrioides M. Martens & Galeotti, Me´m. Foug. Mexique 67, pl. 18. 1842. Dryopteris athyrioides (M. Martens & Galeotti) Kuntze, Revis. Gen. Pl. 2: 811. 1891. Type. Mexico. [Hidalgo:] “Real-del-Monte,” 8000–8500 ft, Galeotti 6425 (P!, photo US!, frag. NY!; isotype K!). Polystichum cystopteroides Nees, Linnaea 19: 685. 1847. Type. Mexico. Aschenborn 192 (B?). Polypodium glanduliferum Liebm., Mexic. Bregn. 206 (reprint 54). 1849. Dryopteris glandulifera (Liebm.) C. Chr., Index Filic. 267. 1905. Syntypes. Mexico. Oaxaca: Inter Comaltepec et Trapiche de la Concepcı´on, Liebmann s.n. [Pl. Mex. 2395, Fl. Mex. 196, 197] (C!; isotype K!). Lastrea indecora Liebm., Mexic. Bregn. 272 (reprint 120). 1849. Dryopteris indecora (Liebm.) C. Chr., Index Filic. 272. 1905. Type. Mexico. Oaxaca: Yavesia, Liebmann s.n. [Pl. Mex. 2417, Fl. Mex. 465] (C!, frag. US!). Aspidium agatolepis Fe´e, Me´m. Foug. 8: 106. 1857. Type. Mexico. S. Agostı´n, Schaffner 309b (RB, cited by Windisch, 1982: 57). Aspidium roseum E. Fourn., Mexic. Pl. 1: 97. 1872. Dryopteris rosea (E. Fourn.) Mickel & Beitel, Mem. New York Bot. Gard. 46: 165. 1988. Lectotype (chosen by Mickel & Beitel, 1988: 165). Mexico. [Veracruz:] Orizaba, Mu¨ller 4bis (P?; isolectotype NY!). Aspidium flaccidum E. Fourn., Bull. Soc. Bot. France 27: 328. 1880, hom. illeg., non Blume, 1828. Nephrodium fournieri Baker, Ann. Bot. (Oxford) 5: 317. 1891, non Baker, op. cit. 328. Dryopteris fournieri (Baker) C. Chr., Index Filic. 266. 1905. Type. Mexico. San Luis Potosı´: Schaffner 85 (P, photo NY!, US!, frag. BM, NY!).

Rhizomes short-creeping to suberect; rhizome scales pale brown to reddish brown, concolorous or with darkened bases at the point of attachment (rarely with a darker central stripe), often lustrous, mostly 10–18 ⫻ 1–3 mm, lanceolate to ovatelanceolate, sometimes with long twisted tips, entire to subentire; fronds 15–75 cm long; stipes stramineous to pinkish to reddish brown, 4–27 cm long, 1⁄4–1⁄2 the frond length, bases scaly, scales mostly 5–18 mm long, with acuminate tips, sometimes also a few scattered scales distally, often densely glandular throughout; blades herbaceous to thin-chartaceous, green to yellow-green, ovate to deltate-lanceolate, 2–3-pinnate-pinnatifid at bases, otherwise mostly 2-pinnate-pinnatifid, 14–30(–50) ⫻ (7–)9–20(–30) cm, with pinnae mostly alternate, ascending 45–60⬚ from the rachises, 5–20 pairs; rachises stramineous, scales absent or nearly so, densely glandular; proximal pinnae (2.5–)4–20 ⫻ (2–)4–10 cm, often slightly shorter than second pinna pair, equilateral or slightly more developed basiscopically than acroscopically; distal pinnae sessile or stalked to 4 mm, equilateral to more developed acroscopically, (3.5–)6–10(–18) ⫻ 2–7 cm, pinnules toothed; indument abaxially of stipitate glands on costae, costules, veins, and laminae between veins, scales none, adaxially the surfaces often stipitate-glandular, sometimes subglabrous; sori submarginal, indusia grayish to tan, 0.8–1.5 mm diam., flat or slightly vaulted, glandular.

Distribution. Terrestrial and epipetric (rocky ledges) in light shade, montane forests; 1000–3100 m. USA (Tex?); Mexico. Selected Specimens Examined. Ags (McVaugh & Koelz 214, CAS, IEB, MEXU, NY). Chih (Correll & Gentry 23002, LL, UC). Coah (Johnston et al. 11983B, LL p.p.). Col (Goldsmith 32, US). DF (Pringle 11773, ARIZ, LL, UC, US). Dgo (Breedlove 59129, CAS, NY). Gro (Lorea 3131, FCME, IEB, NY). Gto (Dı´az B. & Pe´rez 7385, IEB, UC). Hgo (Pringle 13815, ARIZ, ASU, CAS, LL, UC, US). Jal (Machuca N. 6368, IEB, UC). Me´x (Purpus 1606, NY, UC). Mich (Dı´az B. 7422, IEB, UC). Mor (Fisher 35423, MO, NY). Nay (Flores F. 2203, MEXU). Oax (Frame 309, NY, UC). Pue (Purpus 4022, ARIZ, UC, US). Qro (Rzedowski 44507, 51106, IEB, UC). SLP (Rzedowski 11387, NY). Tlax (Sharp & Herna´ndez X. 44467, US). Ver (Ventura A. 10749, ASU). Zac (Rzedowski & McVaugh 953, ASU, ENCB).

Dryopteris cinnamomea is extremely variable and widely distributed in Mexico, but unknown or at least uncertain beyond Mexico. It is distinguished from other members of the D. patula complex by its long (10–18 mm) rhizome apex and stipe base scales with entire or slightly denticulate margins and acute apices, usually smaller frond size, thin-chartaceous blades, stramineous or reddish brown stipes, and glandular laminar surfaces (both sides). However, a significant number of specimens, especially those from western Mexico (Oaxaca to Colima) are eglandular or sparsely glandular adaxially. Indusia are often distinctly grayish (but sometimes tan), and the blades are thinner and more membranaceous than other Mexican Dryopteris. Another rather similar species is D. knoblochii (q.v.). Most collections identified as D. cinnamomea from northwestern Mexico have proven to be either D. rossii, which differs in having smaller, black-tipped stipe base scales, or D. knoblochii, which has a thicker blade texture, strongly inequilateral distal pinnae, and smaller indusia. Occasional mixed collections of D. cinnamomea and D. rossii have been seen, e.g., Zu´n˜iga G. 18288 (UC), from Michoaca´n. Mickel and Beitel (1988) distinguished D. rosea from D. cinnamomea by the former having reddish brown stipes, smaller stipe base scales, and glabrous laminae adaxially. While these differences generally seem to hold, many specimens of D. cinnamomea, with glandular laminae adaxially, have reddish brown stipes typical of D. rosea, e.g., Cervantes C. 35 (IEB), from Edo. Me´xico, and Mickel 3865 (NY), from Oaxaca. Most specimens from Chiapas annotated by Mickel and Beitel as D. rosea, e.g.,

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DRYOPTERIS

Purpus 6725, 6725, Matuda 2452, all NY, are rather typical D. patula. Specimens from the Valle de Me´xico and Edo. Hidalgo are often larger and more finely dissected than specimens elsewhere in the range. The epithet “athyrioides” has aptly been applied to them. Some specimens from Guerrero have somewhat bicolorous stipe base scales, nearly black in a central band, lighter brown along the margins, and the scales are narrower than specimens from elsewhere. These are D. rosea, as the name was applied by Mickel and Beitel (1988). Putative hybrids involving D. cinnamomea and having malformed spores are the following: Rzedowski 45740 (ENCB), from Quere´tero; and King & Soderstrom 5202 (US), from Michoaca´n.

3. DRYOPTERIS FILIX-MAS Dryopteris filix-mas (L.) Schott, Gen. Fil., ad t. 9. 1834. Figs. 120G, H. Polypodium filix-mas L., Sp. Pl. 2: 1090. 1753. Lectotype (chosen by Jonsell & Jarvis, 1994: 149). Europe, Herb. Clifford 475, Polypodium 10 (BM).

Rhizomes suberect to erect; rhizome scales lacking in the sole Mexican specimen, but elsewhere concolorous, orange-tan, of two distinct kinds, one broad, ovate to lanceolate, mostly 5–15 ⫻ 2–6 mm, the other hair-like, mostly 4 ⫻ 0.1 mm, lustrous, irregularly toothed to subentire; fronds 50–80 cm long; stipes stramineous, ca. 15 cm ⫻ 4 mm (frond incomplete), less than 1⁄4 the frond length, with scales near bases and also some distally, 10–18 mm long, with acuminate tips, eglandular; blades herbaceous, green, lanceolate, 40–60 ⫻ 15–25 cm, 2-pinnate-pinnatifid in the proximal 2⁄3–3⁄4, pinnate-pinnatifid in the distal 1⁄3–1⁄4, with pinnae mostly alternate, spreading 75–90⬚ from rachises, 25–35 pairs; rachises stramineous, moderately scaly, eglandular; proximal pinnae 8 ⫻ 3.2 cm, shorter than second pinna pair, equilateral with pinnules about the same length on basiscopic and acroscopic sides, to 2 ⫻ 0.9 cm; distal pinnae sessile or stalked to 3 mm, equilateral, to ca. 13 ⫻ 4 cm, the 5th or 6th pinna pair the longest, pinnules serrate to lobed ca. halfway to costules; indument abaxially of scattered stramineous to light tan scales on costae and costules, glands absent or sparse, adaxially glabrous; sori medial, with tan indusia ca. 1 mm diam., flattish, eglandular; 2n⫽164 (USA).

Distribution. Terrestrial in pine-oak forests; 1500–2250 m. Canada, USA (including Tex, N Mex, Ariz, and Calif); Mexico; Greenland; Europe; Asia. Specimen Examined. Coah (Johnston et al. 11770, LL). Unverified, Doubtful, or Mistaken Reports. NL (Hinton & Hinton 22097, cited by Hinton & Hinton, 1995, but the specimen is D. pseudofilix-mas).

The sole Mexican collection of this species is from near the Texas border and the Big Bend region, Canyon Hundido on N side of Pico de Centinela, Sierra del Jardı´n, 8 km E of Rancho El Jardı´n, 29⬚06–08' N, 102⬚37–38' W. This specimen, which lacks stipe bases and rhizomes, is larger than many from the United States, but matches well the more dissected forms, e.g., Cusick 3250 (UC), from Oregon. Plants of D. filix-mas from the western United States may not be conspecific with those from eastern North America and elsewhere. Dryopteris filix-mas most resembles D. arguta, from which it differs in having the basal basiscopic pinnule of proximal pinnae slightly longer that the next pinnule (slightly shorter in D. arguta), less scaly stipes and rachises, the greater number of pairs of slightly reduced pinnae (the lowermost 8 cm, then the next 5 pairs gradually lengthened to 13.5 cm), and the less pronouncedly spinulose segment margins. From D. pseudofilix-mas it differs in having more dissected leaves (the middle pinnae being fully 2pinnate) and the lighter green blades. The entire D. filix-mas complex is in need of revision (Montgomery & Wagner in FNA Ed. Comm., 1993) throughout its range and especially in the New World.

4. DRYOPTERIS FUTURA Dryopteris futura A. R. Sm., Proc. Calif. Acad. Sci., ser. 4, 40: 216, f. 3F–I. 1975. Type. Mexico. Chiapas: Mpio. El Porvenir, 3–4 km W of El Porvenir along road from Huixtla to Siltepec, Breedlove 31772 (DS). Figs. 120E, F. Rhizomes erect; rhizome scales concolorous, brown to tan, dull to somewhat lustrous, 10–25 ⫻ 2–5 mm, ovate-lanceolate, margins entire or with minute glands; fronds 45–110 cm long; stipes stramineous to light brown at bases, 25–50(–70) cm ⫻ 4–5 mm,

DRYOPTERIS KARWINSKYANA 1

⁄2–2⁄3 the frond length, bases with numerous scales similar to those of rhizomes, glabrous distally; blades thin-herbaceous, green, broadly deltate, 3–4-pinnate-pinnatifid proximally, otherwise 2–3-pinnate-pinnatifid, 20–40 ⫻ 20–35 cm, with pinnae opposite (proximally) to subopposite or alternate (distally), ascending 45–70⬚ from rachises, 10–12 pairs; rachises stramineous, lacking scales, eglandular or sparsely glandular; proximal pinnae the longest, broadly deltate, strongly inequilateral, 13–25 ⫻ 8–17 cm, stalked 0.8–1.5 cm, pinnules to ca. 12 pairs per pinna, with larger basiscopic basal pinnule 6–12 cm long, acroscopic basal pinnule much shorter (10–15 mm long); distal pinnae sessile or stalked to 5 mm, slightly basiscopically developed or nearly equilateral, segment margins denticulate; indument abaxially absent or of sparse to moderate stipitate-glands 0.1 mm long on costae, costules, veins, and sometimes between veins, adaxially the blades glabrous; sori submarginal, indusia tan to red-brown, 0.6– 1 mm diam., flat, glabrous or margins stipitate-glandular; 2n⫽82 (Chis).

275

Rhizomes erect; rhizome scales concolorous, light brown, 7–11 0.5–1 mm, linear-lanceolate, margins with occasional teeth; fronds 20–60 cm long; stipes stramineous, 9–30 cm ⫻ 1.5–3(–4) mm, 1⁄4–1⁄2 the frond length, bases with light brown, linearlanceolate scales, densely glandular; blades chartaceous, green to yellow-green, deltate, 3-pinnate-pinnatifid at bases, otherwise mostly 2-pinnate-pinnatifid, 15–35 ⫻ 11–25 cm, with pinnae opposite to subopposite, spreading nearly 90⬚ from rachises, 12–20 pairs; rachises stramineous, with scattered scales 3–4 ⫻ 0.3–0.5 mm, also with dense glandular hairs; proximal pinnae the longest, 7.5–13 ⫻ 2.5–6 cm, inequilateral, basiscopic basal pinnules 25– 40 mm long, acroscopic basal pinnules shorter, 15–20 mm long; distal pinnae sessile or stalked to 3 mm, equilateral or slightly more developed acroscopically, pinnules acutely lobed or toothed; indument abaxially of dense whitish to yellowish, cylindrical glands 0.1–0.2 mm long on costae, veins, and laminae between veins, also of scattered hair-like scales 1–3 ⫻ 0.1–0.2 mm on costae, adaxially with similar dense glands; sori marginal, fertile zone limited to proximal and central part of fronds near pinna and pinnule bases, usually a single sorus per ultimate segment, indusia red-brown, thick, (1–)1.5–1.8 mm diam., vaulted, bearing dense, whitish, cylindrical glands; 2n⫽82 (Oax). ⫻

Distribution. Terrestrial in wet montane forests; (1750–)2500– 2800 m. Mexico; Guat. Selected Specimens Examined. Chis (Breedlove 32058, DS, FCME, 55907, 65730, CAS; Fraser-Jenkins 13184, UC; Mendoza-Ruiz 239, UAMIZ). Gro (Lorea 1638, FCME, IEB, NY, 2299, 4541, FCME, IEB; Martı´nez S. 1121, MEXU). Hgo (Lyonnet 510900007, MEXU).

Distribution. Terrestrial in oak woodlands, seasonal evergreen forests, especially along streams; 200–1300 m. Mexico; Guat, Hond, Salv, Nic, CR; Easter Island (Rodrı´guez R., 1995).

Dryopteris futura differs from congeners in Mexico by having broadly deltate, 3–4-pinnate-pinnatifid blades, with proximal pinnae ascending and much more strongly developed on the basiscopic side than the acroscopic side. The closest ally is D. nubigena. Both D. futura and D. nubigena are uncommon or at least poorly collected.

Selected Specimens Examined. Chis (Breedlove 40670, DS). Gro (Lorea 4895, FCME, IEB; Lyonnet 1722, CAS, US). Jal (Rzedowski & McVaugh 1372, ENCB). Me´x (Matuda 31464, US). Mich (Lorea 1727, IEB, FCME; Ventura 2496, ENCB, LL, NY). Nay (McVaugh 16549, MICH, TEX, US). Oax (Mickel 6936, NY, UC). Sin (Brandegee s.n., UC, US).

5. DRYOPTERIS KARWINSKYANA

Unverified, Doubtful, or Mistaken Reports. Dgo (Correll & Johnston 20134B, NY, cited by Mickel & Beitel, 1988, is D. cinnamomea). Tres Marias (reported by Lenz, 1995, but not verified).

Dryopteris karwinskyana (Mett.) Kuntze, Revis. Gen. Pl. 2: 813. 1891. Figs. 121C, D. Aspidium karwinskyanum Mett., Abh. Senckenberg. Naturf. Ges. 2: 343. 1858. Nephrodium karwinskyanum (Mett.) Baker in Hooker & Baker, Syn. Fil. 279. 1867. Type. Mexico. [Mexico “Cristo”:] Karwinski s.n. (B!).

This is a very distinctive species, not easily confused with any other species in the genus. The dense whitish or yellowish glandular hairs 0.1–0.2 mm everywhere on the laminae and indusia, large (1.5–1.8 mm), hemispherical, vaulted indusia, and the small ultimate and penultimate segments distinguish D. karwinskyana from members of D. patula complex. Dryopteris karwinskyana is

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DRYOPTERIS

restricted to the western states in Mexico, at relatively low elevations.

6. DRYOPTERIS KNOBLOCHII Dryopteris knoblochii A. R. Sm., sp. nov. Type. Mexico. Chihuahua: La Bufa, on Rı´o Batopilas, Knobloch 556 (holotype US). A D. rossii squamis pallide brunneolis concoloribus stipitum, squamis apicibus denigratis carentibus differt; a D. cinnamomea squamis paucioribus parvioribus obscure brunneolis stipitum, textura crassiore (venis non manifestis) laminarum, pinnis proximalibus inaequilateris, pinnis ascendentibus, indusiis parvioribus (0.5–0.8 mm diam.) caducioribus vel vietis ad maturitatem differt. (Dedicated to the memory of Irving Knobloch, 1907–1999, Professor of Botany at Michigan State University, student of the ferns of Chihuahua, and avid collector in that state.)

Rhizomes short-creeping to suberect; rhizome scales concolorous, light brown, dull, 2–5 ⫻ 1–2 mm, ovate-lanceolate, margins entire or minutely glandular; fronds (12–)25–50 cm long; stipes stramineous, (3–)10–20 cm ⫻ 0.8–2 mm, 1⁄4–1⁄3 the frond length, minutely stipitate-glandular, bases scaly, scales concolorous, mostly 2–5 ⫻ 1–2 mm, entire, tan to orange-brown; blades chartaceous, drying brownish green, deltate, 2-pinnate-pinnatifid to barely 3-pinnate at bases, mostly 2-pinnate-pinnatifid or bipinnate in the middle, 10–35 ⫻ 5–18 cm, with pinnae mostly alternate, ascending 55–70⬚ from rachises, 8–15 pairs; rachises stramineous, with sparse, pale scales, also densely glandular; proximal pinnae Ⳳ equilateral or often a little inequilateral, 2.5–12 ⫻ 1.4– 6.5 cm, with largest basiscopic basal pinnule to 3 cm long, largest acroscopic basal pinnule to 3.5 cm long; distal pinnae stalked up to 3 mm, equilateral or often slightly more developed acroscopically, pinnules and ultimate segments minutely toothed; indument abaxially of stipitate glands on costae, veins, and surfaces between veins, lacking scales or nearly so on costae, adaxially the laminae with rather dense, short-stipitate glands; sori supramedial to submarginal; indusia tan, 0.5–0.8 mm diam., flat, glabrous or glandular, caducous or shrivelled.

Distribution. Terrestrial in pine-oak forests; 1000–2550 m. USA (Ariz); Mexico.

Specimens Examined. Chih (Correll & Gentry 23020, LL, US; Knobloch 163, LL, US, 848, 854, US, 5569, LL, 5959, US, s.n., 6 Aug 1939, DS; McGill & Keil 8376, ASU; Nesom & Vorobik 5734h, TEX, US, 5623, 5728, TEX). Dgo (Breedlove 59072, CAS, NY). Rev (Patin˜o 26, MEXU). Sin (Breedlove 18321, 16935, CAS). Son (Boutin & Kimnach 3676, MEXU, NY; Fishbein 1746, 1849, ARIZ; Gentry et al. 19375, ARIZ, NY, US; Jenkins et al. s.n., ARIZ; Martin et al. s.n., ARIZ; Martin et al. s.n., ARIZ; Pennell 19658, US; Turner 79-374, ARIZ). USA (Lemmon s.n., CAS, DS).

Specimens of this have been determined previously as D. rossii (or Dryopteris patula var. rossii) or D. cinnamomea. From the former, D. knoblochii differs primarily in having tan or light brown, concolorous stipe base scales lacking blackened tips. From D. cinnamomea, it differs in the fewer, smaller, less orangish, dull stipe base scales; thicker, more opaque blade texture with veins not so readily visible; inequilateral, decidedly anadromous proximal pinnae (vs. equilateral and isodromous or nearly so); pinnae ascending (vs. spreading); and by the relatively small (0.5–0.8 mm diam.) indusia that tend to be more caducous or at least greatly shrivelled at maturity. Arizona specimens referred to Dryopteris cinnamomea by Montgomery and Wagner (FNA Ed. Comm., 1993) are a better match for D. knoblochii. A specimen from Chihuahua (12 mi W of Cuauhte´moc, Gould 8952, UC) has strongly malformed spores and may represent a hybrid between D. cinnamomea and D. knoblochii.

7. DRYOPTERIS MAXONII Dryopteris maxonii Underw. & C. Chr., Amer. Fern J. 1: 96. 1911. Type. Mexico. Morelos: Above Cuernavaca, Pringle 6190 (US!; isotypes CAS!, NY!, UC!, US!). Figs. 119A–C. Dryopteris patula (Sw.) Underw. var. moreliae Christ in Lecomte, Notul. Syst. (Paris) 1: 234. 1910. Type. Mexico. Michoaca´n: vicinity of Morelia, Punguato, 2100 m, Arse`ne 2861 (P?; isotypes L, photo US!, US!).

Rhizomes suberect; rhizome scales bicolorous, light brown with dark brown to black central streak, 5–8 ⫻ 2–3 mm, ovatelanceolate, entire; fronds (25–)40–80 cm long; stipes stramineous, 12–32 cm ⫻ (1–)2–5 mm, 1⁄4–1⁄2 the frond length, bases with entire scales, lacking scales distally, eglandular; blades herbaceous to chartaceous, green, ovate-lanceolate to deltate, generally 2pinnate-pinnatifid, 19–40(–60) ⫻ (4–)8–26 cm, with pinnae mostly alternate, spreading or only slightly ascending, 13–20 pairs; rachises stramineous, with scattered, tan lanceolate twisted scales 1–3 ⫻ 0.1–0.4 mm, scales irregular toothed or ciliate, glands absent; proximal pinnae inequilateral, more developed basiscopically, (4–)6–25 ⫻ (1.7–)2.5–10 cm, slightly shorter than or nearly equalling the second pinna pair, basiscopic basal pinnule to 4.5 cm long, shorter that the next pinnule (to 6 cm), acroscopic basal pinnule to 3.5 cm long; distal pinnae sessile or stalked to 5 mm, Ⳳ equilateral, pinnules deeply lobed to toothed; indument abaxially of scattered, tan scales to 3 ⫻ 0.4 mm on costae and costules, glands essentially absent, adaxially the blades glabrous; sori medial, indusia brown, 1.2–1.5 mm diam., vaulted, glabrous; 2n⫽82 (Jal).

DRYOPTERIS NUBIGENA

277

shortened, deltate-lanceolate, to 15 ⫻ 8.5 cm, stalked to 6 mm; distal pinnae sessile or stalked to 1 mm, Ⳳ equilateral, pinnules to ca. 10 pairs per pinna, spinulose at apices; indument abaxially of minute, tan, ovate to linear scales along costae and costules, otherwise glabrous, eglandular, adaxially the blades glabrous; sori medial, indusia reddish brown, ca. 0.8–1 mm diam., glabrous.

Distribution. Shaded banks and rocks in humid or moderately dry oak or pine-oak forests, or tropical deciduous forests; (300–) 1100–2300 m. Mexico; reported from Guatemala, El Salvador, and Costa Rica by Moran (in Davidse et al., 1995), but the identity of these specimens is in doubt and the provenance of the specimen from Costa Rica is also suspect. Selected Specimens Examined. Col (Orcutt 4700, DS, NY, TEX, US). DF (Meave 478, MEXU). Dgo (Ortega 4289, US). Gro (Hinton 9445, NY, TEX, US). Gto (Rzedowski 51756, IEB). Jal (Cutler 4073, UC). Me´x (Hinton 4703, NY). Mich (Pringle 8846, CAS, TEX, UC, US). Mor (Pringle 15718, ARIZ, ASU, CAS, LL, TEX, UC, US). Nay (Solo´rzano Vargas 16916, UC). Pue (Nicolas s.n., UC). Sin (Breedlove 18135, CAS, NY).

Dryopteris maxonii differs from congeners in Mexico by its vaulted, firm, glabrous indusia, essentially glandless blades, and stramineous stipe bases with light tan scales. Hinton 7316 (LL, TEX, US), 16040 (NY, UC, US), and 16155 (ENCB, LL, UC, US) were all collected in Edo. Michoaca´n, Dto. Coalcoman, Aquila, 300 m. The first of these three collections was determined as “D. micradenia Weath., n. sp. ined.,” at GH. These collections differ from other specimens of the species by their smaller size and more dentate pinnules, and were collected at much lower elevation than usual for the species.

8. DRYOPTERIS MUENCHII Dryopteris muenchii A. R. Sm., Proc. Calif. Acad. Sci., ser. 4, 40: 218, f. 3A–B. 1975 [“munchii”]. Type. Mexico. Chiapas: Baduitz, Mu¨nch 116 (DS!; isotype US!). Figs. 120A, B. Rhizomes erect to suberect; rhizome scales concolorous, dark brown, to 10 ⫻ 2 mm, entire to toothed; fronds to ca. 90 cm long; stipes brownish, to 35 cm ⫻ 5 mm, 1⁄3–1⁄2 the frond length, bases scaly, the scales often bicolorous with blackened or darkstreaked bases, apices tan, margins dentate, stipes eglandular; blades herbaceous, dark green adaxially, lighter green abaxially, ovate-lanceolate, subtripinnate at bases, distally 2-pinnatepinnatifid, to 50 ⫻ 20–28 cm, with pinnae mostly alternate, spreading to slightly ascending, 15–18 pairs; rachises tan, bearing scattered to rarely numerous scales, eglandular; proximal pinnae subopposite, basiscopically more developed, slightly or not at all

Distribution. Terrestrial in cloud forests; 2000–2850 m. Mexico. Specimens Examined. Chis (Breedlove 22027, DS, MEXU, NY; Gonza´lez-Espinosa et al. 583, CHAPA; Mendoza-Ruiz 244, UAMIZ; Mu¨nch 167, DS). Hgo (Lyonnet 2007, US; Pin˜a E. 47, ASU; Sa´nchez 375, US).

From D. rossii, D. muenchii differs by its narrower, eglandular, dark green blades and stipe base scales that are blackened or black-streaked at bases (rather than blackish at tips). From D. patula and D. cinnamomea, it differs in the smaller, bicolorous stipe scales, spreading, nearly equilateral, suprabasal pinnae, and eglandular blades. The nearest affinities are uncertain.

9. DRYOPTERIS NUBIGENA Dryopteris nubigena Maxon & C. V. Morton, Proc. Biol. Soc. Wash. 50: 179. 1937. Type. Guatemala. Chimaltenango: Buena Vista above Tecpa´n, Skutch 771 (US!; isotypes NY!, US!). Figs. 122K, 123G, H. Rhizomes erect; rhizome scales concolorous, castaneous to dark brown, 5–10 ⫻ 1.5–2 mm, ovate-lanceolate, margins entire or with minute glands; fronds 33–85 cm long; stipes usually dark brown to atropurpureous at bases, stramineous to light brown distally, 20–40 cm ⫻ 1–3 mm, ca. 1⁄2 or slightly more of the frond length, bases usually with numerous scales similar to those of rhizomes, glabrous distally; blades herbaceous, dark green, broadly deltate, 3-pinnate-pinnatifid proximally, 2-pinnatepinnatifid to 3-pinnate distally, 12–42 ⫻ 9–44 cm, with pinnae opposite to subopposite throughout, spreading to slightly ascending ca. 75–90⬚ from rachises, 8–12 pairs; rachises stramineous, lacking scales, minutely stipitate-glandular or eglandular; proximal pinnae the longest, strongly inequilateral, 5–22 ⫻ 2.5– 13 cm, stalked to 1.5 cm, basiscopic pinnules more strongly devel-

278

DRYOPTERIS

oped, 1.5–7 cm long, acroscopic pinnules shorter, 1–5 cm long; distal pinnae sessile or stalked to 5 mm, Ⳳ equilateral, pinnules to 15 pairs per pinna, margins toothed; indument abaxially lacking or of sparse catenate hairs 0.1–0.3 mm long on costae and costules, also with moderate to dense yellowish short-stipitate glands 0.1 mm on costae, costules, veins, and surfaces between veins, adaxially the surfaces glabrous or often with short-stipitate glands; sori submarginal, indusia tan, 0.8–1 mm diam., flat, stipitate-glandular.

Distribution. Terrestrial pine-oak forests, cloud forests; 2150– 3000 m. Mexico; Guat, Hond, CR. Moran (in Davidse et al., 1995) also cited a collection from Panama; the collection he cited from Honduras (Weatherwax 273, MO!) is actually from Guatemala, but the species does indeed occur in Honduras (Moran 5597, MO!). Selected Specimens Examined. Chis (Breedlove 22069, DS, NY, 22721, DS, MEXU, NY, 22818, 40334, 41739, DS, 55850, CAS, ENCB, NY; Fraser-Jenkins 13119, UC). ?Hgo (Sa´nchez Mejorada 757, MEXU). Me´x (Hinton 3447, K, NY, 3469, BM, K, US). Oax (Barrington 1734, UAMIZ; Mickel 4056, 4136, NY, 5285, ENCB, NY, UC, very atypical).

The glabrous stipes with darkened bases, deltate proximal pinnae much more developed basiscopically, and appressed septate hairs characterize this species. Some specimens from Oaxaca and Chiapas are smaller than the type, but otherwise very similar. The specimen cited from Hidalgo (Plan del Ayacahuite, El Chico) is incomplete, but if it is not this species then it is possibly undescribed. Citation of specimens from Guerrero (LoreaHerna´ndez & Vela´zquez M., 1998) is the result of confusion with D. futura, which see for comparison. Mickel 4056, 4136, and 5285 are very atypical (unusually small, and so less dissected) for this species. In addition to their small size, they differ in having stramineous stipes, lighter stipe base scales, medial sori, and sparsely glandular blades. The description of D. nubigena above does not take into consideration the variation in these collections.

10. DRYOPTERIS PATULA Dryopteris patula (Sw.) Underw., Native Ferns, ed. 4, 117. 1893. Figs. 121G, H, 122H, J.

Aspidium patulum Sw., Kongl. Vetensk. Acad. Handl. 1817: 64. 1817. Type. Brazil. Minas Gerais: Freyreiss s.n. (S). Aspidium paupertinum Kunze, Linnaea 18: 345. 1844. Aspidium mexicanum C. Presl var. obtusilobum Kunze ex Mett., Abh. Senckenberg. Naturf. Ges. 2: 349. 1858. Type. Mexico? “Reg. temp. (Herb. Roem., propr) Coll. No. 42.” Aspidium apertum Fe´e, Me´m. Foug. 8: 106. 1857. Dryopteris aperta (Fe´e) C. Chr., Index Filic. 252. 1905. Syntypes. Mexico. [Veracruz:] Huatusco, Schaffner 73 (P?); [Morelos:] Cuernavaca, Craveri 73b (P!). Dryopteris simplicior Mickel & Beitel, Mem. New York Bot. Gard. 46: 166. 1988. Type. Mexico. Oaxaca: Distrito Putla, 3 km N of Putla, Mickel 3977 (NY!).

Rhizomes erect; rhizome scales concolorous, light brown to often reddish tan, lustrous, (9–)12–25 ⫻ (1.5–)2–3 mm, ovatelanceolate, strongly toothed and twisted; fronds (25–)35–135 cm long; stipes stramineous to tan, 18–35 cm ⫻ 2–9 mm, 1⁄4–1⁄3 the frond length, scaly near bases with twisted scales to 20 mm, margins toothed, stipes also usually glandular; blades chartaceous, green to yellow-green, deltate-lanceolate, 3-pinnatepinnatifid at bases, otherwise mostly 2-pinnate-pinnatifid, (16– )25–50(–70) ⫻ (6.5–)20–40 cm, with pinnae mostly alternate, ascending 45–60⬚ from rachises, 12–20 pairs; rachises stramineous, scaleless or proximally with scattered, tan, linear, twisted scales to 5 ⫻ 0.2–0.4 mm, also with minute stipitate glands ⬍ 0.1 mm; proximal pinnae Ⳳ equilateral to slightly inequilateral, (4–)9–20(–25) ⫻ (2–)5–8 cm, shorter than or about equalling the second pinna pair, basiscopic basal pinnule 2–8 cm long, acroscopic basal pinnule 1.8–5.5 cm long; distal pinnae sessile or stalked to 5 mm, subequilateral or usually inequilateral and more developed acroscopically, pinnules of basiscopic side shorter and more oblique, 7–20 ⫻ 4–8 cm, pinnules pinnatifid to merely shallowly lobed to toothed; indument abaxially of few to many yellowish stipitate glands 0.1 mm long on costae, veins, and sometimes laminar tissue, scales none, adaxially with glands in costal grooves and also sometimes on surfaces; sori medial to submarginal, indusia brown to tan, 0.8–1.5 mm diam., flat, glabrous or glandular; 2n⫽82 (Jam).

Distribution. Terrestrial (rarely epiphytic on old logs) in montane rain forests, seasonal forests, pine-oak forests; (200–)1300– 2500 m. Mexico; Guat, Hond, Salv, Nic, CR, Pan; Jam, Hisp; Col, Ven, Ec, Galapagos, Peru, Braz, Bol, nw Arg. Moran (in Davidse et al., 1995) also included Paraguay in the range.

DRYOPTERIS PSEUDOFILIX-MAS Selected Specimens Examined. Chis (Matuda 2452, K, MEXU, NY; Purpus 6725, NY, UC, US). Hgo (Moore 5362, US). Mich (Arse`ne s.n., 24 Nov 1910, US). Mor (syntype of Aspidium apertum). Oax (Mickel 4862, NY; Solano C. 501, CAS, MEXU). Pue (Orcutt 3995, DS, NY, TEX). Tam (Lof 716, UC). Ver (C. Smith 2223, UC; Purpus 6446, US). Unverified, Doubtful, or Mistaken Reports. DF (reported by Mickel & Beitel, 1988, but not verified). Me´x (reported by Mickel & Beitel, 1988, but not verified; the chromosome voucher, Mickel 668, ISC, cited by Mickel et al., 1966, as this species is D. rossii; Tejero Dı´ez 3002, IZTA, cited by Tejero-Dı´ez & Arreguı´n-Sa´nchez, 2004, is D. cinnamomea). Sin (reported by Vega A. et al., 1989, probably misidentified).

The D. patula complex is confused and in need of monographic study. Within this complex in Mexico we recognize four taxa at species rank (D. patula s.str., D. cinnamomea, D. knoblochii, and D. rossii). In the restricted sense, D. patula has deltatelanceolate fronds, long strongly toothed rhizome scales that are marginally crisped and have long, twisted hair-tips, and usually glabrous blade surfaces (except for costal and costular grooves adaxially and sparse glands on the veins abaxially); however, more glandular variants are known, especially along the Pacific slopes of the Sierra Madre Occidental. Dryopteris flaccisquama A. Rojas, recently described from Costa Rica to Bolivia, and Haiti, is merely a very large form of D. patula, and probably not worthy of species distinction. Dryopteris simplicior was said to differ from D. patula primarily in being glandular on the adaxial surfaces of the laminae (vs. glands along only the midribs), pale brown rhizome scales (vs. reddish brown), and proximal pinna pair equalling or only slightly shorter than the second pinna pair. There is great variation in all these characters when specimens from throughout the range of D. patula are examined, and we are unable to separate the two satisfactorily. One of the collections cited as D. simplicior by Mickel and Beitel (1988) has irregularly shaped malformed spores (Mickel 6093, NY). A collection from Zacuapan, Veracruz (Purpus 3007, UC, US), with scales persistent along stipes and onto proximal part of the rachis, was annotated by Fraser-Jenkins, “. . . a distinct species . . . related to D. patula, but more dissected and more glandular on axes below. Seems to compare with var. serrata and chaerophylloides, but I am looking into its correct name—15 Nov 92.” Purpus 7868 (US) and Purpus 7879 (UC), also from Zacuapan, are similar but larger versions of this entity. Until the Dryopteris patula complex can be studied cytologically and isozymically, it seems premature to describe this variant. The sole collection from Tamaulipas is extremely aberrant in having very narrow, almost hair-like, reddish brown rhizome and stipe base scales (to ca. 10 ⫻ 0.3–0.5 mm) and small fronds (ca. 25 cm) and blades (16 ⫻ 6.5 cm), with pinnae to 4 ⫻ 2 cm; the blades are eglandular or nearly so on both sides, and the indusia are reddish brown. In some respects, it resembles certain specimens from Jamaica, e.g., Jenman s.n. (NY). It may represent an undescribed species in the Dryopteris patula complex, but more material and further study are needed before further subdivision of this species can be attempted. Perhaps the most aberrant element that we include in D.

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patula is a specimen from Chiapas, Breedlove 53929 (CAS), from a sandstone outcrop at Escopetazo (Mpio. Ixtapa). It has broadly deltate blades as broad as or slightly broader than long. Clearly it belongs in D. patula s.l., rather than being assignable to one of the other species with deltate blades (e.g., D. nubigena, D. futura, D. rossii).

11. DRYOPTERIS PSEUDOFILIX-MAS Dryopteris pseudofilix-mas (Fe´e) Rothm., Candollea 10: 96. 1945. Figs. 121E, F. Aspidium pseudofilix-mas Fe´e, Me´m. Foug. 8: 103. 1857. Type. Mexico. “Valle´e du Mexique,” Schaffner 253 (P!; isotype K!).

Rhizomes erect; rhizome scales concolorous, or often dark brown or blackish at the bases near point of attachment, light to medium brown or tan distally, to 20 ⫻ 1–5 mm, ovatelanceolate, margins irregularly ciliate to lacerate; fronds 40–100 cm long; stipes stramineous, 8–20(–25) cm ⫻ 4–8 mm, 1⁄8–1⁄4 the frond length, densely (at base) to moderately (distally) clothed in scales similar to those of rhizomes, eglandular; blades herbaceous, dark green to yellow-green, ovate-lanceolate, 2-pinnatepinnatifid at bases, pinnate-pinnatisect distally, 40–80 ⫻ 14–25 cm, with pinnae mostly alternate, spreading to slightly ascending, 20–36 pairs; rachises stramineous, usually moderately and persistently scaly throughout (larger scales denticulate to strongly toothed), eglandular; proximal pinnae inequilateral, somewhat basiscopically developed, 7–10 ⫻ 2.5–4 cm, slightly shorter than more distal pinnae, short-stalked to 3 mm, largest basiscopic pinnules to 2.5 ⫻ 0.8 cm, pinnatifid; distal pinnae sessile or very short-stalked to 2 mm, equilateral, 5–15 cm long, ultimate segments tapered toward the rounded tips, denticulate; indument abaxially of moderately dense, persistent, fibrillose, often twisted scales to ca. 3 ⫻ 0.4 mm on costae, costules, veins, and tissue between veins, the largest scales strongly toothed, adaxially the lamina with similar but fewer and generally narrower scales; sori medial, indusia tan to brown, ca. 1–1.2 mm diam., slightly vaulted or flat, glabrous; 2n⫽123 (Oax).

Distribution. Terrestrial in pine-oak cloud forests; 2200–3950 m. Mexico; Guat (Molina et al. 16642, NY).

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DRYOPTERIS

Selected Specimens Examined. Chis (Breedlove 24320, DS, atypical). DF (Rzedowski 37660, NY, TEX). Hgo (Pin˜a E. 47, ASU). Me´x (Rzedowski 36088, MEXU, NY, TEX). Mich (Rzedowski 46172, ENCB, IEB). Mor (Sandoval 59, 139, MEXU). NL (Hinton & Hinton 22097, TEX). Oax (Smith 2076, NY, UC, US). Qro (Aguilar H. 55, IEB, MEXU, TEX). Pue (Arse`ne 2181, US p.p.). Ver (Zamora C. et al. 3835, UC).

This species appears to be related to “Dryopteris filix-mas” of the western United States, but it is not the same as eastern North American plants of that name. The relationships remain unresolved. In general, D. pseudofilix-mas differs from D. wallichiana in having fewer and shorter scales on the stipes and rachises and the scales are tan to brown, or with a brownish or blackish base, rather than dark brown or blackish throughout. In addition, the segments of D. pseudofilix-mas are rounded and more tapered at tips rather than blunt and parallel-sided, as in typical D. wallichiana. Probable hybrids involving this species are Huerta H. s.n. (XAL) and Vega Avin˜a 419 (NY), from Edo. Mexico. The spores are mostly highly irregular, but some are seemingly well formed and very large. Similar to this, and with many irregularly formed and small spores is Rzedowski 37660 (NY), from Distrito Federal, Monte Alegre, Serranı´a del Ajusco. The second parent may be D. wallichiana. Rothmaler (Candollea 10: 95–96. 1945) considered D. chrysocarpa (Fe´e) Rothm. (⬅ Aspidium chrysocarpon Fe´e, Me´m. Foug. 8: 103. 1857. Type. Mexico, Popocate´petl, Schaffner 289, P) as the hybrid between D. pseudofilix-mas and D. parallelogramma [⫽ D. wallichiana]. The putative hybrids appear intermediate in various characteristics between the two species and also appear to have malformed or highly irregular spores. The sole chromosome count for the species is indicative of an apogamous life cycle. Specimens of D. pseudofilix-mas from Nuevo Leo´n are all from Mpio. Zaragoza and do not match very well those from the Valle de Me´xico; Hinton 22097 is especially atypical in its less dissected pinnae. These specimens were determined as D. filixmas (see Aguirre-Clavera´n & Arreguı´n-Sa´nchez, 1988; Hinton & Hinton, 1995), but the blades are not as dissected as the sole Mexican specimen of D. filix-mas from near the Texas border. The sole collection from Hidalgo has about three pairs of fully pinnate pinnae, and the pinnules are rather deeply lobed or pinnatifid. The dark scales indicate its affinities are with D. pseudofilix-mas, rather than with D. filix-mas, but it is atypical. The only collection from Chiapas, from the summit of Volca´n Tacana´, is sterile and peculiar in having broad, dentate segments. It is also peculiar in the darkened intervenal tissue, which gives the dried fronds the appearance of being soriate between the veins. This condition is also seen in sterile fronds of Rzedowski 32557 (IEB), from Edo. Me´xico. The cause of this darkening, whether a necrosis or an infection, is unknown.

12. DRYOPTERIS ROSSII Dryopteris rossii C. Chr. in Ross, Mem. Acad. Nac. Ci. “Antonio Alzate” 32: 178. 1912. Figs. 123A–D.

Dryopteris patula (Sw.) Underw. var. rossii (C. Chr.) C. Chr., Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 7, 10: 20. 1913. Lectotype (chosen by Christensen, 1913: 22). Mexico. Morelos: Cuernavaca, Santa Marı´a, Ross 279 (M; isolectotype P!).

Rhizomes short-creeping; rhizome scales concolorous, blackish brown, lustrous, 2.5–5 ⫻ 1–2 mm, ovate, margins entire or with minute glands; fronds 20–80 cm long; stipes stramineous to castaneous proximally, 10–33 cm ⫻ 1–3(–4) mm, 1⁄3–1⁄2 the frond length, densely glandular, bases scaly, with entire scales 5–6(–10) ⫻ 1–2 mm, at least some scales bicolorous with lustrous dark tips and pale brown bases; blades chartaceous, green, deltate, 2–3pinnate-pinnatifid at bases, otherwise mostly 2-pinnatepinnatifid, 25–44 ⫻ (8–)18–30 cm, with pinnae mostly alternate, ascending 50–65⬚ from rachises, 10–15 pairs; rachises stramineous, with sparse, pale scales, also densely glandular; proximal pinnae Ⳳ equilateral to usually a little inequilateral, 8–20(–25) ⫻ 4–7(– 10) cm, with larger basiscopic basal pinnule 3–5.5 cm long, acroscopic basal pinnule 2–3(–5) cm long; distal pinnae stalked to 8 mm, equilateral or often slightly more developed acroscopically, pinnules and ultimate segments toothed; indument abaxially of stipitate glands on costae, veins, and surfaces between veins, also with sparse, hair-like scales 0.5–1 mm long on costae, adaxially glabrous or the lamina quite glandular; sori submarginal, indusia brown to tan, 0.5–0.8(–1) mm diam., flat, glandular; 2n⫽82 (Edo. Me´x., reported as D. patula by Mickel et al., 1966).

Distribution. Terrestrial in pine-oak woodlands, especially along streams; (500–)900–2600 m. Mexico; also cited from Nicaragua and Costa Rica by Moran (in Davidse et al., 1995), but the provenance and/or identity of these collections is suspect. Selected Specimens Examined. Ags (McVaugh & Koelz 214, MICH). BCS (Carter 2368 et al., UC). Col (Alava & Cook 1709, UC). Dgo (Sa´nchez 743, MEXU). Gro (Hinton 9344, LL, NY, US). Gto (Flores 135, IEB, MEXU). Jal (Pringle 1839, UC). Me´x (Bartholomew 2905 et al., CAS, UC). Mich (Pringle 3362, CAS, MEXU, NY, UC, US). Mor (Pringle 15710, ASU, CAS, LL, MO, UC, US). Nay (Alava & Cook 1624, UC). Oax (Mickel 6973, NY). Sin (Mexia 463, UC). Zac (Calzada 13398, UC). Unverified, Doubtful, or Mistaken Reports. DF (reported by Mickel & Beitel, 1988, but not verified). Qro (Carranza 1275, IEB,

DRYOPTERIS WALLICHIANA

281

QMEX, cited by Dı´az-Barriga & Palacios-Rios, 1992, and by Arreguı´n et al., 2001, but this specimen is D. cinnamomea).

This distinctive species in the D. patula complex is common in western and central Mexico, but rare or absent elsewhere. It can be distinguished by its broadly deltate blades, generally longer-stalked pinnae, dark brown or blackish, lustrous, relatively small rhizome scales, and strongly bicolorous stipe scales with tan bases and black tips. The closest relative is D. knoblochii, which differs in the concolorous tan stipe base scales.

13. DRYOPTERIS WALLICHIANA Dryopteris wallichiana (Spreng.) Hyl., Bot. Not. 1953: 352. 1953. Figs. 121A, B. Aspidium paleaceum D. Don, Prod. Flor. Nepal. 4. 1825, hom. illeg., non Sw., 1806. Aspidium wallichianum Spreng., Syst. Veg., ed 16, 4: 104. 1827. Aspidium donianum Spreng., Syst. Veg., ed. 16, 4: 320. 1827, nom. superfl. Dryopteris paleacea Hand.-Mazz., Verh. Zool.-Bot. Ges. Wien 58: 100. 1908, nom. superfl. Type. Nepal. Wallich s.n. (K). Aspidium paleaceum Sw., Syn. Fil. 52. 1806. Dryopteris paleacea (Sw.) C. Chr., Amer. Fern J. 1: 94. 1911, hom. illeg., non Hand.-Mazz., 1908. Type. Peru. Lasgasca s.n. (S–photos GH, US). Aspidium parallelogrammum Kunze, Linnaea 13: 146. 1839. Dryopteris parallelogramma (Kunze) Alston, Amer. Fern J. 47: 92. 1957. Syntypes. Mexico. Hegewisch s.n., Karwinski s.n. (B?). Aspidium crinitum M. Martens & Galeotti, Me´m. Foug. Mexique 66, pl. 17, f. 2. 1842. Type. Mexico. Oaxaca: Llano Verde, Galeotti 6348 (BR!, photos LL!, NY!, US!; isotypes B!, BR!–6 sheets, K!, P!–4 sheets photo US, US!, frag. NY!). For discussion of nomenclature (species epithet and authorship), see Smith and Fraser-Jenkins (1982).

Rhizomes erect, caudex often trunk-like, to 30 cm tall; rhizome scales concolorous, dark brown, to 25 ⫻ 3 mm, linearlanceolate, filiform at tips, with irregularly ciliate to subentire margins; fronds 45–135 cm long; stipes stramineous, (8–)15–30 cm ⫻ 4–12 mm, usually 1⁄4(–1⁄3) or less the frond length, at bases and also distally with dense, spreading, dark brown scales similar to those of rhizomes, the scales nearly concealing the stipes, eglandular; blades thick-herbaceous to subcoriaceous, lustrous dark green above, lighter below, oblong-lanceolate, pinnate-pinnatifid nearly throughout or infrequently fully 2pinnate proximally, (30–)50–100(–120) ⫻ (12–)18–28 cm, with pinnae mostly alternate, spreading nearly 90⬚, 22–40 pairs; rachises stramineous to light tan, copiously scaly (scales similar to those of stipes and often nearly obscuring rachises), eglandular; proximal pinnae linear-lanceolate, sessile or very shortstalked ⬍ 2 mm, reduced, 6–12 cm ⫻ 1.3–2.5 cm, equilateral, without enlarged basiscopic or acroscopic pinnules; distal pinnae sessile or stalked to 2 mm, equilateral, 7–18 ⫻ 1.5–2.5(–3) cm, segments oblong, rounded to often truncate at tips, entire to often denticulate at tips; indument abaxially of numerous linear to lanceolate brownish ciliate scales on costae and costules, veins and surfaces between veins glabrous, adaxially with similar scales on costae, lamina otherwise glabrous; sori medial, indusia brown, 0.8–1.4 mm diam., flat, glabrous; 2n⫽123 (Me´x), 164 (Oax, Jam).

Distribution. Terrestrial in cloud forests, pine-oak forests, especially along streams; 1500–3150 m. Mexico; Guat, Hond, CR, Pan; Jam, Hisp; Col, Ven, Ec, Peru, s Braz, Bol, nw Arg; Africa, Himalayas, India, China, Japan, Malesia, Hawaii. Moran (in Davidse et al., 1995) also included Paraguay in the range. Selected Specimens Examined. Chih (Correll & Gentry 23089, LL, US). Chis (Croat 40744, UC). DF (Ventura 1099, IEB, NY, TEX, XAL). Gro (Hinton 15709, NY, but blade more dissected than usual; Mexia 9070, ARIZ, CAS, NY, UC, US). Gto (Carranza et al. 4365, CAS, IEB). Hgo (Pringle 8750, CAS, LL, NY, UC, US). Jal (Iltis 2389 et al., NY, UC). Me´x (Molseed 544, UC). Mich (Hinton 15709, ARIZ, NY, TEX, US). Mor (Lyonnet 1418, MEXU, NY). Oax (Croat & Hannon 65640, UC). Pue (Pringle 15704, ARIZ, CAS, LL, UC, US). Qro (Ferna´ndez N. 3372, ENCB, IEB, NY, TEX). Tam (Sharp et al. 52225, US). Tlax (Reeves R5941, ASU). Ver (Storer 143, UC, US).

Dryopteris wallichiana appears to be an apogamous triploid or tetraploid in the New World, but an apogamous diploid in the Old World (Fraser-Jenkins, 1989). However, Asian and African material assigned to D. wallichiana is very similar to neotropical specimens. Dryopteris wallichiana has been confused with D. pseudofilix-mas (q.v. for differences); the former is much more common, and the two species are known to hybridize.

Unplaced Names Aspidium leptorhachis Kunze, Linnaea 18: 346. 1844. Type. Provenance not stated, perhaps Mexico, “Reg. temper. (Herb. Roemer, propr.).” “Evidently a less cut form of the same [D. patula]” (Christensen, 1913: 71). Aspidium mexicanum C. Presl var. acutilobum Mett., Abh. Senckenberg. Naturf. Ges. 2: 349. 1858. Type. Mexico. s. coll. s.n. (B?). “Adn. Polypodium angustifrons Kz. Linn. 13. 134 ex specimine incompleto hujus loci an species affinis.” Polypodium angustifrons Kunze, Linnaea 13: 134. 1839. Type. Mexico. Prope los Trojas, Octobri, Schiede 758a (LZ destroyed; isotype B?) “According to Mett. Aspid. 65 [Abh. Senckenberg. Naturf. Ges. 2: 349. 1858] this falls under my D. patula but may be an allied species, if really an Eudryopteris” (Christensen, 1920: 125).

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ELAPHOGLOSSUM

4 4 . EL A PH O G L O S S UM Elaphoglossum Schott ex J. Sm., J. Bot. (Hooker) 4: 148. 1841, nom. cons.. Type: Elaphoglossum conforme Schott ex J. Sm. Aconiopteris C. Presl, Suppl. Tent. Pterid. 236. 1836. Type: Aconiopteris subdiaphana (Hook. & Grev.) C. Presl [⬅ Acrostichum diaphanum Hook. & Grev.] ⫽ Elaphoglossum nervosum (Bory) Christ. Peltapteris Link, Fil. Spec. 147. 1841. Type: Peltapteris peltata (Sw.) C.V. Morton [⬅ Osmunda peltata Sw.] ⬅ Elaphoglossum peltatum (Sw.) Urb. Rhipidopteris Schott ex Fe´e, Me´m. Foug. 2: 14. 1845, hom. illeg., superfl. for Peltapteris peltata, and with the same type. Hymenodium Fe´e, Me´m. Foug. 2: 20, 80. 1845. Type: Hymenodium crinitum (L.) Fe´e [⬅ Acrostichum crinitum L.] ⬅ Elaphoglossum crinitum (L.) Christ.

Generally epiphytic in wet montane forests and evergreen cloud forests, a greater percentage terrestrial at high elevations; rhizomes short- to long-creeping, rarely erect, slender to stout (1–15 mm diam.); rhizome scales orange to black, basally attached or peltate, entire to toothed; fronds 2–200 cm long, erect, arching or pendent; stipes glabrous or scaly, sometimes also with minute glandular hairs, long or very short, bases often darker (phyllopodia) with abscision at their distal demarcation rather than at the rhizome; blades simple (flabellately divided in E. peltatum, pedate in one South American species), linear to ovate or oblanceolate, apices acuminate or caudate to obtuse, bases rounded to long-attenuate; costae grooved adaxially, usually with scales like those of the stipes abaxially; veins generally free, rarely netted or with a marginal commissural vein, simple to 2-forked, ending near the blade margins and lacking hydathodes, or stopping well short of the margins and usually ending in conspicuous hydathodes; blade scales frequently differing from those of the rhizomes or stipes, abundant to absent, often greatly reduced and appearing as stellate hairs; fertile fronds longer or shorter than the sterile fronds but generally with narrower blades and proportionally longer stipes; fertile blades completely covered abaxially with sporangia (acrostichoid sori); indusia absent; sporangia long-stalked, the annuli erect, each interrupted by the stalk; paraphyses generally absent, present in some species; spores bilateral, most with high crests or low ridges, but some echinate or verruculate without ridges or crests; x⫽40, 41. There are probably well over 600 species in Elaphoglossum, about three-fourths of them occurring in tropical America. There are 58 species recognized in Mexico, although our knowledge of them is far from complete. Several are known from only one or a few specimens. The genus is very difficult taxonomically. There are few characters, and the genus has not had a usable subgeneric breakdown until recently. Variation of the characters, such as plant size, blade shape and size, scale color, and scale type, is not fully understood in terms of species delimitation. The characters lie mostly in the scales of the rhizomes and blades. The fertile fronds add characters of relative size, intersporangial scales, and spore details, but virtually all the species can be identified on the basis of vegetative material alone. Unfortunately, collectors are hesitant to collect sterile material, and thus many records have gone uncollected.

In the following descriptions, rhizome diameter excludes the scale covering. Rhizome scales may be appressed or widely spreading and might distort the rhizome diameter measurements were they included. Stipe bases are differentiated into evident phyllopodia in some but not all species. Blade measurements apply to sterile fronds unless otherwise stated. Blade texture is difficult to determine on the basis of dried material and is given only for those species in which it is clearly distinctive. Although the veins seem to run to the blade margins in some species, they end just short of it, leaving pale, narrow margins 0.5–1 mm wide, which are often difficult to see, especially in very coriaceous or heavily indumented fronds. Vein angles and intervein distances are measured at mid-blade, halfway between the costa and margin. Scales of the rhizomes and blades are generally quite distinct from one another, and on the stipes they intergrade or remain distinct. Blade scales, although basically similar on adaxial and abaxial surfaces, are more highly dissected abaxially; in some this means longer teeth, but in more extreme cases scales are reduced to stellate hairs or even to resinous dots. Some dark scales are reduced and dissected so as to resemble spiders, a condition for which the term “arachnidoid” is used (vs. “arachnoid,” which is used for “web-like”). In species with subglabrous fronds, the blade scales are reduced minute stellate trichomidia that are visible only with a hand lens or dissecting scope. On fertile blades, scales of the adaxial surfaces are similar to those of sterile blades, but abaxially the scales are generally limited to the costae and in only a few species are there scales among the sporangia. Another type of indument consists of minute erect glandular hairs, which are found in varying degrees on the stipes and occasionally on the blades in E. lindenii, E. erinaceum, E. pallidum, and their relatives. Elaphoglossum is distinct by its simple blades (rarely with finely dissected blades in subsect. Peltapteris), free veins (rarely netted, but not in ours), and acrostichoid sori. Relationships to other genera are not clear. Elaphoglossum is commonly thought to be close to Lomariopsis, but molecular evidence suggests that Bolbitis is closer. The distinctness of many of the species described in the Rojas publications seems dubious and will require a great deal more study before one has confidence in their definition. Many seem no more than minor variants of already recognized species. Supposed new records are often based on misidentifications or misinterpretations. References Christ, H. 1899. Mongraphie des Genus Elaphoglossum. Neue Denkschr. Allg. Schweiz. Ges. Gesammten Naturwiss. [Mongr. Elaph.] 36: 1–159, t. 1–4; Go´mez P., L. D. 1975. Contribuciones a la pteridologı´a costarricense. VI. El ge´nero Peltapteris Link en Costa Rica. Brenesia 6: 25–31; Mickel, J. T. 1980a. Relationships of the dissected elaphoglossoid ferns. Brittonia 32: 109–117; Mickel, J. T. 1980b. Nine new elaphoglossums (Elaphoglossaceae) from Mexico. Brittonia 32: 334–339; Mickel, J. T. & L. Atehortu´a G. 1980. Subdivision of the genus Elaphoglossum. Amer. Fern J. 70: 47–68; Morton, C. V. 1955. Notes on Elaphoglossum. III. The publication of Elaphoglossum and Rhipidopteris. Amer. Fern J. 45: 11–14; Rojas-Alvarado, A. F. 1996. Twelve new species of Elaphoglossum (Elaphoglossaceae) from Costa Rica and Panama. Brenesia 45–46: 7–26; RojasAlvarado, A. F. 1997. Fourteen new species of Elaphoglossum (Elaphoglossaceae) from Mesoamerica. Brenesia 47–48: 1–16; Rojas-Alvarado, A. F. 2002. New spe-

ELAPHOGLOSSUM cies, new combinations and new distributions in neotropical species of Elaphoglossum (Lomariopsidaceae). Revista Biol. Trop. 50: 969-1006; Rojas-Alvarado, A. F. 2003a. New taxa, new records and redefined concepts in the Elaphoglossum sect. Elaphoglossum subsect. Pachyglossa (Lomariopsidaceae) from Mexico and Central America. Revista Biol. Trop. 51: 1-32; Rojas-Alvarado, A. F. 2003b. Notes

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on Elaphoglossum (Lomariopsidaceae) section Polytrichia subsection Hybrida in Mexico and Central America. Revista Biol. Trop. 51: 33-48; Skog, J. E., J. T. Mickel, R. C. Moran, M. Volovsek, & E. A. Zimmer. 2004. Molecular studies of the New World species in the fern genus Elaphoglossum based on chloroplast DNA sequences rbcL, trnL, rps4 and trnS. Int. J. Pl. Sci. (submitted)

Key to the Mexican Species of Elaphoglossum 1. Sterile blades 4–6 times flabellately divided; fertile fronds undivided, slightly bilobed; sterile fronds 3–15 cm long. ...................... 35. E. peltatum. 1. Sterile and fertile blades entire, undivided; sterile fronds 6–75 cm long. 2. Sterile fronds bearing linear-lanceolate scales that are usually inrolled and appear hair-like (subulate), at least on stipes, but generally throughout the fronds; veins often ending in hydathodes. 3. Blade scales dark brown, reddish brown, or black, rarely orange-tan, concentrated on stipes, blade margins, and costae, with virtually none on blade surfaces, rarely absent; hydathodes absent. 4. Scales conspicuous on blade costae and margins. ................................................................................................................. 8. E. erinaceum. 4. Scales very sparse to absent on blade costae and margins. 5. Rhizome scales dark reddish brown, with flexuous apices; blades elliptic with apices caudate. ............................ 33. E. pallidum. 5. Rhizome scales orange to brown, plane; blades lanceolate with apices acute to acuminate, rarely caudate. ....................................................................................................................................................................... 3. E. albomarginatum. 3. Blade scales orange or brown, more or less evenly distributed on the blade although scales of the surface may be much smaller than those of the costae and margins; hydathodes evident or not. 6. Blades oblanceolate; hydathodes absent. ..................................................................................................................................... 20. E. latum. 6. Blades linear to lanceolate; hydathodes present or absent. 7. Blades linear-elliptic, margins crenulate; marginal and costal scales larger than those of blade surfaces. .............. 9. E. eximium. 7. Blades linear to lanceolate, margins entire; marginal and costal scales not especiallylarger than those of blade surface. 8. Blades spatulate or obovate-lanceolate, rarely narrowly elliptic, apices obtuse; hydathodes often inconspicuous, generally in pits; fronds mostly 2–14 cm long. 9. Sterile blades oblong; fertile blades ovate to suborbicular, plane, with scales same color as those of sterile blades; hydathodes apparent; blades membranous; veins visible but not conspicuous. ............................ 41. E. pusillum. 9. Sterile blades spatulate, obovate-lanceolate, rarely to narrowly elliptic; fertile blades rounded, strongly conduplicate, with distinctly darker scales than those of the sterile blades; hydathodes inconspicuous to absent; blades chartaceous; veins obscure. 10. Fronds 2–4(–8) cm long; blades 1–3.5 ⫻ 0.4–1 cm; scales of the stipes and blades 1–2 mm long. ........................................................................................................................................................... 37. E. piloselloides. 10. Fronds 12–14 cm long; blades 7–8.5 ⫻ 1–1.3 cm; scales of the stipes and blades ca. 3 mm long. ............................................................................................................................................................ 18. E. jaliscanum. 8. Blades ovate to narrowly elliptic, apices acuminate, rarely cuspidate or rounded; hydathodes conspicuous; fronds mostly 8–60 cm long. 11. Rhizomes long-creeping; blade scales plane, not hair-like. ....................................................................... 51. E. squarrosum. 11. Rhizomes short-creeping; blade scales hair-like, with inrolled margins (subulate). 12. Blades cuneate at bases; fertile fronds with scales among the sporangia. 13. Fronds 8–18 ⫻ 0.8–1.5 cm; blade scales conspicuous; blade margins entire. ............................... 49. E. setosum. 13. Fronds 9–34 ⫻ 1.2–2.7 cm; blade scales sparse; blade margins slightly crenulate. .................... 19. E. lanceum. 12. Blades truncate to subcordate at bases; fertile fronds with no scales among the sporangia. 14. Blades elliptic to linear-lanceolate, bases truncate to rounded or broadly cuneate, apices acuminate; spores echinate. 15. Blades linear-lanceolate, length/width ratio 10–15:1. ............................................................. 48. E. setigerum. 15. Blades oblong-elliptic, length/width ratio 3–6:1. .................................................................... 28. E. monicae. 14. Blades ovate-lanceolate, bases rounded, apices obtuse to cuspidate, not acuminate; spores with low crests. 16. Blade apices obtuse to acute. .................................................................................................... 22. E. leonardii. 16. Blade apices cuspidate. 17. Blades broadly cuneate to rounded at bases; stipes and blades sparsely scaly to glabrous. ................................................................................................................................. 24. E. lindenii. 17. Blades rounded to truncate at bases; stipes and blades conspicuously scaly. ................ 38. E. pilosius. 2. Sterile fronds virtually glabrous with only minute stellate hairs to densely scaly; scales various (broadly lanceolate or ovate, ciliate, stellate, round-peltate) but not subulate; hydathodes absent. 18. Scales present on stipes and blades. 19. Abaxial blade surfaces glabrous or with stellate hairs/scales, trichomidia, and/or resinous dots (other types of indument may be present also).

284

ELAPHOGLOSSUM

20. Rhizome scales orange, dull; abaxial blade surfaces with stellate hairs, simple hairs, or 3-armed trichomidia. 21. Abaxial blade surfaces with stellate hairs. ................................................................................................................ 29. E. muelleri. 21. Abaxial blade surfaces with simple to 3-armed trichomidia. .......................................................................... 39. E. potosianum. 20. Rhizome scales castaneous to black, lustrous, at least in part; abaxial blade surfaces glabrous, glandular-punctate, or with stellate to substellate hairs/scales. 22. Rhizomes moderately or long-creeping, the fronds to 1 cm distant. .................................................................. 14. E. hartwegii. 22. Rhizomes compact to short-creeping, the fronds approximate. 23. Scales of stipes and adaxial blade surfaces round, peltate. ............................................................................... 52. E. tectum. 23. Scales of stipes and to a lesser degree the adaxial blade surfaces ovate, lanceolate, or occasionally absent. 24. Blade apices obtuse; blade scales ovate, entire, mostly along adaxial margins; rhizome apices ascending. ..................................................................................................................................................... 16. E. huacsaro. 24. Blade apices acute to acuminate; blade scales lanceolate or stellate, along margins or on blade surfaces; rhizomes horizontal. 25. Abaxial blade scales stellate or substellate. 26. Abaxial blade scales substellate, with at least a narrow scale body. ....................................... 40. E. pringlei. 26. Abaxial blade scales stellate, lacking any scale body. 27. Stellate hairs (lacking scale body) present abaxially; substellate scales with some scale body adaxially; rhizome scales sparsely toothed. ......................................................... 17. E. ipshookense. 27. Stellate hairs present both abaxially and adaxially; rhizome scales entire. .............. 53. E. tenuiculum. 25. Abaxial blade scales linear-lanceolate, or with trichomidia or resinous dots, rarely the blades lacking indument altogether. 28. Rhizome scales entire to faintly dark-toothed, scales of stipe bases deeply toothed; stipes 1⁄3– 2 ⁄3 the frond length; blades 4–9 cm long; resinous dots absent. ............................................. 12. E. gratum. 28. Scales of rhizomes and stipe bases entire; stipes ca. 1⁄8–1⁄2 the frond length; blades 10–30 cm long; resinous dots commonly on one or both blade surfaces. 29. Abaxial blade surfaces with 1–3-armed trichomidia. .................................................. 39. E. potosianum. 29. Abaxial blade surfaces with scales or resinous dots. 30. Abaxial blade surfaces usually with resinous dots, occasionally with scattered linear-lanceolate scales or substellate hairs. ...........................................................36. E. petiolatum. 30. Abaxial blade surface with abundant ciliolate scales. ........................................... 45. E. rzedowskii. 19. Abaxial blade surfaces bearing scales but lacking resinous dots, stellate hairs, and/or trichomidia. 31. Blades 1.9–4.8 cm wide, ovate-lanceolate to elliptic, apices rounded; spores verruculate. 32. Rhizome scales with short teeth; stipe scales erose; blade scales with cilia shorter than the width of the scale body; rhizomes short-creeping to erect. ................................................................................................................ 30. E. muscosum. 32. Rhizome scales ciliate; stipe scales cilio-denticulate; blade scales with cilia as long as the width of the scale body; rhizomes moderately to short-creeping, horizontal. ........................................................................................ 7. E. engelii. 31. Blades 0.6–3.2 cm wide, lanceolate, linear-lanceolate, or narrowly elliptic, apices acute to acuminate, rarely obtuse; spores with crests or ridges, not verruculate. 33. Rhizome scales tan to orange with irregular dark streaks. ................................................................................. 2. E. alansmithii. 33. Rhizome scales uniformly colored or with dark tips, lacking dark streaks. 34. Abaxial blade scales linear or inrolled and erect, scattered. 35. Rhizome scales orange-tan; Oax, Ver, Chis. ......................................................................................... 4. E. auricomum. 35. Rhizome scales castaneous. 36. Stipe scales 2–8 mm long; blades 3–6 cm wide; abaxial blade scales erect; Ver, Hgo, Pue. .... 31. E. obscurum. 36. Stipe scales ca. 2 mm long; blades 1.4–3.2 cm wide; abaxial blade scales more or less plane or with margins turned up, rarely whole scale erect. 37. Rhizome scales weakly toothed; abaxial blade scales with teeth and margins upcurved; Chis. ............................................................................................................................................. 43. E. rubescens. 37. Rhizome scales strongly toothed; abaxial blade scales more or less plane, rarely erect; Qro, Hgo, Pue, Ver, Oax, Chis. ......................................................................................................... 55. E. vestitum. 34. Abaxial blade scales lanceolate, scattered to imbricate. 38. Stipes 1⁄10–1⁄5 the sterile frond length; fertile blades linear; Gro, Oax. ..................................... 26. E. manantlanense. 38. Stipes 1⁄4–1⁄2 the sterile frond length; fertile blades elliptic. 39. Rhizome scales brown to black, strongly toothed, teeth mostly persistent, rigid; abaxial scales scattered to imbricate; stipes 1⁄4–1⁄3 the sterile frond length; central and southern Mexico. .............................................................................................................................................. 32. E. paleaceum. 39. Rhizome scales castaneous to blackish brown, weakly toothed, the teeth somewhat deciduous; abaxial blade scales imbricate; stipes 1⁄3–1⁄2 the sterile frond length; central and western Mexico. ................................................................................................................................................ 44. E. rufescens. 18. Scales absent on stipes and blades except, in some cases, for stellate trichomidia or sparse inconspicuous linear scales (1–2 mm) on the blades or ovate scales on the stipes.

ELAPHOGLOSSUM AFFINE

285

40. Rhizomes (short-) moderately to long-creeping, fronds (1–)3–30 mm distant. 41. Phyllopodia absent; fronds 5–15 cm long; rhizomes ca. 1 mm diam. 42. Blades spatulate; (2000–)2450–3050 m. ................................................................................................................ 50. E. squamipes. 42. Blades narrowly elliptic; 600–1000 m. .................................................................................................................. 42. E. revolutum. 41. Phyllopodia distinct; fronds 8–50 cm long; rhizomes (1–)2–10 mm diam. 43. Rhizome scales mostly orange-tan but at least some with black or dark brown streaks. 44. Blade apices acute to acuminate; fronds 21–40 cm long; blades 1.5–2.9 cm wide. ................................. 27. E. mcvaughii. 44. Blade apices acute to rounded, or if acuminate, 1 cm or less wide; fronds 6–30 cm long; blades 0.6–2.6 cm wide. 45. Blades 1.2–2.6 cm wide; rhizomes short- to long-creeping; fronds 3–30 mm apart. ............................... 1. E. affine. 45. Blades 0.6–1 cm wide; rhizomes short- to moderately creeping; fronds 1–8 mm apart. ............. 54. E. tenuifolium. 43. Rhizome scales dark brown to tan, concolorous. 46. Rhizome scales lustrous dark brown. .......................................................................................................... 13. E. guatemalense. 46. Rhizome scales dull tan. 47. Rhizomes 3 mm diam.; fronds 2–4 cm distant. ................................................................................. 47. E. seminudum. 47. Rhizomes 4–6 mm diam.; fronds 0.5–1.7 cm distant. 48. Rhizome scales ovate to ovate-lanceolate. ...................................................................................... 6. E. dissitifrons. 48. Rhizome scales linear-lanceolate. ........................................................................................................... 56. E. viride. 40. Rhizomes compact or short-creeping, fronds approximate. 49. Blades thin-textured, veins evident, vein ends laterally expanded and sometimes joining. 50. Stipes 1⁄5–1⁄3 the frond length. .......................................................................................................................... 25. E. lonchophyllum. 50. Stipes nearly absent, 1–2 cm long. .......................................................................................................................... 5. E. decursivum. 49. Blades firm to coriaceous, veins obscure (or if barely evident, not fusing at margins). 51. Stipes essentially absent (0–2 cm long on sterile fronds); blades linear-elliptic; sterile fronds pendent, 0.5–2 m long; rhizome scales often 1.5–3 cm long, bright orange; fertile fronds much shorter than the sterile fronds, 1⁄5–1⁄3 the sterile frond length. ................................................................................................................... 15. E. herminieri. 51. Stipes present (over 4 cm long); blades elliptic to oblanceolate; sterile fronds erect or spreading; 0.2–0.7 mm long; rhizome scales 1–10 mm long; fertile fronds 1⁄2 to exceeding the sterile frond length. 52. Rhizome scales linear, dark reddish brown to black. 53. Blade apices acute to obtuse or rounded; veins obscure; stipes 1⁄2–3⁄5 the frond length; blades lacking subulate scales; phyllopodia distinct. ........................................................................................... 11. E. glaucum. 53. Blade apices caudate; veins evident; stipes 1⁄3–1⁄2 the frond length; blades often with a few subulate scales; phyllopodia absent. ........................................................................................................................ 33. E. pallidum. 52. Rhizome scales lanceolate to deltate-lanceolate or linear-lanceolate, orange or tan, to dark brown or black. 54. Stipes with tan, lanceolate to ovate scales. ........................................................................................ 23. E. lepidopodum. 54. Stipes lacking scales, or if scales present then linear or minute or reduced to trichomidia. 55. Blades 0.5–2 cm wide; rhizome scales lustrous brown to black; rhizome 2–4 mm diam. 56. Fertile fronds shorter than sterile; sterile blades 0.5–1 cm wide, glabrous; phyllopodia 4–8 mm long. .................................................................................................................................... 10. E. glabellum. 56. Fertile fronds longer than the sterile; sterile blades 1–2 cm wide, often with minute black stellate trichomidia; phyllopodia 1–1.5 cm long. ...................................................................... 34. E. parduei. 55. Blades (1.5–)2.4–7 cm wide; rhizome scales orange to dull tan, or dirty brown, occasionally somewhat lustrous; rhizomes (3–)4–10 mm diam. 57. Rhizome scales 5–20 mm long. 58. Rhizome scales linear, 5–10 mm long, concolorous orange-tan. .............................. 58. E. yourkeorum. 58. Rhizome scales linear-lanceolate, 15–20 mm long, orange-tan tinged lustrous black .................................................................................................................................. 21. E. leebrowniae. 57. Rhizome scales mostly 5–8 mm long. 59. Fertile blades linear to narrowly oblong; sterile blade bases 65–95⬚ angle. ......... 57. E. xanthopodum. 59. Fertile blades narrowly elliptic; sterile blade bases 30–50⬚ angle. .................................. 46. E. sartorii.

1. ELAPHOGLOSSUM AFFINE Elaphoglossum affine (M. Martens & Galeotti) T. Moore, Index Fil. 4. 1857. Figs. 127Q–S. Acrostichum affine M. Martens & Galeotti, Me´m. Foug. Mexique 24, pl. 3, f. 1. 1842. Type. Mexico. Veracruz: Volca´n Orizaba, Galeotti 6454 (BR!; isotype LE!).

Rhizomes short- to long-creeping, 1–2 mm diam.; rhizome scales dense to sparse, ovate, orange-tan to tan with varying degrees of dark, lustrous, sclerotic streaking, ca. 3 mm long, deeply cordate, appearing nearly peltate, entire or with occasional small, irregular teeth; fronds 3–30 mm apart, 7–30 ⫻ 1.2–2.6 cm; phyllopodia distinct; stipes 1⁄2–2⁄3 the frond length, glabrous or with a few tan, ovate scales; blades narrowly elliptic, coriaceous, apices acute to rounded, bases cuneate; veins obscure, free, simple or

286

ELAPHOGLOSSUM

once-forked, ca. 1 mm apart, at 60–70⬚ angle to costa; hydathodes absent; blade surfaces with sparse, minute, stellate trichomidia abaxially, trichomidia sparse and larger adaxially; fertile fronds similar to the sterile in blade size and shape but often somewhat longer, intersporangial scales absent.

size and shape but usually somewhat longer, intersporangial scales absent.

Distribution. Epiphytic in cloud forests; 1900–3000 m. Mexico. Specimens Examined. Chis (Breedlove 22732, DS, 40809, DS, 53377, CAS). Oax (Mickel 3735, NY).

Distribution. Terrestrial or epipetric in montane cloud forests; 2650–3150 m. Mexico; Guat, Salv, CR, Pan; Col, Ven. Selected Specimens Examined. Chis (Breedlove 24307, DS). DF (Lyonnet 750, CAS). Gro (Lorea 4202, FCME, IEB). Me´x (Rzedowski 21862, ENCB). Mich (Hinton 15710, NY). Mor (Cha´vez 34, UAMIZ). Oax (Camp 2683, NY; Mickel 1154, NY). Ver (Galeotti 6454, BR; Va´zquez 1673, XAL)

This species and E. tenuifolium, E. mcvaughii, and E. alansmithii form a closely related group, all with darkly streaked rhizome scales; they differ in their blade shape and details of blade scaliness. Elaphoglossum affine also closely resembles E. guatemalense, which has shorter rhizomes, and uniformly dark, lustrous brown rhizome scales (scales orange-tan with dark streaks in E. affine).

2. ELAPHOGLOSSUM ALANSMITHII Elaphoglossum alansmithii Mickel, Brittonia 32: 334. 1980. Type. Mexico. Oaxaca: Distrito Ixtla´n, trail from San Pedro Nolasco to the Llano Verde, Mickel 3739 (NY!). Figs. 141F–H, J, K. Rhizomes moderately creeping, ca. 2 mm diam.; rhizome scales dense to sparse, ovate-lanceolate, tan to orange with varying degrees of dark, lustrous, sclerotic streaking, 2–4 mm long (deeply cordate, appearing peltate), entire or with occasional small, irregular teeth; fronds 2–10 mm apart, (4–)9–22(–41) ⫻ 0.9–2.6 cm; phyllopodia distinct; stipes 1⁄6–1⁄3 the frond length, with sparse tan ovate scales, at least at the bases; blades narrowly elliptic, coriaceous, apices acuminate, bases cuneate; veins obscure, free, simple or once-forked, ca. 1 mm apart, at 60–70⬚ angle to costa; hydathodes absent; adaxial blade surfaces essentially glabrous; abaxial surfaces with linear scales with hastate or stellate bases, or scales deeply stellate; fertile fronds similar to the sterile in blade

This species closely resembles E. affine in the dark streaking of the rhizome scales and blade size and shape but is noticeably more scaly, and the blade apices are distinctly acuminate. The type collection consists of plants about the size of specimens of E. affine (4–22 cm long), but Mickel 3735, from the same locality, has longer fronds (28–41 cm).

3. ELAPHOGLOSSUM ALBOMARGINATUM Elaphoglossum albomarginatum A. R. Sm., Proc. Calif. Acad. Sci., ser. 4, 40: 220, f. 4A. 1975. Type. Mexico. Chiapas: Unio´n Jua´rez, Volca´n Tacana´, Breedlove 31629 (DS!). Figs. 133A–C. Rhizomes short-creeping, stout, to 10 mm diam.; rhizome scales orange to brown, ribbon-like or nearly hair-like, to 10 mm long; fronds clumped, 26–60 ⫻ 5–8.5 cm; phyllopodia absent; stipes 1⁄3–1⁄2 the frond length, with spreading, subulate scales or glabrous, usually with minute, erect, glandular hairs; blades lanceolate, apices acute to acuminate, rarely caudate, bases rounded; veins evident, free, 1–2-forked, 1.5–2 mm apart, at 70–80⬚ angle to costa; hydathodes absent; margins pale, ca. 0.5 mm wide; blades completely lacking scales, but with minute, black, stellate hairs; fertile fronds smaller than the sterile, with a conspicuous sterile margin ca. 1 mm wide, scales absent on abaxial costa and among the sporangia.

ELAPHOGLOSSUM DECURSIVUM

Distribution. Epiphytic, epipetric, or terrestrial, in montane rain forests; 2200 m (1800–2500 m in Guat). Mexico; Guat. Known in Mexico only from the type. Elaphoglossum albomarginatum closely resembles a large specimen of E. pallidum, but differs not only in size but also in the rhizome scales and the pale margins on both the sterile and fertile blades. Marginal blade scales are absent in E. albomarginatum, whereas in E. pallidum at least an occasional marginal scale is found.

4. ELAPHOGLOSSUM AURICOMUM Elaphoglossum auricomum (Kunze) T. Moore, Index Fil. 7. 1857. Figs. 132D, E. Acrostichum auricomum Kunze, Linnaea 9: 28. 1834. Type. Peru. Pampayaco, Jul 1829, Poeppig s.n. (LZ destroyed, frag. P!).

Rhizomes short-creeping, to 5 mm diam.; rhizome scales linearlanceolate, orange-tan, to 7 mm long, margin with hair-like teeth; fronds clumped, 18–50 ⫻ 1.6–3.2 cm; phyllopodia inconspicuous or absent; stipes 1⁄8–1⁄4 the frond length, densely clothed with spreading, orange scales 4 mm long; blades narrowly elliptic, papyraceous, apices acuminate, bases narrowly cuneate; veins indistinct, free, simple or once-forked, 1–2 mm apart, at 60–70⬚ angle to costa; hydathodes absent; blade scales orange, scattered, not overlapping, skeletonized, linear, 1–4 mm long with long, hair-like teeth, the teeth at least half as long as the scales, some scales reduced to nearly stellate hairs; fertile fronds with longer stipes, ca. 1⁄3 the fertile frond length, with a few stellate hairs on the abaxial costa.

287

Distribution. Epiphytic in montane rain forests; 180–1250 m. Mexico; Guat, Hond, CR; Hisp; Col, Ven, Ec, Peru, Bol. Specimens Examined. Chis (Breedlove 32937, DS; 38933, DS; Martı´nez S. 23961, MEXU). Oax (Mickel 6442, NY; Yatskievych 83-407, ARIZ). Ver (Herna´ndez M. 1385, MEXU; Lira 227, XAL; Mu¨ller 386, NY).

This species is close to E. vestitum in the skeletonized blade scales and the general laxness of the fronds, but differs in several features. Elaphoglossum auricomum lacks caudate blade tips, the rhizome scales are not as dark, and the stipes are densely covered with spreading orange scales. It also resembles E. paleaceum, but the rhizome scales are dark orange or maroon, not black as usually found in E. paleaceum, and the blade scales are more sparse and more highly dissected than in that species. Some specimens have very short stipes, but apparently this character is quite variable in E. auricomum.

5. ELAPHOGLOSSUM DECURSIVUM Elaphoglossum decursivum Mickel, Brittonia 32: 334. 1980. Type. Mexico. Oaxaca: Distrito Ixtla´n, 29 km S of Valle Nacional, 80 km N of Ixtla´n de Jua´rez, trail E of Rte 175 at Campamento Vista Hermosa toward Ladu´, 1 hour hike down to Rı´o de la Trucha, Mickel 6417 (NY!; isotypes ENCB!, NY!, UC!). Fig. 131A. Rhizomes short-creeping, to 3 mm diam.; rhizome scales lanceolate to ovate, concolorous light brown, not resinous, to 3 mm long, entire; fronds clumped, 55–62 ⫻ 2.2–4 cm; phyllopodia very short, indistinct; stipes nearly absent, only 1–2 cm long, with minute, stellate, appressed hairs; blades narrowly elliptic, very thin, apices acuminate, bases narrowly cuneate, long-decurrent nearly to the rhizomes; veins evident, free, simple or once-forked, but vein tips expanded laterally, sometimes uniting, at 75⬚ angle to costa; hydathodes absent; blade scales absent, but minute, stellate, reddish, resinous hairs present, mostly on the abaxial surfaces; fertile fronds shorter than the sterile, 40–46 cm long, stipe 1 ⁄3–1⁄2 the fertile frond length, blades 1.2–1.6 cm wide, intersporangial scales absent.

288

ELAPHOGLOSSUM

Distribution. Epiphytic in wet montane forests; 450–850 m. Mexico; Bel, Hond, CR, Pan. Specimen Examined. Oax (Mickel 7205, NY).

Elaphoglossum decursivum closely resembles E. lonchophyllum in its slender rhizomes, minute stellate hairs on stipes and blades, thin blades, and the vein tips laterally extended and sometimes uniting, but the former differs in the lack of resinous rhizome scales and the virtually sessile fronds.

6. ELAPHOGLOSSUM DISSITIFRONS Elaphoglossum dissitifrons Mickel, Brittonia 32: 335. 1980. Type. Mexico. Guerrero: Ⳳ2 km al NE del Campamento El Gallo, estribacio´nes suroccidentales del Cerro Teotepec, Rzedowski & McVaugh 174 (NY!; isotypes ENCB!, MICH!). Figs. 132F–H, 140A–D.

Distribution. Terrestrial, epipetric, or epiphytic in moist pineoak forests; 1900–2900 m. Mexico. Selected Specimens Examined. Gro (Hinton 14303, NY, US; Lorea 2341, FCME, IEB, 2514, NY, 4213, FCME, IEB; Rzedowski & McVaugh 73, ENCB, MICH, 143, ENCB, MICH). Me´x (Hinton 3455, LL, NY; Matuda et al. 38095, US). Oax (Mickel 4046, NY; Zamudio 8381, IEB).

The creeping rhizomes are similar to those of E. seminudum and E. mcvaughii, but the latter has narrower blades and bicolorous rhizome scales. Elaphoglossum dissitifrons is superficially most like E. sartorii in frond size and shape but is more closely related to E. affine, as evidenced by the broader, ovate rhizome scales.

7. ELAPHOGLOSSUM ENGELII Elaphoglossum engelii (H. Karst.) Christ, Monogr. Elaph. 81. 1899. Figs. 135A–F.

Elaphoglossum clewellianum Mickel, Mem. New York Bot. Gard. 46: 172. 1988. Type. Mexico. Oaxaca: Distrito Centro, N slope of Cerro San Felipe, Mickel 4034 (NY!; isotypes ENCB!, US!).

Acrostichum engelii H. Karst., Fl. Columb. 1: 119, t. 59. 1860. Type. Colombia. Karsten s.n. (LE).

Rhizomes long-creeping, 4–6 mm diam.; rhizome scales ovate to ovate-lanceolate, dull tan, concolorous, mostly 5–7 mm long, entire or each with an occasional tooth or short marginal hair, scales dense, not deciduous; fronds 5–17 mm apart, 25–34(–47) ⫻ 2.6–4.3 cm; phyllopodia distinct, 2–3 cm long; stipes 1⁄5–2⁄5 (–1⁄2) of the frond length, with scattered ovate to ovate-lanceolate scales, tan, occasionally with dark sclerotic tips, entire to irregularly toothed; blades narrowly elliptic, apices acute, bases broadly cuneate, coriaceous; veins obscure, free, simple or onceforked, ca. 1 mm apart, at 70–75⬚ angle to costa; hydathodes absent; blade surfaces glabrous or subglabrous with sparse, minute, stellate trichomidia (0.1 mm), these sometimes reduced to mere dots; costae with occasional broader scales; fertile fronds equal to or slightly longer that the sterile, the stipes 2⁄5–2⁄3 of the fertile frond length, the blades similar in shape to those of sterile fronds, but narrower (1.8–2 cm wide), scales absent on abaxial blade surfaces.

Rhizomes to 4 mm diam., short- to moderately creeping; rhizome scales 8 mm long, linear, reddish orange, lustrous, with long hair-teeth; fronds 24–30 cm, approximate to slightly distant; phyllopodia inconspicuous or absent; stipes 1⁄2 of sterile frond length, orange-tan; stipe scales mostly 3–5 mm long, lanceolate, spreading, orange or with black centers, strongly ciliodenticulate, the shorter ones with longer hair-teeth; sterile blades 6–15 ⫻ 2–4.2 cm, narrowly elliptic or lanceolate, subcoriaceous, bases broadly cuneate, apices obtuse; veins obscure, c. 1.5 mm apart, at 60–70⬚ angle to costa; hydathodes absent; blade scales mostly 1.5–3 mm long, abundant, cilio-denticulate, adaxially whitish with age, imbricate to deciduous, abaxially orange with some lamina visible between the scales; fertile fronds slightly longer than the sterile, stipes 2⁄5 of frond length, blades narrower than those of sterile fronds, scales abundant adaxially and on abaxial costa, intersporangial scales conspicuous and abundant or deciduous, with long hair-teeth.

ELAPHOGLOSSUM ERINACEUM

289

8a. Elaphoglossum erinaceum var. erinaceum Figs. 127E, 139D–F. Acrostichum crinitum M. Martens & Galeotti, Me´m. Foug. Mexique 25. 1842, hom. illeg., non L., 1753. Acrostichum blepharodes Fe´e, Me´m. Foug. 2: 48, pl. 24, f. 3. 1845, nom. nov. Elaphoglossum blepharodes (Fe´e) T. Moore, Index Fil. 7. 1857. Type. Mexico. Veracruz: Zacuapa´n, Galeotti 6297 (P, not found; isotype BR!). Acrostichum nitidum Liebm., Mexico. Bregn. 168 (reprint 16). 1849. Elaphoglossum fimbriatum T. Moore, Index Fil. 356. 1862, nom. nov., non Elaphoglossum nitidum Brack., 1854. Type. Mexico. Puebla: Chinautla, Liebmann s.n. [Pl. Mex. 2421] (C!; isotypes P!, US!). Acrostichum mexicanum E. Fourn., Mexic. Pl. 1: 63. 1872. Syntypes. Mexico. Puebla: Chinautla, Tizutla´n, Liebmann s.n. (P!; isotype K!); Veracruz: Misantla, Hahn s.n. (P!).

Distribution. Epiphytic in mixed forests; 3000–3200 m. Mexico; Guat, CR, Pan; Hisp; Col, Ven, Ec, Peru, Bol.

Differing from var. occidentale by the black to dark reddish brown (vs. orange-tan) blade margin scales and its occurrence in southern and eastern (vs. western) Mexico.

Specimens Examined. Me´x (Lyonnet 2024, ENCB, MEXU, US; Rzedowski 23279, ENCB, NY, 33968a, 36023, IEB). Mor (Lyonnet 1413, CAS, ENCB, FCME, UAMIZ, UC, US, 2024, CAS; Sa´nchez S. 465, MEXU, US).

Elaphoglossum engelii is closely allied to E. muscosum, but distinct by the hair-toothed rhizome scales and fringed (especially the larger scales) vs. erose to short-toothed stipe scales.

8. ELAPHOGLOSSUM ERINACEUM Elaphoglossum erinaceum (Fe´e) T. Moore, Index Fil. 9. 1857. Acrostichum erinaceum Fe´e, Me´m. Foug. 2: 41. 1845. Lectotype (chosen by Proctor 1977: 215). Guadeloupe. Rivie`re de Saint-Louis, de Thiouville s.n. (P!).

Rhizomes short-creeping, to 7 mm diam.; rhizome scales linear, orange to castaneous, to 17 mm long, entire or with very irregular teeth near the tips; fronds clumped, 23–73 ⫻ 2.3–8 cm; phyllopodia absent; stipes 1⁄3–1⁄2 the frond length, with scales dark reddish brown, subulate, widely spreading, to 5 mm long, also with minute erect glandular hairs on stipes and costae; blades lanceolate to elliptic, chartaceous, apices acuminate to caudate, bases cuneate to rounded; veins evident, free, simple or once-forked, 1–2 mm apart, at 70–80⬚ angle to costa; hydathodes absent; blade scales subulate, dark reddish brown, often appearing black (orange-tan in var. occidentale), mostly on the midveins and spreading from the margins, also with minute, simple, branched, or stellate hairs on the abaxial surfaces; fertile fronds considerably shorter than the sterile (ca. 1⁄2 the length of the sterile) and somewhat narrower, lanceolate to linear-lanceolate, with a noticeable sterile margins, scales absent on the abaxial surfaces; spores with broad, low, smooth ridges.

Distribution. Epiphytic and epipetric in montane rain forests and cloud forests; 800–2800 m. Mexico; Guat, Salv, Hond, Nic, CR, Pan; Gr & L Ant; Col, Ven, Ec, Peru, Braz, Bol. Selected Specimens Examined. Chis (Breedlove 22347, NY; Me´ndez G. 8041, MEXU; Riba 1373, UAMIZ). Hgo (Alonzo s.n., 29 Jul 1978, FCME). Oax (Anderson & Anderson 4728, NY; Mickel 4904, 5646, NY). Pue (Campos V. 499, NY; Orcutt 4009, MEXU). Ver (Lorence 3873, NY).

Elaphoglossum erinaceum belongs to a very perplexing group. The Mexican material shows considerable variation in blade size and relative width, rhizome scale size and color, degree of scaliness, and habitat. Further study may show this material to represent more than one taxon. Of special note are Mickel 1083 and 5646 (both Oaxaca, NY), which have very large blades (8.5–10 cm wide, 45–50 cm long), long stipe scales (4–8 mm) and large rhizome scales (ca. 1 cm long and 2 mm wide). These may in fact represent a distinct species, in which case the name would be E. fimbriatum T. Moore.

Key to the Mexican Varieties of Elaphoglossum erinaceum 1. Scales of blade margins black to dark reddish brown; eastern Mexico. ........................................................... 8a. var. erinaceum. 1. Scales of blade margins orange-tan; western Mexico. ....................................................................... 8b. var. occidentale.

8b. Elaphoglossum erinaceum var. occidentale Mickel, var. nov. Type. Mexico. Me´xico: About 6 mi S of Tenancingo and 8 mi N of Villa Guerrero on Rte 55, Mickel 706 (holotype NY!; isotypes ENCB, MICH!). Fig. 133K.

290

ELAPHOGLOSSUM

Ab var. erinaceo paleis laminarum melleis brevibus et distributione geographica occidentali distincta. (L., occidentalis, western, alluding to the westerly distribution of this variety in Mexico.)

Distribution. Epiphytic, less commonly epipetric and terrestrial in wet forests; 1850 m. Mexico; Guat, Hond, Salv, CR, Pan; Hisp; Col, Ven, Ec, Peru, Bol. Distribution. Terrestrial on moist shaded banks in pine-oak or broad-leaved evergreen forests; 1400–2600 m. Mexico. Specimens Examined. Gro (Lorea 2225, NY; Rzedowski 23682, 26011, NY). Jal (Boutin & Brandt 2570, NY; McVaugh 14319, 21439, 23196, NY). Me´x (Bartholomew 2968, NY; Purpus 1594, UC). Mich (Arse`ne s.n., 13 Dec 1910, MICH; Garcı´a & Nava 3069, NY; Hinton 15609, PH; Rzedowski 25175, ASU, NY; Yatskievych 81-417, NY). Mor (Lyonnet 2863, US; Pringle 6546, CAS, MEXU, MSC, UC, 15713, MSC, UC, US; Sa´nchez 394, MEXU). Qro (Dı´az B. & Carranza 6557, IEB, MEXU). Sin (Quin s.n., 20 Jan 1982, ARIZ; Vega A. 3131, ENCB).

Specimens from western Mexico, Sinaloa to Guerrero, generally have scales of blade margins orange-tan rather than brown to black and often shorter (1–2 vs. 3–4 mm), though the scale size is quite variable.

9. ELAPHOGLOSSUM EXIMIUM Elaphoglossum eximium (Mett.) Christ, Monogr. Elaph. 107. 1899. Figs. 133D–G. Acrostichum eximium Mett., Ann. Sci. Nat. Bot., se´r. 5, 2: 199. 1864. Type. Colombia. Manzanos, Lindig 315 (B!; isotypes BM!, P!–2 sheets.).

Rhizomes ca. 5 mm diam., compact to short-creeping; rhizome scales 3–5 mm long, linear, dark brown, lustrous, entire; fronds (16–)20–44 cm long, approximate; phyllopodia absent; stipes ca. 1 ⁄8 of sterile leaf length, orange-tan; stipe scales ca. 2 mm long, abundant, brown, spreading, subulate; blades 13–27 ⫻ 0.9–1.8 cm, linear-elliptic, chartaceous, bases attenuate, apices acuminate, margins crenulate; veins evident, ca. 2 mm apart, at 45–50⬚ angle to costa; hydathodes present; blade scales subulate, brown, conspicuous but sparse on costae, smaller on margins, mostly between crenulations, indument of stellate trichomidia or punctae, less commonly of scattered small (0.5 mm) linear or lanceolate scales; fertile fronds shorter than the sterile, stipes 3⁄4 of the fertile frond length, blades narrowly lanceolate, entire, bases truncate to cordate, apices acute, intersporangial scales absent.

Specimens Examined. Chis (Gittins 4171, NY). Oax (Geith 95-M017 bis, NY).

Elaphoglossum eximium is distinct by the linear, crenulate blades with small subulate scales and hydathodes.

10. ELAPHOGLOSSUM GLABELLUM Elaphoglossum glabellum J. Sm., London J. Bot. 1: 197. 1842. Figs. 126K, L. Acrostichum glabellum (J. Sm.) Klotzsch, Linnaea 20: 421. 1847. Type. Guyana. Schomburgk 447 (BM!; isotypes P!, US!).

Rhizomes short- to moderately creeping, 2–3 mm diam.; rhizome scales lanceolate, lustrous dark brown to black, with pale brown bases, margin, and tip, 1–2 mm long, entire or with a few weak teeth, scales often somewhat deciduous and leaving partially naked rhizomes; fronds slightly spaced, to 33 ⫻ 0.5–1 cm; phyllopodia distinct, 4–8 mm long; stipes 1⁄10–1⁄5 the frond length, glabrous or with minute appressed scales; blades linear-elliptic, coriaceous, gradually acuminate at both ends; veins obscure, free, simple or once-forked, ca. 1 mm apart, at ca. 70⬚ angle to costa; hydathodes absent; blade surfaces glabrous or with very sparse minute stellate trichomidia on the abaxial surfaces; fertile fronds slightly shorter than the sterile fronds, the stipes longer (1⁄3–1⁄2 the fertile frond length) and the blades broader (ca. 10 mm wide), scales absent; 2n⫽ca. 164 (Trin).

ELAPHOGLOSSUM GRATUM

Distribution. Epiphytic in moist oak woods; 450–1300 m. Mexico; Salv, CR, Pan; Gr & L Ant; Col, Ven, Trin, Guy, Sur, Ec, Peru, Bol. Specimens Examined. Oax (Mickel 4800, 6522, 6673, 6696, NY).

See E. parduei for comparison with that species.

11. ELAPHOGLOSSUM GLAUCUM Elaphoglossum glaucum T. Moore, Index Fil. 10. 1857. Figs. 124L–N. Acrostichum glaucum Fe´e, Me´m. Foug. 8: 67. 1857, hom. illeg., non Cav., 1799. Type. Mexico. Veracruz: Co´rdoba, 1854, Schaffner 16 (P!). Elaphoglossum eucraspedum Christ, Bull. Herb. Boissier, se´r. 2, 7: 415. 1907. Type. Mexico. Chiapas: Mu¨nch 1902 (P!).

Rhizomes short-creeping, ca. 5 mm diam.; rhizome scales linear-deltate, often hair-like most of their length, lustrous dark reddish brown to nearly black, to 17 mm long, entire; fronds clumped or a few mm apart, 13–62 ⫻ 1.7–3.8 cm; phyllopodia distinct; stipes 1⁄2–3⁄5 the frond length, glabrous; blades narrowly elliptic to oblanceolate, coriaceous, apices acute to obtuse or rounded, bases cuneate; veins obscure, free, curved, simple or once-forked, 1–1.5 mm apart, at ca. 70⬚ angle to costa; hydathodes absent; blade surfaces glabrous to sparsely beset with minute, linear, hastate scales to 1–2 mm long; fertile fronds slightly shorter to somewhat longer than the sterile, but otherwise nearly equal in blade size and shape.

291

Distribution. Epiphyte in wet oak or pine-oak woods, cloud forests; 1100–3200 m. Mexico; Guat, Hond, Salv. Selected Specimens Examined. Chis (Breedlove 22731, NY). Gro (Rzedowski 23684, NY). Hgo (Moore 5307, UC, US). Jal (McVaugh 23106, MICH). Me´x (Hinton 8997, NY). Mich (Hinton 15372, NY). Mor (Herrera s.n., MEXU). Nay (Kral 2566, ENCB). Oax (Mickel 4670, NY). Pue (Pringle 10836, US). Qro (Dı´az B. 4979, IEB). Ver (Valdivia Q. 2212, IEB, XAL).

This species is most easily distinguished by the obtuse to rounded blade tips and the dark rhizome scales. Elaphoglossum guatemalense may sometimes have obtuse blades but has shorter, broader, darker rhizome scales and consistently has more slender rhizomes and minute stellate trichomidia on the abaxial blade surfaces. Elaphoglossum sartorii has orange or brownish rhizome scales, and its blades are acute to acuminate. The type has golden rhizome scales in contrast to the dark maroon scales typical of most specimens. Specimens from Orizaba at NY have castaneous scales and material from Jalisco, Guerrero, and Michoaca´n have golden scales, so apparently scale color is variable in the northern part of the range. Material in the southern part of the range (Chiapas to Honduras) has narrower, more bristle-like rhizome scales.

12. ELAPHOGLOSSUM GRATUM Elaphoglossum gratum (Fe´e) T. Moore, Index Fil. 10. 1857. Figs. 132A–C. Acrostichum gratum Fe´e, Me´m. Foug. 8: 69. 1857. Lectotype (chosen by Mickel in Davidse et al, 1995: 267). Mexico. Me´xico: Popocate´petl, Schaffner 279 (P!; isotype RB). Acrostichum callolepis Fe´e, Me´m. Foug. 8: 69. 1857. Elaphoglossum callolepis (Fe´e) T. Moore, Index Fil. 7. 1857. Type. Mexico. Galeotti s.n. (P?). Acrostichum deppei Kunze ex Christ, Monogr. Elaph. 18. 1899. Type. Mexico. Schaffner s.n. [76]. (P!).

Rhizomes short- to moderately creeping, to 4 mm diam.; rhizome scales linear-lanceolate, dark brown to black, lustrous, entire to only faintly toothed, ca. 5 mm long; fronds clumped, 8– 20(–30) ⫻ 0.8–1.2(–3) cm; phyllopodia distinct to indistinct; stipes 1 ⁄3–2⁄3 the frond length, with scales linear-lanceolate and darktoothed or dark-tipped, spreading, 3 mm long, some more dissected, like stellate hairs, appressed; blades narrowly elliptic,

292

ELAPHOGLOSSUM

chartaceous, apices acute to acuminate, bases narrowly cuneate; veins obscure, free, simple or once-forked, 0.6–1 mm apart, at 55–65⬚ angle to costa; hydathodes absent; blade scales evenly and liberally distributed adaxially but not overlapping, linearlanceolate, 1–1.5 mm long, with short hair-teeth, on abaxial surfaces less abundant, often folded lengthwise and curved; fertile fronds generally longer than the sterile and very long-stiped (ca. 5 ⁄6 the length), apices more acute or obtuse, bases more broadly cuneate, scales on abaxial costae but not among the sporangia.

Acrostichum salvinii Baker ex Hemsl., Biol. Cent.-Amer., Bot. 3: 688. 1886. Type. Guatemala. Volca´n de Fuego, Godman & Salvin 54226 (K!).

Rhizomes short- to long-creeping, 2–4 mm diam.; rhizome scales lanceolate, concolorous lustrous brown, to 5 mm long, entire or with a few weak, short hairs, sometimes deciduous, leaving naked rhizomes; fronds approximate to slightly spaced, to 60 cm long, but usually half that long, 1.5–4 cm wide; phyllopodia distinct; stipes 1⁄4–1⁄2 the frond length, naked or with minute, reduced, appressed scales; blades lanceolate to nearly elliptic, coriaceous, apices rounded to acuminate, bases narrowly cuneate; veins obscure, free, simple or once-forked, ca. 1 mm apart, at ca. 70⬚ angle to costa; hydathodes absent; blade surfaces glabrous or with minute stellate trichomidia, especially on the abaxial surfaces; fertile fronds slightly longer than the sterile fronds, the stipes longer (1⁄2–3⁄4 the length) and the blades narrower (1–1.9 cm), scales absent.

Distribution. Terrestrial in cloud forests; (1750–)2450–3300 (–3800) m. Mexico; Guat, CR. Selected Specimens Examined. DF (Ventura A. 5, NY). Hgo (Sa´nchez Mejorada 707, MEXU). Jal (Gonza´lez Tamayo s.n., 14 Oct 1984, XAL). Me´x (Pringle 4487, NY). Mich (Garcı´a Lo´pez 3104, MEXU; Rzedowski 46151, MEXU). Mor (Lyonnet 3465, MEXU). Oax (Mickel 7437, NY). Pue (Reeves 6192, ASC). Ver (Nee 23129, NY).

Elaphoglossum gratum is closely allied to E. hartwegii and E. pringlei. It is somewhat like E. muelleri, which is also scaly, terrestrial, and of high elevations, but can be distinguished by its linear-lanceolate, coarsely toothed blade scales. Elaphoglossum muelleri has blade scales with little body to them and the rhizome scales are tan to orange. Elaphoglossum gratum also resembles E. petiolatum, which has entire rhizome scales, and the adaxial blade scales are usually substellate rather than linear-lanceolate. Elaphoglossum gratum is close to E. hartwegii in its heavier scaliness on adaxial blade surfaces, dark entire rhizome scales, and tendency for the rhizomes to creep, but E. hartwegii has longer creeping rhizomes, and blade scales that are more deltatelanceolate and subentire, broader with darker middle, and pale margins rather than the usual white. Both may have the resin dots on the abaxial blade surfaces.

Distribution. Epiphytic, rarely terrestrial, in wet montane forests; 500–2500 m. Mexico; Guat, Bel, Hond, Salv. Selected Specimens Examined. Chis (Breedlove 25345, 25365, NY). Hgo (Moore 5303, US). Oax (Hallberg 1580, NY; Mickel 4726, NY). Pue (Riba et al. 304, MEXU, NY—atypical in having stipes only 1⁄10 the sterile frond length and the fertile fronds narrower and longer than the sterile). Qro (Calzada 595, XAL; Carranza 3867, NY; Dı´az B. & Carranza 6559, NY). Ver (Ibarra Manriquez 3012, UAMIZ). Unverified, Doubtful, or Mistaken Reports. Tam (Lof, 1980, f. 61, drawing of E. sp. appears to be E. guatemalense, but no specimen cited, not verified).

13. ELAPHOGLOSSUM GUATEMALENSE

Under this name we have placed those essentially glabrous specimens that have slender, creeping rhizomes and short, lanceolate, lustrous, brown rhizome scales. There may in fact be more than one element here, as some specimens seem especially narrow with acuminate apices, some broader with bluish green blades and acuminate apices; however, most specimens have obtuse to acute apices.

Elaphoglossum guatemalense (Klotzsch) T. Moore, Parker’s Cat. 1858. Figs. 131B, 138F–H, J, K.

14. ELAPHOGLOSSUM HARTWEGII

Acrostichum guatemalense Klotzsch, Allg. Gartenzeitung 23: 66. 1855. Type. From cultivation in Potsdam, Germany, grown from spores of unknown locality in Guatemala (B?).

Elaphoglossum hartwegii (Fe´e) T. Moore, Index Fil. xvi, 10. 1857. Figs. 135G, H, J, K.

ELAPHOGLOSSUM HUACSARO Acrostichum hartwegii Fe´e, Me´m. Foug. 2: 53, pl. 9, f. 2. 1845. Type. Colombia. Bogota´, Hartweg 1486 (G-DEL; isotypes GH!, K!, RB).

Rhizomes long-creeping, 1–1.5 mm diam.; rhizome scales linear-lanceolate, reddish brown to black, lustrous with markedly paler margins and sagittate bases, entire, ca. 3 mm long; fronds often 1 cm apart, 7–21 ⫻ 1–2.1 cm; phyllopodia distinct; stipes 1⁄2–2⁄3 the frond length, with scales mostly 1–2 mm long, brown with pale margins, appressed to slightly spreading, entire to slightly toothed; blades narrowly elliptic, chartaceous to subcoriaceous, acuminate, acute, or obtuse at apices, cuneate at bases; veins inconspicuous, 0.8–1 mm apart, at 60–70⬚ angle to costa; hydathodes present but inconspicuous; blade scales of the adaxial surface generally abundant but not overlapping, lanceolate to narrowly deltate, peltate, brown with pale margins, the abaxial surfaces often essentially glabrous; fertile fronds somewhat taller than the sterile, with longer stipes (2⁄3–3⁄4 the frond length), scales sparse abaxially on the costae, not present among the sporangia.

Distribution. Epipetric and terrestrial on shaded ledges in pine or fir forests; 3000–4100 m. Mexico; Guat, CR; Col, Ven, Ec, Peru, Bol, Chile. Selected Specimens Examined. Chis (Breedlove 24315, DS, NY; Matuda 2314, US). DF (Beaman 2800, MSC). Jal (McVaugh 10074, MICH, NY). Me´x (Beaman 2823, MEXU, MSC, UC, US; Matuda 27573, MEXU, UC, US; Rzedowski 23469, LL, MEXU, TEX, US, WIS). Mich (Leavenworth & Hoogstraal 1190, MO). Pue (Herna´ndez X. & Sharp X-168, MEXU). Ver (Nee & Diggs 24844, F, NY, XAL).

Specimens in Mexico and Guatemala tend to have somewhat broader leaf blades than those from South America, but further study is necessary in this complex before species distinctions can be made with confidence. Mickel and Beitel (1988) treated E. hartwegii as a synonym of E. mathewsii (Fe´e) T. Moore, but now, with more plentiful material of this complex at hand from Peru (cf. J. T. Mickel, Elaphoglossum, in Pteridophyta of Peru, IV. Fieldiana, Bot., n.s., 27: 111-166. 1991), we believe the two to be distinct. More study is needed to determine fully the ranges of both species. See E. gratum for discussion of the hybrid between that species and E. hartwegii.

293

15. ELAPHOGLOSSUM HERMINIERI Elaphoglossum herminieri (Bory ex Fe´e) T. Moore, Index Fil. xvi. 1857. Figs. 137D, E. Acrostichum herminieri Bory ex Fe´e, Me´m. Foug. 2: 43, pl. 11. 1845. Lectotype (chosen here). Guadeloupe, L’Herminier s.n. (P!; isolectotypes GH, RB).

Rhizomes short-creeping, to ca. 8 mm diam.; rhizome scales linear, concolorous bright orange, lax, 15–30 mm, entire or rarely with irregular teeth; fronds approximate, to 200 ⫻ 4 cm, usually pendent; phyllopodia absent; stipes essentially absent (0–2 cm); blades linear-elliptic, apices acuminate, bases narrowly cuneate, coriaceous; veins obscure, free, simple or once-forked, ca. 1 mm apart, running at ca. 70⬚ to costa; hydathodes absent; adaxial surfaces essentially glabrous; abaxial surfaces sparsely clothed with minute, dissected, orange scales, to nearly glabrous; fertile fronds rare, lanceolate, 10–25 cm, only 1⁄5–1⁄3 the sterile frond length, glabrous, stipes 1⁄5–1⁄4 the frond length; 2n⫽216 (Trin).

Distribution. Epiphytic in wet lowland forests; 0–600 m. Mexico; Guat, Bel, Hond, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec. Specimen Examined. Chis (Castillo C. et al. 3949, XAL).

Elaphoglossum herminieri probably has the longest fronds of any Elaphoglossum, reaching 2 m in length. It is one of the few species found at low elevations. The bluish green cast to the leathery fronds and the long, slender, orange rhizome scales are distinctive.

16. ELAPHOGLOSSUM HUACSARO Elaphoglossum huacsaro (Ruiz) Christ, Monogr. Elaph. 96. 1899. Figs. 135Q–S. Acrostichum huacsaro Ruiz, Mem. Calaguala 57. 1805. Type. Peru. “Habitat in Andinum montibus ad Sanatolmas et Tarmac provinciae,” Ruiz 54 (B!; isotype BR, photos F, US).

Rhizomes compact, often 5–10 cm long, 3–5 mm diam., apices ascending; rhizome scales lanceolate, concolorous, castaneous to

294

ELAPHOGLOSSUM

black, 1–3 mm long, lustrous, entire, each often with a tortuous hair-like tip; fronds 17–34 cm long, approximate; phyllopodia present but obscured by scales; stipes 1⁄4–1⁄3 of sterile frond length, orange to brown; stipe scales ca. 1 mm long, mostly appressed, some spreading, lanceolate, dark to pale, the larger ones often dark-tipped; sterile blades (6–)15–24 ⫻ 0.8–1.6 cm, linear, chartaceous to subcoriaceous, bases narrowly cuneate, apices obtuse to rounded; veins obscure, ca. 1 mm apart, at 60–75⬚ angle to costa; hydathodes absent; blade scales reduced to resinous dots on both surfaces, costal scales ca. 1 mm long, lanceolate, spreading, and scales along adaxial lamina margins 0.2–0.6 mm long, appressed, ovate, entire; fertile fronds far exceeding the sterile in length, stipes 2⁄3 of frond length, nearly as long as the entire sterile leaf, blades narrow, (6–)8–11 mm wide, intersporangial scales absent.

absent; adaxial blade surfaces subglabrous with sparse substellate scales, costa with sparse, short, dark, arachnidoid scales and occasional inconspicuous, pale, peltate, fimbriate scales; abaxial surfaces with sparse, orange, stellate hairs, costae with sparse, dark-sclerotic to pale arachnidoid scales; fertile fronds ca. equal to sterile in length but narrow, linear, 3–4 mm wide with inrolled margins, adaxial surfaces with abundant, orange, round, peltate, fimbriate scales ca. 0.5 mm across (incl. teeth), adaxial costae with similar scales and scattered, small, dark, sclerotic, arachnidoid scales, abaxial costae with only dark arachnidoid scales.

Distribution. Epiphytic in wet montane forest; 800–2200 m. Known only from the type collection.

Distribution. Terrestrial or epipetric on steep roadbanks in pineoak and wet forests. 1700–2200 m. Mexico; Guat, CR; Jam, Hisp; Col, Ven, Ec, Peru, Bol. Specimens Examined. Chis (Breedlove 22412, ENCB, NY, 23167, 32336, DS; Lloyd 4064, ENCB).

Elaphoglossum huacsaro is distinct by the linear fronds with resinous dots abaxially and small marginal scales adaxially, obtuse blade apices, ascending rhizomes, and terrestrial habitat.

Unverified, Doubtful, or Mistaken Reports. Chis (Pe´rez-Farrera 435, UAMIZ, UNICACH, cited by Pe´rez-Farrera, Garcı´a, Riba, & Lo´pez-Molina, Amer. Fern J. 93: 152, 2003, but not verified.

Elaphoglossum ipshookense is closely allied to E. tectum, resembling that species in blade shape, dark rhizome scales, and stellate hairs abaxially, but differing from it in smaller size and in lacking the distinctive round, peltate scales on stipes, rachises, and adaxial blade surfaces of the sterile fronds. The peltate scales on the fertile fronds confirm the relationship with E. tectum. This is one of the few species in Elaphoglossum in which the scales of the fertile fronds differ from those of the sterile fronds.

17. ELAPHOGLOSSUM IPSHOOKENSE Elaphoglossum ipshookense Mickel, Mem. New York Bot. Gard. 46: 175. 1988. Type. Mexico. Oaxaca: Distrito Mixe, N slope of Cerro Zempoalte´petl, trail from Totontepec to La Candelaria, Mickel 4748 (NY!). Figs. 129H, J, K. Rhizomes short-creeping, compact, horizontal, 1.2–2 mm diam.; rhizome scales linear-lanceolate, castaneous to dark reddish brown, sclerotic, lustrous, 2–3 mm long, subentire with sparse teeth; fronds clumped to 2 mm apart, 19–30 ⫻ 1–1.4 cm; phyllopodia distinct, 5–7 mm long; stipes 1⁄6–1⁄4 the frond length, with scales sparse, appressed, sclerotic, dark reddish brown, arachnidoid; blades linear-elliptic, chartaceous, bases narrowly cuneate, apices long-acuminate; veins obscure, free, simple or 1-forked, mostly 0.7–1 mm apart, at ca. 75⬚ angle to costa; hydathodes

18. ELAPHOGLOSSUM JALISCANUM Elaphoglossum jaliscanum Mickel in McVaugh, Fl. Nov.-Gal. 17: 271. 1992. Type. Mexico. Jalisco: 29 km S of Puerto Vallarta, then 2 km along dirt road E to Zimapa´n, Stevenson 570 (NY!). Figs. 134J–L. Similar to E. piloselloides in the spatulate blades, orange subulate scales, flattened stipes, and relatively inconspicuous hydathodes, but distinct in its much larger size (fronds 12–14 ⫻ 1–1.3 cm instead of 2–4(–8) ⫻ 0.4–1 cm), the blades 7–8.5 cm long (vs.1–3.5 cm), the stipes ca. 1⁄5 the length of the sterile fronds (vs. 1⁄2–3⁄4), and the scales of the stipes and blades ca. 3 mm long (vs. 1.5–2); fertile fronds unknown.

ELAPHOGLOSSUM LATUM

295

Distribution. Epipetric in arroyo; 100 m. Known only from the type collection.

Distribution. Epiphytic in pine-oak cloud forests; 2300–2500 m. Mexico; Guat, Hond, Salv.

Elaphoglossum piloselloides, the most similar species, is widespread from Sinaloa to Chile but differs in its remarkably uniform blades (size and shape).

Specimens Examined. Oax (Breedlove 65911, 65926, CAS; Mickel 5565, NY).

19. ELAPHOGLOSSUM LANCEUM Elaphoglossum lanceum Mickel, Amer. Fern J. 69: 101. 1979. Type. Mexico. Oaxaca: Distrito Mixe, SE shoulder of Cerro Zempoalte´petl, below Patio de Arena, Mickel 4900 (NY!; isotypes F!, NY!, UC!, US!). Figs. 130D–F. Rhizomes short-creeping to erect, 1 mm diam.; rhizome scales linear-lanceolate, reddish brown, 7 mm long; fronds clumped, 9–34 ⫻ 1.2–2.7 cm; phyllopodia absent; stipes 1⁄2 the frond length, with scales red-brown, strongly spreading, subulate, to 6 mm long; blades lanceolate to linear-lanceolate, membranaceous, bases cuneate, apices acuminate, margins entire to crenulate; veins conspicuous, free, simple or once-forked, 2–3 mm apart, at 35–40⬚ angle to costa; hydathodes evident; blade scales orangebrown, subulate, 3–5 mm long, sparse; fertile fronds shorter than the sterile, ca. 16 cm long, bases cuneate, blade apices acuminate to subcaudate, blades shorter in proportion to the length of the sterile blades (to 13 cm long, 4.8 cm wide), stipes 2⁄3–4⁄5 length, scales mixed with the sporangia.

Elaphoglossum lanceum is known only from a few collections, most of which have fronds only 12–20 cm long and distinctly lance-shaped. It resembles E. smithii (Baker) Christ of the West Indies, but E. lanceum is readily distinguished by its small size, thin texture, somewhat more blade scales, and highly fenestrate spore crests (vs. imperforate crests in E. smithii).

20. ELAPHOGLOSSUM LATUM Elaphoglossum latum (Mickel) Atehortu´a in Mickel, Novon 2: 377. 1992. Figs. 133H, J. Elaphoglossum apodum (Kaulf.) Schott ex J. Sm. var. latum Mickel, Amer. Fern J. 69: 100. 1979. Type. Guatemala. Alta Verapaz: along Rte 5 between Semococh and La Laguna on road to Chajmarjic, Steyermark 46368 (F!; isotype US!).

Rhizomes 4–6 mm diam., short-creeping; rhizome scales 2.5 cm long, linear, orange- to red-brown, lustrous, entire; fronds 25–48 cm long, approximate; phyllopodia 0.5–1 cm long; stipes 0.5–1 cm long, nearly absent, less than 1⁄30 of sterile frond length, stout, stramineous to brown; stipe scales 3–6 mm long, spreading, subulate, orange, lustrous, entire, the stipes also bearing minute glandular hairs; sterile blades 24–47 ⫻ 4.5–5.7 cm, oblanceolate, papyraceous to coriaceous, bases narrowly (or rarely broadly) cuneate, apices long-acuminate to caudate; veins ca. 1 mm apart, at 60–70⬚ angle to costa; hydathodes absent; blade scales 2–3 mm long, subulate, orange, lustrous, especially concentrated on the costae and margins, scattered on abaxial blade surfaces, entire, very sparse on adaxial surfaces; lamina also with minute, erect, glandular hairs; fertile fronds much smaller than the sterile, stipes 1 ⁄15–1⁄5 the frond length, with long, linear, orange scales on the abaxial costae, intersporangial scales absent.

296

ELAPHOGLOSSUM

Distribution. Epiphytic, rarely epipetric in secondary growth forests, dry forests, and rain forests; 280–550 m. Mexico; Guat, Bel, Hond, Nic, CR, Pan; Hisp; Col, Ec. Specimens Examined. Chis (Breedlove 48328, CAS, 57355, CAS, ENCB; Martı´nez S. 25922, MEXU).

Distribution. Epiphytic in wet pine-oak forests; 1800–2800 m. Mexico. Specimens Examined. Chis (Breedlove 15292, 22014, 22093, 23036, NY; Laughlin 166, NY; Shilom Ton 694, NY). Oax (Diggs 3886, NY; Lorence 4207, NY; Mickel 4124, 4362, 6521, NY; Torres C. 10703, NY).

Elaphoglossum latum has fronds that are virtually sessile (stipes to 1 cm) and oblanceolate with orange subulate scales. It resembles E. apodum of the West Indies, Venezuela, and Nicaragua, but E. latum is distinct by its broader blades (4.5–5.7 cm vs. 0.9– 4.4 cm).

Elaphoglossum leebrowniae is distinct in the E. sartorii complex by its rhizome scales large, spreading, tinged with black, and fertile fronds about half as broad as the sterile.

21. ELAPHOGLOSSUM LEEBROWNIAE

Elaphoglossum leonardii Mickel, Brittonia 32: 336. 1980. Type. Mexico. Oaxaca: Distrito Ixtla´n, Cerro de Malacate, N of Capulalpan, ca. 8 km E of Ixtla´n de Jua´rez, Mickel 5296 (NY!; isotypes MEXU!, UC!, NCU (as Leonard 4069), US!). Figs. 130L–N.

Elaphoglossum leebrowniae Mickel, sp. nov. Type. Mexico. Chiapas: 3 mi N of San Cristo´bal las Casas on road up Cerro Zontehuitz, Mickel 1230 (holotype NY!). Figs. 125A–C. Ab E. sartorii squamis rhizomatum grandioribus nigrescentibus differt. (Dedicated to the senior author’s research assistant, Leone (Lee) Brown, who did so much to bring this manuscript to completion.)

Rhizomes compact, stout, 5–8 mm diam.; rhizome scales linear-lanceolate, mostly 15–20 ⫻ 2 mm, orange-tan, tinged with lustrous black, with attenuate apices and irregular hair-like lateral projections; fronds 30–60 ⫻ 3.7–6.5 cm, 0.5–1.5 cm distant; phyllopodia distinct, ca. 1 cm long; stipes 1⁄3–2⁄5 the frond length; blades narrowly elliptic, bases cuneate at 25–50⬚ angle, apices acute to acuminate, coriaceous; veins obscure, free, simple to 2forked; hydathodes absent; adaxial surfaces glabrous or with sparse dark trichomidia; abaxial surfaces with scattered dark brown stellate trichomidia, mostly 0.2–0.4 mm; fertile fronds slightly shorter to slightly longer than the sterile, stipes ca. 1⁄2 the frond length, the blades 2⁄5–3⁄4 the width of the sterile, intersporangial scales absent.

22. ELAPHOGLOSSUM LEONARDII

Rhizomes short-creeping, 2–3 mm diam.; rhizome scales linear, lustrous, maroon, ca. 2 mm long, with short teeth; fronds clumped, 7–14 ⫻ 1.5–2.8 cm; stipes 1⁄2–3⁄5 the frond length, with scales orange to maroon, subulate, widely spreading, ca. 1 mm long, and with minute erect glandular hairs; phyllopodia absent; blades ovate to lanceolate-oblong, chartaceous, bases rounded, apices obtuse to acute; veins evident, free, simple or once-forked, 1.5–2 mm apart, at 60–70⬚ angle to costa; hydathodes distinct; blade scales subulate, orange to maroon, generally abundant on abaxial surfaces, sparse to absent adaxially, 1–2 mm long; fertile fronds shorter than the sterile fronds (2⁄3 to nearly as long), their blades much smaller (1⁄3–1⁄2 the dimensions of the sterile blade), often folded at the costa before maturity, scales scattered among the sporangia.

ELAPHOGLOSSUM LINDENII

297

Distribution. Epipetric in cloud forest; 2700 m. Known only from the type collection.

Distribution. Epiphytic in wet pine-oak woods; 850–1700 m. Mexico.

Elaphoglossum leonardii closely resembles E. lindenii, from which it differs in having the blade apices acute to obtuse rather than cuspidate.

Specimens Examined. Chis (Heath & Long 1045, MEXU). Oax (Campos 3103, MEXU; McVaugh 22254, ENCB, NY; Mickel 5046, NY; Rzedowski s.n., 22 Sep 1965, ENCB).

23. ELAPHOGLOSSUM LEPIDOPODUM Elaphoglossum lepidopodum Mickel, Brittonia 32: 336. 1980. Type. Mexico. Oaxaca: Distrito Juquila, 24–25 km n of San Gabriel, wooded banks by stream with Pinus strobus and oaks, Mickel 6175 (NY!; isotypes ENCB!, MEXU!, UC!). Figs. 124J, K, 138A–E. Rhizomes short-creeping, 2–5 mm diam.; rhizome scales lanceolate to deltate-lanceolate, brown with dark lustrous center with age, to 4 mm long, with sparse irregular marginal processes; fronds slightly spaced, 18–43 cm; phyllopodia distinct, ca. 1 cm long; stipes 1⁄8–1⁄5(–1⁄3) the frond length, scaly with tan, spreading, lanceolate to ovate scales with isodiametric cells, distally ovate to lanceolate, brown to orange-tan with irregular short processes, and minute black, appressed, round to arachnidoid scales; blades narrowly elliptic, firm, 1.5–2.8 cm wide, longacuminate at both ends to obtuse at apices; veins scarcely visible, free, simple, or once-forked, often slightly expanded laterally at tips, 1 mm apart, at ca. 70–80⬚ angle to costa; hydathodes absent; adaxial blade surfaces glabrous; abaxial surfaces with minute black to brown stellate trichomidia; fertile fronds slightly longer than sterile and only slightly narrower, stipes 2⁄5 the frond length, with scattered scales on costal bases, glabrous adaxially.

This species is probably closest to E. guatemalense, agreeing with that in its slender rhizomes, lustrous rhizome scales (at least with age), irregularly toothed stipe scales, distinct phyllopodia, reduced black stellate blade scales, and distribution on the Pacific slope, but E. lepidopodum differs in its very slender fronds, thin blade texture, and scaly stipes.

24. ELAPHOGLOSSUM LINDENII Elaphoglossum lindenii (Bory ex Fe´e) T. Moore, Index Fil. 360. 1862. Figs. 131F–H. Acrostichum lindenii Bory ex Fe´e, Me´m. Foug. 2: 48, pl. 18, f. 3. 1845. Type (isolectotypes chosen by Mickel in Stolze, 1981: 226). Mexico. Veracruz: Orizaba, Galeotti 6263 (G-DEL; isotypes BR–2 sheets, P!). Acrostichum pumilum M. Martens & Galeotti, Me´m. Foug. Mexique 23, pl. 2, f. 2. 1842, non Elaphoglossum pumilum H. J. Lam & Verhey ex H. J. Lam, 1945. Type. Mexico. Veracruz: Orizaba, Galeotti 6263 (BR!; isotype P!). Acrostichum venustum Liebm., Mexic. Bregn. 168 (reprint 16). 1849, non Fe´e, 1857, nec Elaphoglossum venustum T. Moore, 1857. Elaphoglossum liebmannii T. Moore, Index Fil. 360. 1862. Type. Mexico. Veracruz: Orizaba, Liebmann s.n. [Pl. Mex. 2420] (C!; isotypes P!, US!).

Rhizomes short-creeping to ascending, ca. 3 mm diam.; rhizome scales linear, maroon, to 8 mm long, with short teeth; fronds clumped, 7–40 ⫻ 1.5–4 cm; phyllopodia absent; stipes 1⁄2–3⁄4 the frond length, with scales orange to maroon, subulate, widespreading, 2–3 mm long, also with a few minute erect glandular hairs; blades ovate to ovate-lanceolate, chartaceous, bases rounded or subcordate, apices cuspidate; veins evident, free, simple or once-forked, ca. 2 mm apart, at 60–70⬚ angle to costa; hydathodes distinct; blade scales subulate, orange to maroon, abundant on abaxial surfaces, few adaxially, especially spreading from the margins, 2–3 mm long; fertile fronds usually slightly shorter than the sterile, the blades more elliptic, to 4.2 ⫻ 2.4 cm, stipes to 4⁄5 the frond length, apices obtuse, blades often folded at the costa, scales on the abaxial midvein but not among the sporangia.

298

ELAPHOGLOSSUM

Distribution. Terrestrial or epipetric, rarely epiphytic, in cloud forests; 2800–3600 m. Mexico; Guat, CR, Pan; Col, Ven, Ec, Peru, Bol, Chile. Selected Specimens Examined. Chis (Breedlove 22065, DS, 22806, 24316, NY). Gro (Hinton 14295, NY; Lorea 4211, IEB). Jal (McVaugh 10060, US p.p.). Me´x (Rzedowski 23278, NY). Oax (Mickel 9652, NY; Somers 345, NY). Ver (Galeotti 6263, BR). Unverified, Doubtful, or Mistaken Reports. Mich (reported by Dı´az-Barriga & Palacios-Rios, 1992, referring to Ibarra, 1983, but not verified and doubtful).

The plants are usually small but the species is highly variable. It is distinguished by the blade shape, the generally cuspidate blade tips, and the folded fertile blades. It may be mistaken for E. monicae, but the latter has lighter colored blade scales, acuminate blade apices, and echinate spores. Very small specimens of E. lindenii closely resemble plants of E. piloselloides, but the former can be distinguished by blade shape, more evident hydathodes, and higher elevation (confined to Pacific slope).

25. ELAPHOGLOSSUM LONCHOPHYLLUM Elaphoglossum lonchophyllum (Fe´e) T. Moore, Index Fil. 360. 1862. Figs. 126C–E. Acrostichum lonchophyllum Fe´e, Me´m. Foug. 8: 68. 1857. Type. Mexico. Veracruz: Huatusco, in 1854, Schaffner 19 (RB). Elaphoglossum acutissimum Christ, Bull. Herb. Boissier, se´r. 2, 5: 730. 1905. Type. Mexico. Chiapas: San Pablo, Mu¨nch 168 (P?; isotypes DS!, US!).

Rhizomes short-creeping, to 3 mm diam.; rhizome scales linearlanceolate, lustrous, concolorous, brown, somewhat resinous, to 4(–20) mm long, entire; fronds clumped, 24–74 ⫻ 1.7–4.5 cm; phyllopodia indistinct; stipes 1⁄5–1⁄3 the frond length, sparsely clothed with minute, stellate, appressed hairs; blades narrowly elliptic, very thin, bases cuneate, apices long acuminate; veins evident, free, simple or once-forked, 1.5–2 mm apart, at 60–70⬚ angle to costa; hydathodes absent, but vein tips expanded laterally, sometimes uniting; blade scales absent except for a few minute, stellate, reddish, resinous hairs, mostly on the abaxial blade surfaces; fertile fronds about equal to the sterile in length, but with longer stipes (ca. 1⁄2 the fertile frond length) and narrower blades, scales absent from abaxial blade surfaces.

Distribution. Epiphytic in montane rain forests; 450–2500 m. Mexico; Guat, Hond, Nic, CR, Pan; Hisp, PR. Selected Specimens Examined. Chis (Breedlove 31627, DS, 32218, NY, 32994, DS). Gro (Lorea 1004, FCME; Nu´n˜ez 1040, NY). Oax (Mickel 1003, 1084, 5706, NY). Ver (Bourgeau 2888, K, NY; Mu¨ller 435, NY).

This is distinguished from other species by its very thintextured blades and laterally expanded vein ends. Specimens from Chiapas seem to have shorter, more resinous rhizome scales than do those of Guatemala, but since both conditions are found in Oaxaca, it is presumed to be a variable character in the species. Breedlove 32218, NY, has rhizome scales to 20 mm long. From E. decursivum, E. lonchophyllum differs in having longer stipes and blades not long-decurrent.

26. ELAPHOGLOSSUM MANANTLANENSE Elaphoglossum manantlanense Mickel in McVaugh, Fl. Nov.Gal. 17: 272. 1992. Type. Mexico. Jalisco: E of [Rancho] Manantla´n, about 15 mi SSE of Autla´n, Wilbur & Wilbur 1971 (US!; isotypes MEXU!, MICH!). Figs. 136F–H, J–L. Rhizomes short-creeping, horizontal, 2–3 mm diam.; rhizome scales linear-lanceolate, castaneous, lustrous, 3–4 mm long, inconspicuously toothed; fronds clumped, 20–33 ⫻ 1.5–2.3 cm; phyllopodia distinct; stipes 1⁄10–1⁄5 the frond length, densely scaly, the scales spreading, orange-tan, concolorous or with dark central streak, 2–3 mm long; blades narrowly elliptic, narrowly cuneate at bases, acuminate at apices; veins obscure, ca. 1 mm apart, at 80⬚ angle to costa; hydathodes absent; blade scales scattered to overlapping, lanceolate, 1–2 mm long (to 3 mm long on the costae abaxially), orange-tan (on adaxial surfaces becoming white with age, with long hair-teeth); fertile fronds longer than the sterile, the stipes ca. 3⁄4 the frond length, the blades linear, ca. 6 mm wide, usually inrolled to appear 3–4 mm wide, the scales lanceolate abaxially along the costae, those of the adaxial surfaces much reduced.

ELAPHOGLOSSUM MONICAE

Distribution. On vertical, moss-covered walls of gorges in mixed pine-hardwood forests; 1550–1950 m. Mexico. Specimens Examined. Gro (Lorea 3136, 3144, 3175, all FCME; Rzedowski & McVaugh 66, ENCB, MICH). Jal (only the type collection). Oax (Mickel 6195, NY).

Elaphoglossum manantlanense resembles E. paleaceum in the scaly blades but is distinct in the castaneous, nearly edentate rhizome scales, very short stipes (1⁄10–1⁄5 the length of the frond instead of 1⁄5–1⁄4), concolorous stipe scales (or with a dark central streak), and linear fertile blades, slightly longer than the sterile.

27. ELAPHOGLOSSUM MCVAUGHII Elaphoglossum mcvaughii Mickel, Brittonia 32: 336. 1980. Type. Mexico. Michoaca´n: 8–10 mi NW and WNW of Ciudad Hidalgo, among mountains west of Cerro San Andre´s and 6–7 mi N of village of San Pedro Aguaro, McVaugh 9904 (US!; isotypes LL!, MICH!). Figs. 137A–C. Rhizomes long-creeping, ca. 3 mm diam.; rhizome scales ovate to ovate-lanceolate, tan to orange-tan, at least some with lustrous black or dark brown streaks, mostly 3–5 mm long, generally entire, scales somewhat deciduous; fronds 1–2 cm apart, 21–40 ⫻ 1.5–2.9 cm; phyllopodia distinct, 1–2 cm long; stipes 1⁄3–1⁄2 of the frond length, with scattered ovate scales similar to those of the rhizomes, entire or with occasional weak marginal hairs; blades narrowly elliptic, apices acute to acuminate, bases narrowly cuneate; coriaceous; veins obscure, free, simple or once-forked, ca. 1 mm apart, at ca. 70⬚ angle to costa; hydathodes absent; blade surfaces subglabrous, with sparse, minute, stellate trichomidia and occasional broader scales on the costae, especially on the abaxial surfaces; fertile fronds longer than the sterile fronds, the stipes longer (1⁄2–2⁄3 of the frond length) and the blades slightly narrower, scales absent from abaxial surfaces.

299

Distribution. On rocks in ravines in pine-fir and pine-oak forests; 2250–3000 m. Mexico; Guat, Salv. Specimens Examined. Chis (Breedlove 65796A, CAS). Mich (Dı´az B. & Zamudio 2671, IEB; Rzedowski 46175, IEB; Santos Martı´nez 1999, ENCB, IEB). Oax (Lorence 4619, ENCB, MEXU; Mickel 6614, NY; Pe´rez Cruz C12, MEXU).

Elaphoglossum mcvaughii is similar to E. affine in the bicolorous rhizome scales, creeping rhizomes, rhizome scale morphology (entire with occasional weak teeth or marginal hairs), fertile fronds longer than the sterile, blade texture, and indument; however, the former species has acuminate rather than acute to rounded blades and generally greater dimensions. It also resembles E. seminudum, but differs in having bicolorous rhizome scales that are more spreading (vs. appressed).

28. ELAPHOGLOSSUM MONICAE Elaphoglossum monicae Mickel in McVaugh, Fl. Nov.-Gal. 17: 273. Type. Mexico. Me´xico: Mpio. Tlalmanalco, Can˜ada de San Rafael, near San Rafael, Rzedowski 26695 (NY!; isotype ENCB!). Figs. 134O–Q. Rhizomes short-creeping to ascending, ca. 3 mm diam.; rhizome scales linear, maroon, to 8 mm long, with short teeth; fronds clumped, 7–40 ⫻ 1.5–4 cm; phyllopodia absent; stipes 1⁄2–2⁄3 the length of the sterile fronds; stipe scales orange to maroon, subulate, wide-spreading, 2–3 mm long; stipes also with a few minute erect glandular hairs; blades oblong-elliptic, 3–6 times as long as wide, rounded to broadly cuneate at bases, acute to acuminate or sometimes cuspidate at apices; veins evident, ca. 2 mm apart, at 60–70⬚ angle to costa; hydathodes distinct; blade scales subulate, orange to maroon, sparse on the abaxial surfaces, rare adaxially, spreading from the margins, 2–3 mm long; fertile fronds shorter than the sterile, the stipes 3⁄4–4⁄5 the length of the fertile fronds, the blades ovate, rounded to truncate at bases, 1.7–5 ⫻ 0.8–1.5 cm, the apices obtuse, the blades often folded at the midveins, scales present on the midveins abaxially, but lacking among sporangia.

300

ELAPHOGLOSSUM

surfaces, ca. 1 mm long, occasionally with a few linear-lanceolate scales on the abaxial costae; fertile fronds usually exceeding the sterile and with longer stipes (nearly 1⁄2 the length) and narrower blades (to 0.8 cm wide), stellate hairs on abaxial costae, but absent among sporangia.

Distribution. Terrestrial or epipetric, in pine or pine-fir forests; 2150–3300 m. Mexico. Selected Specimens Examined. DF (Rzedowski 27242, NY). Dgo (McVaugh 23599, MICH). Gro (Rzedowski & McVaugh 250, ENCB, MICH). Jal (McVaugh 10060, ENCB, LL, MICH, NY, TEX, US p.p., 26140, NY). Me´x (Hinton 7221, LL, MO, NY, TEX, US; Purpus 1585, NY). Mor (Lyonnet 1407, US). Sin (Ownbey & Ownbey 1965, US). Ver (Dorantes 566, MEXU).

This species is closest to E. setigerum, which is distinguished by its sterile blade length/width ratio of 10–15:1 (vs. 3–6:1), and more southern distribution (Gro, Oax, Chis). Elaphoglossum monicae closely resembles E. lindenii in the sometimes cuspidate blade apices, subulate scales, glands on the stipes, and red-brown rhizome scales, but in E. monicae the blades are more oblongelliptic and the spores are echinate-reticulate, 37–49 (avg. 43) ␮m long, rather than with fenestrate crests and 23–33 (avg. 29) ␮m long.

29. ELAPHOGLOSSUM MUELLERI Elaphoglossum muelleri (E. Fourn.) C. Chr., Index Filic. 311. 1905. Figs. 129L–O. Acrostichum muelleri E. Fourn., Mexic. Pl. 1: 68, t. 1, f. 2. 1872. Type. Mexico. Veracruz: Ulnapa, Mu¨ller 41 quinter (P!, photo NY!). “Mu¨ller 41 4parte” (NY) is from Orizaba, aserradero de Santa Cruz, 20 Aug 1853. Acrostichum intermedium Fe´e, Me´m. Foug. 8: 69. 1857, non Elaphoglossum intermedium Brack., 1854. Type. Mexico. Veracruz: Huatusco, Schaffner 22 (P!, RB). Acrostichum araneosum D. C. Eaton, Proc. Amer. Acad. Arts 22: 461. 1887. Elaphoglossum araneosum (D. C. Eaton) C. Chr., Index Filic. 303. 1905. Type. Mexico. Jalisco: Rı´o Blanco, Palmer 333 (YU!; isotypes LE!, NY!, US!).

Rhizomes short-creeping, to 5 mm diam.; rhizome scales dull orange, ovate-lanceolate to nearly deltate, ca. 2 mm long, entire or weakly toothed; fronds clumped, 15–33 ⫻ 1.2–2.5 cm; phyllopodia absent; stipes 1⁄5–1⁄3 the frond length, with scales orange, few, linear-lanceolate or mostly stellate and hair-like, somewhat deciduous with age; blades narrowly elliptic, chartaceous, bases narrowly cuneate, apices acuminate; veins visible but not conspicuous, free, simple to 2-forked, 0.4–0.7 mm apart, at ca. 70⬚ angle to costa; hydathodes absent; blade scales appearing as stellate hairs with virtually no scale body on both adaxial and abaxial

Distribution. Terrestrial or epipetric in dry to moist oak and pine-oak forests; (300–) 900–2600 m. Mexico; Guat, Hond, Nic, CR. Selected Specimens Examined. Chis (Breedlove 23083, DS). Col (Goldsmith 41, UC). DF (Lyonnet 891, MEXU). Gro (Hinton 14882, NY). Gto (Ventura & Lo´pez 7446, MEXU). Hgo (Matuda 32531, MEXU). Jal (Pringle 2590, MEXU, MSC, NY). Me´x (Hinton 1811, NY). Mich (McVaugh 22693, MICH). Nay (Mexia 708, NY). Oax (Mickel 5057, NY). Pue (Nicolas s.n., 12 Nov 1909, NY). Sin (Breedlove 17056, NY). SLP (Schaffner s.n., Oct 1876, NY). Ver (Mu¨ller 41, NY). Unverified, Doubtful, or Mistaken Reports. Qro (Ferna´ndez 3560, cited as E. pilosum [E. muelleri] by Dı´az-Barriga & Palacios-Rios, 1992, but not verified and possibly E. petiolatum).

This species is quite variable in scaliness and plant size. It can be distinguished from Elaphoglossum petiolatum and E. gratum by its orange rhizome scales. It is most abundant in Mexico and only doubtful south of Guatemala. Elaphoglossum muelleri has generally been treated under the name of E. pilosum (Humb. & Bonpl. ex Willd.) T. Moore, the type of which is from Venezuela; it has narrower, nonciliate scales on the rhizomes and stipe bases. Reports of E. pilosum in Mexico (e.g., Conzatti, 1939; Knobloch & Correll, 1962) may apply to E. muelleri, E. petiolatum, E. potosianum, or E. rzedowskii.

30. ELAPHOGLOSSUM MUSCOSUM Elaphoglossum muscosum (Sw.) T. Moore, Index Fil. 12. 1857. Figs. 128A–F. Acrostichum muscosum Sw., Prodr. 128. 1788. Type. Jamaica. Swartz s.n. (S; isotypes B-Willd. 19523, photo GH, BM!). Elaphoglossum hookerianum Underw. ex Maxon, Contr. U.S. Natl. Herb. 13: 6. 1909. Type. Guatemala. Alta Verapaz: Coba´n, Tu¨rckheim II. 1862 (US!; isotype NY!).

Rhizomes short-creeping to erect, to 12 mm diam.; rhizome scales linear, reddish orange, to 13 mm long, with short teeth;

ELAPHOGLOSSUM PALEACEUM

fronds clumped, 13–30 ⫻ 1.9–4.8 cm; phyllopodia inconspicuous or absent; stipes 2⁄5–3⁄5 the frond length, with scales light orange, erose, to 6 mm long, spreading; blades narrowly elliptic or ovatelanceolate, subcoriaceous, bases broadly cuneate to rounded, apices rounded; veins inconspicuous, free, simple or once-forked, ca. 1 mm apart, at ca. 60⬚ angle to costa; hydathodes absent; blade scales with cilia shorter than the width of the scale body, scales of adaxial surfaces white, or with slightly darker centers, peltate, ovate-lanceolate, long-toothed, abundant, longer, darker orange, and less peltate on abaxial surface; fertile fronds as long as the sterile fronds, but with smaller blades and longer stipes (ca. 1⁄3 the fertile frond length), scales sparse, small and inconspicuous among the sporangia.

301

free, simple to once-forked, ca. 1.5 mm apart, at ca. 70⬚ angle to costa; hydathodes absent; adaxial surfaces with narrowly lanceolate scales 1–2 mm long, plane, with long hair-teeth, glabrous with age; abaxial blade scales orange, lanceolate, long-ciliolate, margins somewhat upcurved, scattered on the abaxial surfaces, mostly 1–1.5 mm long; abaxial costal scales 2–5 mm long and with small substellate scales as on stipes; fertile fronds slightly shorter than the sterile fronds, blades linear, 10–14 mm wide, stipes 1⁄2 the frond length, with scales on abaxial midveins as on sterile costa but lacking among sporangia, abaxially with many ciliolate scales 0.5–1 mm long.

Distribution. Epipetric in wet montane forests (alder-oak, pineoak) and oak cloud forests; 1350–1800 m. Mexico. Distribution. Epiphytic in wet montane forests and pine-oak cloud forests; 1300–3000 m. Mexico; Guat, Hond, Salv, CR, Pan; Jam, Hisp; Col, Ven, Ec, Peru. Selected Specimens Examined. Chis (Breedlove 22734, NY; Luteyn 11583, NY; Martı´nez 22423, NY; Mickel 1246, NY). Oax (Anderson & Anderson 4725, MICH; Mickel 1126, 4314, 5379, NY). Ver (Chazaro & Herna´ndez 3648, WIS; Houck 51, MICH).

Elaphoglossum muscosum belongs to a very distinct small group that is distinguished by the blade shape and densely clothed blades and stipes. This species is distinguished from E. engelii by having inconspicuous scales among the sporangia, less toothed rhizome and stipe scales, the more compact rhizome habit, and occurrence at lower elevations.

Specimens Examined. Hgo (Lorence 4902, MEXU). Pue (Boege 1146, MEXU; Rzedowski 31762, MEXU). Ver (Diggs & Nee 2782, NY, UC; Nee & Diggs 24900, F; Nee & Hansen 18697, NY; Taylor et al. 122a, NY; Ventura A. 21, NY; Riba et al. 402, NY).

Elaphoglossum obscurum resembles E. vestitum but is distinct in the long, abundant conspicuous scales, especially on the stipes and costae, and the broader blades (3–6 vs. 1.4–3.1 cm).

32. ELAPHOGLOSSUM PALEACEUM Elaphoglossum paleaceum (Hook. & Grev.) Sledge, Bull. Brit. Mus. (Nat. Hist.) Bot. 4: 95. 1967. Figs. 128G, H, J–P.

Acrostichum obscurum E. Fourn., Mexic. Pl. 1: 68. 1872. Type. Mexico. Veracruz: Misantla, Hahn s.n. (P!).

Acrostichum paleaceum Hook. & Grev., Icon. Filic. 2: t. 235, plate as “paleaceum,” text as “vestitum.” 1832. Type. Madeira. Ribeiro Frio, Lowe s.n., 16 Apr 1828 (K!). Acrostichum squamosum Sw., J. Bot. (Schrader) 1800(2): 11. 1801, hom. illeg., non Cav., 1799. Elaphoglossum squamosum J. Sm., J. Bot. (Hooker) 4: 148. 1841. Type. None cited. Elaphoglossum hirtum sensu auct., non Acrostichum hirtum Sw., J. Bot. (Schrader) 1800(2): 10. 1801.

Rhizomes short-creeping, 5–8 mm diam.; rhizome scales castaneous, lustrous, linear-lanceolate, with long hair-teeth, 5–8 mm long; fronds approximate, 32–75 ⫻ 3–6 cm; phyllopodia distinct; stipes ca. 1⁄3 the frond length, with scales linear-lanceolate, tan-orange, 2–8 mm long, ciliolate, wide-spreading, others smaller, substellate; blades narrowly elliptic, chartaceous, bases cuneate to rounded, apices caudate; veins evident to obscure,

Rhizomes short-creeping, to 10 mm diam.; rhizome scales linear-lanceolate, black (rarely dark brown) and opaque, ca. 5 mm long, margin with rigid hair-like teeth; fronds clumped, 10– 53 ⫻ 0.9–4 cm; phyllopodia distinct; stipes 1⁄4–1⁄3 the frond length, 1–1.5 mm diam., densely clothed with spreading, black or orange scales similar in shape and toothing to those of rhizomes, to 3 mm long; blades narrowly elliptic, chartaceous, bases narrowly

31. ELAPHOGLOSSUM OBSCURUM Elaphoglossum obscurum (E. Fourn.) C. Chr., Index Filic. 312. 1905. Figs. 136M–P.

302

ELAPHOGLOSSUM

to broadly cuneate to rounded, apices acuminate to caudate; veins obscure, free, simple or once-forked, ca. 1 mm apart, at ca. 70⬚ angle to costa; hydathodes absent; abaxial blade scales abundant, plane, scattered or overlapping, lanceolate, ca. 2 mm long, with long hair-teeth, the hairs 1–2 times as long as width of scale body, usually those of the stipes and costae dark, dark ones sometimes scattered in proximal part of blades; adaxial scales similar, but less abundant; fertile fronds shorter than the sterile, to 28 ⫻ 1.4 cm but with same shape, scales along the abaxial costae and only occasionally among the sporangia.

Acrostichum pallidum Baker ex Jenman, J. Bot. 17: 263. 1879. Type. Jamaica. Jenman s.n. (K!; isotype NY!).

Rhizomes short-creeping, 5–8 mm diam.; rhizome scales linear, dark reddish brown, lustrous, 6–12 mm long, each with a flexuous tip; phyllopodia absent; fronds clumped, 24–55 ⫻ 4.1–5.6 cm; stipes 1⁄3–1⁄2 the frond length, with scales subulate, spreading, sparse or absent, and with a few minute, erect, glandular hairs; blades elliptic, chartaceous, apices caudate, bases usually rounded (less commonly broadly cuneate); veins evident, free, once- or twice-forked, 1.5–2 mm apart, at 70–80⬚ angle to costa; hydathodes absent; blade scales black, subulate, mostly along the margin, generally early-deciduous, mature specimens with only a few scales or scales totally absent, blades also with minute, black, stellate hairs; fertile fronds much shorter than the sterile (1⁄2–2⁄3), intersporangial scales absent.

Distribution. Epiphytic and epipetric in wet montane forests and cloud forests; (800–)1200–3000 m. Mexico; Guat, Hond, Salv, CR, Pan; Cuba, Jam, Hisp; Col, Ven, Ec, Peru, Braz, Bol; Azores & Madeira, Hawaii. Selected Specimens Examined. Chis (Breedlove 22735, NY; Martı´nez S. 22422, NY). Gro (Lorea 3128, IEB, NY, 3144, NY). Jal (McVaugh 23166, MICH). Me´x (Tejero-Dı´ez 2701, IZTA). Oax (Lorence 4040, NY; Mickel 4812, 6562, NY). Pue (Riba et al. 332, MEXU). Ver (Ventura A. 2249, NY).

Distribution. Terrestrial, epipetric, or epiphytic in wet pineoak woods and cloud forests; 1850–3000 m. Mexico; Guat, CR, Pan; Jam, Hisp; Col, Ven, Ec.

Unverified, Doubtful, or Mistaken Reports. Mich (reported by Dı´az-Barriga & Palacios-Rios, 1992, but no specimen cited and not verified). Qro (Dı´az-Barriga & Carranza 6556, IEB, cited by Dı´azBarriga & Palacios-Rios, 1992, but not verified).

Selected Specimens Examined. Chis (Breedlove 22804, DS, NY, 32046, NY). Gro (Vela´squez 257, FCME). Mich (Arse`ne s.n., 13 Dec 1910, MICH). Mor (Lyonnet 1359, MEXU, 2863, UC, US). Oax (Lorence 4214, NY; Mickel 3772, 7040, NY; Somers 371, NY).

There is considerable variation in this species in America. One group has black scales on stipes, costae, and broad blades (2.4–4 cm wide) that are densely to only moderately covered with scales. The other large group has only orange-tan scales. None of the Mexican specimens quite match the type of E. paleaceum from Madeira. There are close relatives in Mesoamerica and South America as well as in Africa and India. Unfortunately, there is an earlier name for this species, E. semicylindricum (T. Bowdich) Benl, also of Madeira. For now, we prefer to continue to use the widely applied name E. paleaceum until species limits for neotropical and Hawaiian material in this taxonomically difficult complex can be refined.

33. ELAPHOGLOSSUM PALLIDUM Elaphoglossum pallidum (Baker ex Jenman) C. Chr., Index Filic. 312. 1905. Figs. 127A–D.

Elaphoglossum pallidum is very close to E. erinaceum but can be distinguished by the former having nearly glabrous blade margins and narrow, dark brown rhizome scales with flexuous tips, and lanceolate (vs. elliptic) blades. It was treated previously (Mickel & Beitel, 1988) as a synonym of E. tambillense (Hook.) T. Moore, which occurs from Costa Rica to Ecuador. Antillean material seems to lack marginal subulate scales entirely, whereas Mexican material is variable, either lacking scales or with a few scales.

34. ELAPHOGLOSSUM PARDUEI Elaphoglossum parduei Mickel, Brittonia 32: 337. 1980. Type. Mexico. Oaxaca: Distrito Ixtla´n, 4 km S of Vista Hermosa, 74 km N of Ixtla´n de Jua´rez on Rte 175, Mickel 6615 (NY!; isotypes MEXU!, UC!). Figs. 126M, 131C, D.

ELAPHOGLOSSUM PETIOLATUM

303

Rhizomes moderately creeping, 2–4 mm diam.; rhizome scales deltate-lanceolate, lustrous brown, ca. 2 mm long, entire or with a few weak short hairs, somewhat deciduous, leaving partially naked rhizomes; fronds somewhat spaced, to 30 ⫻ 1–2 cm; phyllopodia distinct, usually 1–1.5 cm long; stipes 1⁄4–1⁄2 the frond length, naked or with a few lanceolate scales proximally; blades narrowly elliptic, coriaceous, bases narrowly cuneate, apices acute to acuminate; veins obscure, free, simple or once-forked, ca. 1 mm apart, at ca. 70⬚ angle to costa; hydathodes absent; blade surfaces glabrous or with minute, black, stellate trichomidia, especially on the abaxial surfaces; fertile fronds slightly longer than the sterile fronds, the stipes longer (1⁄2–3⁄4 the frond length) and the blades slightly narrower, intersporangial scales absent. Distribution. Epiphytic on mossy trunks and fallen logs, in wet montane forests at middle and upper elevations; 600–2450 m. Mexico; Guat, Hond, Salv, Nic, CR, Pan; Jam, Hisp; Col, Ven, Guy, Sur, Fr Gui, Ec, Peru, Bol. Selected Specimens Examined. Chis (Breedlove 21662, NY; Me´ndez G. 7833, NY). Gro (Hinton 14273, NY; Lorea 3168, NY). Hgo (Pringle 10304, NY). Jal (McVaugh 21443, NY). Oax (Hallberg 1514, NY; Mickel 6607, NY). Pue (Tenorio L. 14278, NY). Ver (Purpus 1993, NY).

Distribution. Epiphytic in wet montane forests; 1300–2050 m. Mexico.

Several varieties or forms have been recognized, based on dissection, from undivided to very finely divided; ours are all f. peltatum. This species, the most widespread representative of the splinter genus Peltapteris, is closely allied to E. squamipes, differing only in blade shape and dissection. Molecular studies (Skog et al., 2004) show Peltapteris nested within Elaphoglossum.

Specimens Examined. Oax (Hallberg 1460, NY; Mickel 1009, 1544, NY).

36. ELAPHOGLOSSUM PETIOLATUM

This species is much like E. glabellum but with shorter phyllopodia, broader blades with minute, black, stellate scales, and fertile fronds longer than the sterile.

Elaphoglossum petiolatum (Sw.) Urb., Symb. Antill. 4: 61. 1903. Figs. 129E–G.

35. ELAPHOGLOSSUM PELTATUM Elaphoglossum peltatum (Sw.) Urb., Symb. Antill. 4: 60. 1903. Fig. 130K. Osmunda peltata Sw., Prodr. 127. 1788. Acrostichum peltatum (Sw.) Sw., J. Bot. (Schrader) 1800(2): 11. 1801. Rhipidopteris peltata (Sw.) Schott ex Fe´e, Me´m. Foug. 2: 14. 1845. Peltapteris peltata (Sw.) C. V. Morton, Amer. Fern J. 45: 13. 1955. Type. Jamaica. Swartz s.n. (S!).

Rhizomes long-creeping, slender, ca. 1 mm diam.; rhizome scales lanceolate, tan; fronds dimorphic; sterile fronds 3–15 ⫻ 2.5–5 cm, distant; stipes ca. 3⁄4 the frond length, green to stramineous, sparsely scaly; phyllopodia absent; blades flabellate, mostly 4–6 times dichotomously divided, the segments linear, 0.5–1.5 mm wide, laminae with a few small scales; fertile fronds longer than the sterile and nearly undivided, usually two-lobed, 5–20 mm wide, with sterile margins ca. 0.5 mm wide; 2n⫽80 (CR).

Acrostichum petiolatum Sw., Prodr. 128. 1788. Acrostichum viscosum Sw., Syn. Fil. 10, 193. 1806, nom. superfl. Elaphoglossum viscosum (Sw.) J. Sm., J. Bot. (Hooker) 4: 148. 1841. Type. Jamaica. Swartz s.n. (S!, frag. US!). Acrostichum schiedei Kunze, Analecta Pteridogr. 10. 1837. Type. Mexico. Veracruz: Jalapa, Schiede s.n. (B!; isotypes BR!, NY!, P!). Acrostichum schmitzii Mett. ex Kuhn, Linnaea 36: 51. 1869. Syntypes. Mexico. Schmitz 3 (B), Karwinski s.n. (B?), Schaffner s.n. (B?).

Rhizomes short-creeping, compact, horizontal, 3–5 mm diam.; rhizome scales linear, blackish maroon to castaneous, each with tortuous hair tip, ca. 3 mm long, entire; fronds clumped, 10– 39(–51) ⫻ 0.9–3.4 cm; phyllopodia distinct, but very short (2–3 mm long); stipes ca. 1⁄4 the frond length, with scales 1–2 mm long, some spreading and broader, some narrow and appressed, entire or toothed to stellate; blades narrowly elliptic, chartaceous, bases cuneate, apices acuminate; veins visible, free, simple or once-forked, ca. 1 mm apart, at 60–70⬚ angle to costa; hydathodes absent; blade scales sparse, linear-lanceolate with hair teeth, mostly on the adaxial surfaces and on young fronds, often more abundant near the margins, occasionally with resin dots; abaxial surfaces usually with resin dots, occasionally with small linear-

304

ELAPHOGLOSSUM

lanceolate scales or substellate hairs as on adaxial surface, rarely glabrous; fertile fronds slightly shorter than to far exceeding the sterile fronds but with proportionally longer stipes, ca. 2⁄3 the fertile frond length, scales on the abaxial costae, absent among sporangia absent; 2n⫽164 (Jam).

Acrostichum piloselloides C. Presl, Reliq. Haenk. 1(1): 14, t. 2, f. 1. 1825. Type. “Hab. in montanis Peruviae.” Haenke s.n. (PR).

Rhizomes erect or ascending, ca. 2 mm diam.; rhizome scales linear, orange, entire, 3–6 mm long; fronds clumped, 2–4(–8) ⫻ 0.4–1 cm; phyllopodia absent; stipes slender, usually 2⁄3–3⁄4 the frond length, moderately clothed with tan to orange, very narrow, subulate scales, 1.5–2 mm long; blades spatulate, obovatelanceolate to narrowly elliptic, chartaceous, bases broadly cuneate, apices obtuse; veins obscure, free, simple or rarely once-forked, ca. 1 mm apart, at 40–50⬚ angle to costa; hydathodes present but inconspicuous; blade scales somewhat subulate (slightly inrolled at base), tan to dull orange, 1–2 mm, uniformly and moderately covering the blade surfaces; fertile fronds as tall as or taller than the sterile, more erect, stipes ca. 4⁄5 the length, blades smaller than those of the sterile fronds, often folded in half, spathulate to nearly round, scales of blades and distal part of stipes dark brown to black, lustrous, scale bases not inrolled but spreading and slightly toothed, soral scales absent except for bases of costae.

Distribution. Epiphytic or terrestrial in wet montane forests; 1700–3000 m. Mexico; Guat, Hond, Salv, Nic, CR, Pan; Gr & L Ant; Col, Ven, Ec, Peru, Braz. Selected Specimens Examined. Chis (Matuda 2584, F, K, MEXU, NY, US). Col (Goldsmith 41, NY). DF (Lyonnet 911, NY). Gro (Lorea 3148, NY). Gto (Rzedowski 47827, XAL). Hgo (Pringle 10837, MSC, WIS). Jal (McVaugh 14040, MICH). Me´x (Rzedowski 26741, NY). Mich (Hinton 15368, NY). Mor (Lyonnet 591, NY). Nay (Rose 2210, US). Oax (Mickel 3773, NY). Pue (Marcks & Marcks 758, NY). Qro (Arguelles 1310, MEXU). Sin (Vega A. 3126, ENCB). Tlax (Sohns 654, UC). Ver (Purpus 2350, NY). Zac (Rose 2728, US).

There is considerable variation in the degree of scaliness, especially on the abaxial surfaces, and the degree of resinous dotting, but the significance of this variation is not yet understood. Elaphoglossum petiolatum is widespread in tropical America, and if construed in a broad sense occurs also in the Old World tropics. In Africa it is usually called E. salicifolium (Willd. ex Kaulf.) Alston; in St. Helena, E. nervosum (Bory) Christ; in India, E. stelligerum (Wall. ex Baker) T. Moore ex Alston & Bonner; and in China, Southeast Asia, and the Pacific, E. yunnanense (Baker) C. Chr. and E. blumeanum (Fe´e) J. Sm. The E. petiolatum complex is characterized by having resinous dots on the abaxial blade surfaces, usually black, entire rhizome scales, and dark-tipped scales on the proximal portions of the stipes. Mexico has several taxa in this group: E. rzedowskii in the northwest, E. potosianum in the northeast, E. pringlei in the south, and E. petiolatum throughout much of the country. Elaphoglossum petiolatum closely resembles E. huacsaro in the resinous dots of the abaxial blade surfaces, the frequent concentration of scales along the blade margins, and the fertile fronds noticeably longer than the sterile the but E. huacsaro has more erect rhizomes and obtuse blade apices.

37. ELAPHOGLOSSUM PILOSELLOIDES Elaphoglossum piloselloides (C. Presl) T. Moore, Index Fil. 13. 1857. Figs. 130A–C.

Distribution. Terrestrial or epipetric in oak and pine-oak forests; 1000–2600 m. Mexico; Guat, Hond, Salv, CR, Pan; Hisp; Col, Ven, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol, Chile. Selected Specimens Examined. Chis (Ghiesbreght 313, F, K, NY). Gro (Rzedowski 26015, ENCB, NY). Hgo (Pringle 15628, CAS, US). Jal (McVaugh 13997, MICH). Me´x (Hinton 8205, NY). Mich (Pringle 13253, UC). Mor (Lyonnet 3333, US). Nay (Norris & Taranto 14602, MICH). Oax (Pringle 4964, GH, US). Sin (Breedlove 44967, ENCB, CAS). Ver (Bourgeau 3273, NY).

Elaphoglossum piloselloides is often placed under E. spatulatum (Bory) T. Moore of Africa, Madagascar, Re´union, and Sri Lanka, but differs from it in having the scales of the fertile fronds black as opposed to orange-tan as on the sterile fronds. African material is also highly variable in blade shape, often being somewhat elongate to linear.

38. ELAPHOGLOSSUM PILOSIUS Elaphoglossum pilosius Mickel, Brittonia 39: 324. 1987. Type. Venezuela. Me´rida: trail leading from La Negrita to the Boquero´n of the Quebrada de las Can˜as, Luteyn 6139 (NY!). Figs. 134A–E.

ELAPHOGLOSSUM PRINGLEI

Rhizomes compact, ca. 2 mm diam.; rhizome scales spreading, linear-lanceolate, dark reddish brown, lustrous, 7–10 mm long, denticulate; fronds approximate, (6–)9–16 cm long; phyllopodia absent; stipes light brown, 1⁄2–2⁄3 the frond length, with scales patent, subulate, reddish brown, lustrous, 2–3 mm long, denticulate, mixed with minute glandular hairs; blades lanceolate, 4–7 ⫻ 1.8–2.7 cm, bases rounded to truncate, apices cuspidate; veins evident, at ca. 75⬚ angle to costa; hydathodes present; blade scales scattered, subulate, reddish brown, lustrous, 2–3 mm long on adaxial surfaces, costae and margins, 1–1.5 mm long on abaxial surfaces; fertile fronds longer than the sterile, stipes 3⁄4 the frond length, blades oblong, bases rounded to truncate, apices obtuse, often folded along costae (conduplicate), with costal scales dense, subulate.

305

once-forked, ca. 1 mm apart, at 60–70⬚ angle to costa; hydathodes absent; blade scales adaxially scattered, substellate, mostly ca. 0.5 mm, abaxially the blades with simple to 3-armed trichomidia subtended or not by resinous dots; fertile fronds surpassing the sterile in length, stipes ca. 2⁄3 the frond length, with a few lanceolate scales on abaxial costae.

Distribution. Terrestrial in pine-oak woods; 2000–2540 m. Mexico. Specimens Examined. DF (Schaffner 93, NY). Gto (Correll & Correll 2882, LL). Hgo (Knobloch 682, MSC; Orcutt 885, DS). NL (Dorr 2597, TEX; Hinton 22119, ARIZ, TEX, 22679, NY). SLP (Schaffner 92, NY). Tam (Gonza´lez Medrano 17496, MEXU).

Distribution. Epipetric on pine-oak slopes; 2750–3200 m. Mexico; CR, Pan; Col, Ven, Ec, Peru. Specimens Examined. Chis (Breedlove 22065, NY, 22806, DS, 55848, CAS, ENCB; Martı´nez S. 22435, MEXU; Martı´nez S. & Soto 24298A, MEXU, XAL). Gro (Lorea 4206, 4211, ENCB).

Elaphoglossum pilosius is close to E. lindenii, but the former has rounded to truncate blade bases, and the stipes and blades are conspicuously scaly.

39. ELAPHOGLOSSUM POTOSIANUM

This species is similar to E. petiolatum except: abaxial blade indument of simple to 3-armed stellate trichomidia with or without small resinous subtending dots (vs. large dots and/or substellate scales), and rhizome scales usually orange, occasionally black with age (vs. always black). Elaphoglossum potosianum has been mistakenly identified as E. muelleri because of the orange rhizome scales.

40. ELAPHOGLOSSUM PRINGLEI Elaphoglossum pringlei (Davenp.) C. Chr., Index Filic. 313. 1905. Figs. 128R–W.

Elaphoglossum potosianum Christ, Monogr. Elaph. 119. 1899. Type. Mexico. San Luis Potosı´: Schaffner 77 (P!). Figs. 139A–C.

Acrostichum pringlei Davenp., Bot. Gaz. 21: 253. 1894. Type. Mexico. Oaxaca: Sierra de San Felipe, Pringle 5605 (GH!; isotypes GH!–2 sheets, K!, MEXU!, P!, US!).

Elaphoglossum trichomidiatum Mickel, Brittonia 53: 488. 2001[2002]. Type. Mexico. San Luis Potosı´: In the region of San Luis Potosı´, 1878, Parry & Palmer 1007 (NY!; isotype US!).

Rhizomes short-creeping, compact, horizontal, 2–4 mm diam.; rhizome scales linear-lanceolate, dark brown, lustrous, entire, ca. 2 mm long; fronds clumped, to 18 ⫻ 0.5–3 cm; phyllopodia distinct to indistinct; stipes 1⁄5–1⁄2 the frond length, with scales both spreading, lanceolate and reduced, appressed, substellate; blades narrowly elliptic, chartaceous, bases cuneate, apices acute to acuminate; veins obscure, free, simple or onceforked, 0.6–1 mm apart, at 60–70⬚ angle to costa; hydathodes absent; blade scales of adaxial surfaces lanceolate, deeply toothed, on abaxial surface substellate (stellate with some scale body); fertile fronds longer than the sterile and very long-stiped, stipes about 2⁄3 the frond length, blades narrower than those of sterile

Rhizomes short-creeping, 2–3 mm diam.; rhizome scales lanceolate to linear-lanceolate, orange, occasionally black with age (especially in Hidalgo), ca. 3 mm long, apices attenuate, subentire with occasional short teeth; fronds approximate, 12–21 ⫻ 1.4–2.2 cm; phyllopodia distinct, 3–4 mm; stipes 1⁄3–1⁄2 the sterile frond length, with scales ca. 1 mm, scattered, mostly appressed, stellate, proximally with a few lanceolate scales like those of rhizomes; blades narrowly elliptic, chartaceous, bases narrowly to broadly cuneate, apices acute to acuminate; veins free, simple or

306

ELAPHOGLOSSUM

fronds, 0.7–1.2 cm wide, scales on the abaxial costae but not among the sporangia.

Distribution. Terrestrial in dry pine-oak forests; 1300–2750 m. Mexico. Selected Specimens Examined. Oax (Anderson & Anderson 4778, NY; Campos V. 2650, NY; Hallberg 1811, NY; Hellwig 260, NY; Mickel 822, 3737, 4593, 5329, 6809, NY; Rzedowski 21009, NY). Unverified, Doubtful, or Mistaken Reports. Mich (reported by Mickel & Beitel, but specimens redetermined as E. rzedowskii).

It is surprising to find so many collections from Oaxaca and none from other states. Elaphoglossum pringlei resembles E. muelleri but differs in having dark, lustrous rhizome scales and substellate abaxial blade scales, with at least a narrow scale body. Elaphoglossum pringlei is closely allied with E. petiolatum in its entire rhizome scales and blade size and shape, but there are no resin dots on the blades and the scales are substellate abaxially.

41. ELAPHOGLOSSUM PUSILLUM Elaphoglossum pusillum (Mett. ex Kuhn) C. Chr., Index Filic. 314. 1905. Figs. 134M, N. Acrostichum pusillum Mett. ex Kuhn, Linnaea 36: 43. 1869. Type. Cuba, “Cuba orientali,” Wright 794 (B!; isotypes GH–empty folder, P!, US!, YU!).

Rhizomes 1–2 mm diam., compact; rhizome scales 2.5–3.5 mm long, linear, orange-tan, lustrous, entire; fronds 3.6–11.5 cm long, approximate; phyllopodia absent; stipes 1⁄5–1⁄3 the sterile frond length, brown; stipe scales 1.4–2.6 mm long, spreading, subulate, orange-brown, lustrous, denticulate, mixed with minute erect glandular hairs; sterile blades 2.5–6.5 ⫻ 0.9–1.6 cm, oblong, chartaceous, bases broadly cuneate, apices obtuse; veins visible but not conspicuous, ca. 2 mm apart, at ca. 50⬚ angle to costa; hydathodes present; blade scales 1–2 mm long, scattered, spreading, linear, subulate, orange-brown to dark brown, lustrous, denticulate; fertile fronds slightly shorter than the sterile, stipes 4⁄5 the fertile frond length, fertile blades 8–16 mm long, ovate to suborbicular, plane, bases broadly rounded, apices obtuse, intersporangial scales absent.

Distribution. Epiphytic, epipetric, and terrestrial in wet forests; 1350 m (300–1600 m in Mesoamerica). Mexico; Guat, Nic, CR, Pan; Cuba; Col, Ven.. Specimens Examined. Chis (Breedlove 32527, DS, MEXU, NY). Oax (Cedillo T. 1896, IEB, MEXU).

This species resembles E. piloselloides, but the blades are thinner, oblong rather than spatulate, and the hydathodes are more conspicuous.

42. ELAPHOGLOSSUM REVOLUTUM Elaphoglossum revolutum (Liebm.) T. Moore, Index Fil. 365. 1862. Figs. 132I–K. Acrostichum revolutum Liebm., Mexic. Bregn. 163 (reprint 11). 1849. Lectotype (chosen by Mickel & Beitel, 1988: 181). Mexico. Oaxaca: Chinantla, Lobani to Petlapa, Liebmann s.n. [Pl. Mex. 2427, Fl. Mex. 4] (C!; isolectotypes BM, US!, frag. NY!). Acrostichum rampans Baker in Hooker & Baker, Syn. Fil., ed. 2, 518. 1874. Elaphoglossum rampans (Baker) Christ, Monogr. Elaph. 56. 1899. Type. Cuba. Wright 3959 (K!; isotypes NY!, P!).

Rhizomes long-creeping, ca. 1 mm diam.; rhizome scales ovatelanceolate, orange, ca. 3 mm long, entire; fronds usually 4–8 mm apart, 5–15 ⫻ 0.6–1.1 cm; phyllopodia absent; stipes ca. 1⁄4(–1⁄2) the frond length, with orange, ovate, somewhat spreading scales 2 mm long; blades narrowly elliptic, coriaceous, bases cuneate, apices obtuse; veins obscure, free, simple or once-forked, ca. 0.8 mm apart, at ca. 60⬚ angle to costa; hydathodes absent; blade surfaces glabrous or with a few ovate scales on the abaxial costae; fertile fronds exceeding the sterile fronds, stipes 1⁄2–3⁄4 the frond length, blade narrow, to 4 mm wide, scales absent on the abaxial surfaces.

ELAPHOGLOSSUM RUFESCENS

307

Distribution. Epiphytic in wet montane forests; 600–1000 m. Mexico; Guat, Hond, CR, Pan; Cuba, Jam, Hisp; Col, Ec, Peru, Bol, Braz.

Distribution. Epiphytic in montane rain and pine-oakLiquidambar forests; 800–1300(–2150) m. Mexico; Guat, Hond, Nic.

Selected Specimens Examined. Chis (Breedlove 21894, NY, 27432, MEXU; Martı´nez S. 18661, 18758, MEXU, 23981, NY; Nelson 3048, US). Gro (Lorea 11, FCME, IEB). Oax (Liebmann s.n. [Pl. Mex. 2427], C; Mickel 7326, NY). Ver (Valdivia Q. 942, XAL).

Selected Specimens Examined. Chis (Breedlove 22346, 27568, 32131, 32896, DS, 47901, 53264, 53576, 56659, CAS; Martı´nez 1841, MEXU; Martı´nez S. 23501, MEXU).

Elaphoglossum revolutum is closely allied to E. squamipes and E. peltatum, as evidenced by the rhizome habit, scales, and spores. It is distinguished from E. affine, E. tenuifolium, and small specimens of E. guatemalense by the orange, ovatelanceolate rhizome scales and the lack of phyllopodia, and from E. squamipes by the generally more elliptic blades and occurrence at much lower elevations.

43. ELAPHOGLOSSUM RUBESCENS Elaphoglossum rubescens Christ, Monogr. Elaph. 93, f. 47, 48. 1899. Type. Guatemala. Verapaz: Bernoulli 320 (P, not found). Figs. 137F, G, H, J. Rhizomes short-creeping, ca. 5 mm diam.; rhizome scales castaneous, lustrous, linear-lanceolate, 2–4 mm long, entire or with short teeth; fronds clumped, 30–65 ⫻ 1.5–3.2 cm; phyllopodia distinct; stipes 1⁄4–1⁄3 the frond length, sparsely clothed with lanceolate, spreading, toothed scales ca. 2 mm long, more copiously covered with small, appressed, shorter scales with long hair-teeth, scales sometimes reduced nearly to stellate hairs, the stipes often with resin dots; blades linear-elliptic, bases broadly cuneate, apices caudate; veins obscure, free, simple or once-forked, 1–1.2 mm apart, at 70⬚ angle to costa; hydathodes absent; blade scales abundant but not overlapping, adaxially delicate, white, appressed, round to subdeltate, with short hair-teeth, or scales absent with age; abaxial surfaces with scales orange, narrow to deltate, with long hair-teeth and distinctly upturned margins; blade margins especially scaly with scales flat and overlapping; fertile fronds about equal to sterile fronds in length, but with narrower blades (0.7–0.9 cm wide) and longer stipes (1⁄3–1⁄2 the frond length), scales on costae abaxially but not among sporangia.

Elaphoglossum rubescens is most closely allied to E. vestitum, resembling it in rhizome scale color and texture, ciliolate blade scales lightly covering the laminae, and lax fronds with caudate blade tips. The former can be distinguished by the rhizome scales not (or only weakly) toothed, the stipes often with resin dots, and the blade scales with the teeth and margins upturned. These two species may be only varietally distinct, overlapping in Chiapas with several intermediate specimens. Elaphoglossum auricomum belongs to the same complex, but has more abundant spreading stipe scales, the rhizome scales are lighter in color, and the blades have acuminate apices. Some specimens of E. paleaceum have blades only lightly covered with scales, but they have black scales on the stipes and proximal part of abaxial rachises.

44. ELAPHOGLOSSUM RUFESCENS Elaphoglossum rufescens (Liebm.) T. Moore, Index Fil. 365. 1862. Figs. 136A–E. Acrostichum rufescens, Liebm., Mexic. Bregn. 171 (reprint 18). 1849. Type. Mexico. Volca´n Orizaba, Liebmann s.n. (C!, frag. NY!; isotypes GH!, K!, P!, US!). Acrostichum roezlii W. Schaffn. in Fe´e, Me´m. Foug. 8: 69. 1857. Elaphoglossum roezlii (W. Schaffn.) T. Moore, Index Fil. 14. 1857. Type. Mexico. Me´xico: Popocate´petl, Schaffner 280 (P!). Elaphoglossum laxipes Mickel in McVaugh, Fl. Nov.-Gal. 17: 271. 1992. Type. Mexico. Me´xico: Temascaltepec, Meson Viejo, Hinton 2750 (NY!; isotypes MO!, NY!).

Rhizomes compact, horizontal, 2–3 mm diam.; rhizome scales linear-lanceolate, castaneous to blackish brown, lustrous, 5–6 mm long, with long slender apices, the hair-teeth deciduous or sparse; fronds clumped, (7–)16–30 ⫻ (0.8–)1.2–1.7 cm; phyllopodia distinct; stipes 1⁄3–1⁄2 the frond length, 0.7–0.9 mm diam., with scales sparse, appressed to spreading, brownish black, often

308

ELAPHOGLOSSUM

orange distally, concolorous, 1–2 mm long; blades narrowly elliptic, narrowly cuneate at bases, acuminate at apices (rounded in juvenile plants); veins obscure, ca. 1 mm apart, at 70⬚ angle to costa; hydathodes absent; blade scales lanceolate, mostly 1.5–2 mm long, short-toothed with the teeth ca. 1⁄2 the width of the scale body, scattered on the adaxial surfaces, orange-tan to white, abaxially abundant, imbricate, orange-tan, those of the costae 3– 4 mm long, often dark-centered; fertile fronds shorter than (ca. 1⁄2 to nearly as long as) the sterile, the blades narrowly elliptic, 1–1.1 cm wide, bases broadly cuneate to rounded, apices acute, the costae scaly abaxially, the adaxial scales imbricate, absent among sporangia.

25 ⫻ 0.5–0.8 cm; phyllopodia indistinct or absent; stipes 1⁄8–1⁄3 of the frond length, clothed with linear-lanceolate, entire, spreading, dark-tipped scales (similar to those of rhizomes) and much reduced stellate or hastate, appressed, resin-based scales (similar to those of the adaxial blade surfaces); blades linear-elliptic, acuminate at both ends; papyraceous; veins obscure, free, simple or once-forked, ca. 1 mm apart, at ca. 60⬚ angle to costa; hydathodes absent; adaxial blade surfaces with abundant but not overlapping lanceolate, hastate, or stellate scales with resinous bases; abaxial surfaces densely scaly with overlapping ciliolate scales, costae with sparse scales similar to those of rhizomes (linear-lanceolate, entire, dark-tipped); fertile fronds about same length as the sterile but much longer-stiped (1⁄2 of the fertile frond length), intersporangial scales absent.

Distribution. Epipetric in pine-fir forest; 2600–3350 m. Mexico. Selected Specimens Examined. DF (Lyonnett 3469, US; Matuda 18794, US; Pringle 7088, MEXU; Purpus 1587, US; Rzedowski 34688, IEB). Jal (McVaugh 10044, MICH). Me´x (Hinton 2750, US; Matuda 18794, MEXU; Rzedowski 23280, ENCB, NY). Pue (Balls 4469, US).

Elaphoglossum rufescens resembles E. paleaceum in the abundant blade scales but is distinct in the castaneous, sparsely toothed rhizome scales, slender stipes (0.7–0.9 mm diam. instead of 1–1.5 mm), and longer stipes (1⁄3–1⁄2 the length of the frond instead of 1⁄5–1⁄4); the scales of the stipes and costae are dark-centered, not black-tipped, and the blade scales have short, stiff teeth (shorter than the width of the scale body instead of equal to, or longer than, the width). Elaphoglossum laxipes was distinguished by having longer, more slender stipes, but subsequent observations on Popocatepe´tl showed transitional forms to E. rufescens.

45. ELAPHOGLOSSUM RZEDOWSKII Elaphoglossum rzedowskii Mickel, Brittonia 32: 337. 1980. Type. Mexico. Jalisco: Municipio de San Martı´n de Bolan˜os, Las Vidrieras, 10 km NW of El Platanar, Rzedowski 26161a (MICH!; isotype IEB!). Figs. 135L–P. Rhizomes short-creeping, horizontal, compact, 2–3 mm diam., obscured by old frond bases and appearing ca. 1 cm diam.; rhizome scales linear-lanceolate with long, hair-like apices, lustrous dark brown, mostly 2–3 mm long; fronds clumped, to

Distribution. Epipetric on shaded ledges in pine-oak forest; 1900–2700 m. Mexico. Selected Specimens Examined. Ags (McVaugh 18431, MICH). Chih (Correll & Gentry 22972, NY). Dgo (Torres C. 3515, MEXU). Gro (Rzedowski 26032, NY). Gto (Rzedowski 45444, IEB). Jal (Anderson & Laskowski 3784, ENCB). Mich (Leavenworth 717, NY). Sin (Sanders 21170, UCR). Son (s. coll. 118, MEXU). Zac (Rzedowski & McVaugh 1006, ENCB).

Elaphoglossum rzedowskii is a member of the E. petiolatum complex, which has entire or subentire rhizome and stipe scales and dissected blade scales, with a tendency for the scales to be reduced to resinous dots. The former species is distinct in the scaliness of the abaxial blade surfaces, and in this is close to E. pringlei. Elaphoglossum pringlei can be distinguished by the broader blades with acute rather than acuminate blade apices. It is possible that E. pringlei and E. rzedowskii are conspecific and represent regional variation of a single taxon.

46. ELAPHOGLOSSUM SARTORII Elaphoglossum sartorii (Liebm.) Mickel, Mem. New York Bot. Gard. 46: 181. 1988. Figs. 124A–H, 126F–H, J, 138L–P. Acrostichum sartorii Liebm., Mexic. Bregn. 166 (reprint 14). 1849. Lectotype (chosen by Mickel & Beitel, 1988: 181). Mexico. Veracruz: Hac. de Mirador, Liebmann s.n. [Pl. Mex. 2430, Fl. Mex. 14] (C!; isolectotypes P!–2 sheets). Elaphoglossum nanchititlense Matuda, Helechos Edo. Me´x. 36. 1956. Type.

ELAPHOGLOSSUM SETIGERUM

309

Mexico. Me´xico: Can˜ada de Nanchititla, Matuda et al. 31589 (MEXU!; isotype US!). Acrostichum scapellum Kunze ex Fe´e, Me´m. Foug. 9: 1. 1857. Elaphoglossum scapellum (Kunze ex Fe´e) T. Moore, Index Fil. 365. 1862. Type. Mexico. Veracruz: “Bois de la Cordillie`re,” Galeotti 6304 (P!).

row fertile blades and bases of sterile blades broadly cuneate at 60–95⬚ angles). Elaphoglossum viride is also very similar and is distinct by its incurved rhizome scales with concave isodiametric cells.

Rhizomes short-creeping, rarely moderately creeping, 4–10 mm diam.; rhizome scales linear-lanceolate, orange to dull tan or dirty brown, occasionally somewhat lustrous or with a blackish cast, to 8 mm long, entire; fronds clumped, rarely slightly spaced, (9–)16–70 ⫻ (1.5–)2.4–7 cm; phyllopodia distinct; stipes usually 1 ⁄4–1⁄3 the frond length, glabrous or sparsely clothed with spreading, lanceolate, irregularly toothed scales, rarely with abundant scales; blades narrowly elliptic, coriaceous, bases cuneate to broadly cuneate, apices acuminate; veins obscure, free, simple to twice-forked, ca. 1 mm apart, at 70–80⬚ angle to costa; hydathodes absent; blade surfaces glabrous or with minute stellate black hairs on the abaxial surfaces; fertile fronds about equal to the sterile fronds in size but sometimes with slightly narrower blades or longer stipes, intersporangial scales absent.

47. ELAPHOGLOSSUM SEMINUDUM Elaphoglossum seminudum Mickel, Brittonia 32: 338. 1980. Type. Mexico. Oaxaca: Distrito Mixe, SE shoulder of Cerro Zempoalte´petl, below Patio de Arena, Mickel 4908 (NY!; isotypes MEXU!, UC!). Figs. 126A, B. Rhizomes very long-creeping, ca. 3 mm diam.; rhizome scales ovate, pale brown, black at peltate attachment, mostly 3–4 mm long, entire, somewhat deciduous, leaving mostly naked rhizomes; fronds 2–4 cm apart, to 38 ⫻ 2.5–3.5 cm; phyllopodia distinct, 1.5–3 cm long; stipes 1⁄3–1⁄2 the frond length, with scattered ovate scales similar to those of rhizomes and minute, black, appressed, stellate hairs; blades lanceolate to narrowly elliptic, coriaceous, bases broadly cuneate, apices acute; veins obscure, free, simple or once-forked, ca. 1 mm apart, at ca. 70⬚ angle to costa; hydathodes absent; blade surfaces nearly glabrous with minute stellate hairs and a few broader scales on the costae, especially on the abaxial surfaces; fertile fronds slightly to much longer than the sterile fronds, the stipes longer (1⁄2–2⁄3 the frond length) and the blades slightly narrower, scales absent throughout.

Distribution. Epiphytic or terrestrial in wet montane forests and cloud forests; 300–3000 m. Mexico. Material outside of Mexico over much of Latin America is closely allied and goes under the name of E. latifolium s.l., or by regional names. Selected Specimens Examined. Chis (Breedlove 14891, NY). Dgo (Ortega 4461, US). Gro (Hinton 14303, NY). Hgo (Pringle 10307, UC). Jal (Mexia 1658, MICH). Me´x (Hinton 3455, NY). Mich (Dı´az Barriga 4674a, IEB). Mor (Hinton 17448, NY). Nay (McVaugh 18897, MICH). Oax (Mickel 5299, NY). Pue (Herna´ndez X. 813, US). Sin (Gonza´lez Ortega 4461, MEXU). Ver (Purpus 1452, NY).

Elaphoglossum sartorii is very close to, but probably distinct from, E. latifolium (Sw.) J. Sm. of the West Indies, and these species represent perhaps the most difficult complex in the genus. The blades are essentially glabrous (at most with minute, stellate trichomidia abaxially), and the rhizomes generally compact, stout, with linear to linear-lanceolate, orange-tan to dirty brown scales. There is great variation in frond size, blade shape, and rhizome scale color, shape, and size. Some of the variation falls into consistent patterns, and we are here describing three recognizable variants at the species level: E. xanthopodum (with long, linear, orange rhizome scales), E. leebrowniae (with large rhizome scales tinged with black), and E. yourkeorum (with nar-

Distribution. Terrestrial in wet montane and cloud forests; 1750–2450 m. Mexico. Selected Specimens Examined. Hgo (Herrera & Riba s.n., Jan 1961, MEXU; Lorence 4919, MEXU; Pringle 10307, CAS, CHAPA, ENCB, IEB, MICH). Oax (Breedlove 65989, CAS; Hellwig 375, NY; Mickel 4656, 5592, NY). Ver (Bohs et al. 1838, GH, XAL; Taylor & Nee 257, NY; Ventura A. 357, IEB, NY).

Elaphoglossum seminudum is close to E. lingua (Raddi) Brack. of Central and South America and the West Indies, but has more slender blades, and tan, rather than black, rhizome scales.

48. ELAPHOGLOSSUM SETIGERUM Elaphoglossum setigerum (Sodiro) Diels in Engler & Prantl, Nat. Pflanzenfam. 1(4): 333. 1899. Figs. 130G, H, J.

310

ELAPHOGLOSSUM

Acrostichum setigerum Sodiro, Anales Univ. Centr. Ecuador 4: 174. 1890. Type. Ecuador. Cordillera Occidental, cerca de Nono, Sodiro s.n. (QPLS). Elaphoglossum crinipes C. Chr., Kongl. Svenska Vetenskapsakad. Handl. III, se´r. 3, 16(2): 76, t. 17, f. 1–2. 1936. Type. Haiti. Croix-des-Bouquets, Badeau, Ekman 7708 (BM!; isotype US!).

Rhizomes short-creeping, to 9 mm diam.; rhizome scales linear-lanceolate, reddish orange, to 12 mm long, with short teeth; fronds clumped, 10–57 ⫻ 1.2–4 cm; phyllopodia absent; stipes 1⁄4–1⁄2 the frond length, with scales reddish orange, spreading, subulate, slightly toothed, to 4 mm long; blades linearlanceolate, 10–15 times as long as wide, chartaceous, bases rounded, truncate, or broadly cuneate, apices acuminate; veins distinct, free, simple or once-forked, 1.5–2.5 mm apart, at 60– 70⬚ angle to costa; hydathodes evident; blade scales abundant, reddish orange, subulate, slightly toothed, denser at margins, spreading to 5–6 mm long, usually less scaly on the adaxial surfaces; fertile fronds slightly shorter that the sterile, to 38 cm long, 2.5 cm wide, with longer stipes ca. 3⁄4 the frond length, bases subcordate, apices acute, intersporangial scales absent.

5. 1909. Type. Guatemala. Alta Verapaz: Pansamala´, Tu¨rckheim s.n. [Donn. Sm. 1003] (US!; isotypes K!, NY!, UC!, YU!).

Rhizomes short-creeping, 1–2 mm diam.; rhizome scales linear, with sparse and irregular short teeth, brown to dull orange, 3–5 mm long; fronds clumped, 8–18 ⫻ 0.8–1.5 cm; phyllopodia absent; stipes slender, 1⁄5–2⁄5 the frond length, moderately clothed with slender, dull orange, subulate, spreading scales 2–3 mm long; blades narrowly elliptic, bases cuneate, apices gradually acuminate, lamina thin; veins conspicuous, free, mostly simple or rarely once-forked, 1.5–2 mm apart, at ca. 40⬚ angle to costa; hydathodes conspicuous; blade scales subulate, dull orange, moderately and uniformly clothing blade surfaces, more so on abaxial surfaces, longer and spreading at blade margins; fertile fronds nearly as long as the sterile but more elliptic, ca. 3 cm long and 1.1 cm wide, with acute apices and longer stipes ca. 3⁄4 the frond length, intersporangial scales abundant.

Distribution. Epiphytic in montane rain forests; 750–1400 m. Mexico; Guat, Salv, Nic, CR. Distribution. Epiphytic, rarely epipetric or terrestrial, in montane oak and pine-oak rain forests and cloud forests; 1850– 2700 m. Mexico; Guat, Pan; Hisp; Col, Ec, Peru.

Specimens Examined. Chis (Breedlove 32246, 38908, DS; 53467, 56239, 56551, 57767, CAS; Dressler 1599 GH, US). Oax (Geith 95-MO16 bis, NY). Ver (only the type).

Selected Specimens Examined. Chis (Breedlove 31888, DS; Ghiesbreght 308, NY). Gro (Lorea 2300, FCME, IEB, NY). Oax (Lorence 4213, NY; Mickel 3785, 3798, 4118, 4905, 4906, 7043, NY).

This species is superficially much like E. crinipes, but it can be distinguished by the smaller frond size and the presence of scales among the sporangia. It is also like E. monicae, which has more elliptic blades and darker blade scales.

This species closely resembles E. villosum (Sw.) J. Sm. of the West Indies, but is distinct by E. villosum having scales among the sporangia and blades with narrowly cuneate bases and caudate apices.

50. ELAPHOGLOSSUM SQUAMIPES Elaphoglossum squamipes (Hook.) T. Moore, Index Fil. 15. 1857. Figs. 127F–H, J–M.

49. ELAPHOGLOSSUM SETOSUM Elaphoglossum setosum (Liebm.) T. Moore, Index Fil. 366. 1862. Figs. 134F–H. Acrostichum setosum Liebm., Mexic. Bregn. 169. (reprint 17) 1849. Acrostichum recognitum Kunze ex E. Fourn., Mexic. Pl. 1: 67. 1872, nom. superfl. Type. Mexico. Veracruz: Barranca de Huitamalco, Liebmann s.n. [Pl. Mex. 2433, Fl. Mex. 29, 30] (C!; isotypes K!, P!, US!). Elaphoglossum catharinae Underw. ex Maxon, Contr. U.S. Natl. Herb. 13:

Acrostichum squamipes Hook., Icon. Pl., ser. 1, 2: t. 197. 1837. Type. Peru. Chacapoyas, Mathews s.n. (K!). Acrostichum ovatum Liebm., Mexic. Bregn. 164 (reprint 12). 1849, hom. illeg., non Hook. & Grev., 1829. Elaphoglossum subaequale T. Moore, Index Fil. 368. 1862. Type. Mexico. Oaxaca: Cerro Zempoalte´petl, Liebmann s.n. [Pl. Mex. 2431, Fl. Mex. 5] (C!; isotypes K!, P!, US!).

Rhizomes long-creeping, ca. 1 mm diam.; rhizome scales ovate to linear-lanceolate, dull orange, peltate, entire, 3–6 mm long; fronds 1–2 cm distant, 5–8 ⫻ 1.2–1.6 cm; phyllopodia absent; stipes

ELAPHOGLOSSUM TECTUM

slender, ca. 1⁄2 the frond length, moderately clothed with ovate, dull orange scales 2–3 mm long; blades ovate-lanceolate or spatulate, chartaceous, bases broadly cuneate, apices obtuse; veins obscure, free, simple or once-forked, ca. 1 mm apart, at 60–70⬚ angle to costa; hydathodes absent; blade scales sparse, small, inconspicuous, 1–2 mm long, linear, often coarsely and irregularly toothed, especially toward the scale bases, more frequent on the abaxial surfaces; fertile fronds as tall as, or slightly taller than, the sterile fronds, with longer stipes, ca. 3⁄4 the length, blades spathulate, smaller than the sterile with a few scales along the abaxial costae but absent among the sporangia.

311

once-forked, 1.5–2.5 mm apart, at 50–60⬚ angle to costa; hydathodes evident; blade scales brown, linear-lanceolate with flared bases, evenly but sparsely distributed on blades, scarce along margins, 2 mm long, with short teeth; fertile fronds smaller than the sterile, blades ca. 3.5 ⫻ 1.2 cm, stipes ca 4⁄5 the fertile frond length, intersporangial scales absent.

Distribution. On moist banks and rocks in wet montane forests; 1700–2500 m. Mexico; Guat, Hond, CR, Pan; Hisp; Col, Ven.

Distribution. Epiphytic in cloud forests; (2000–)2450–3050 m. Mexico; Guat, Salv, CR, Pan; Cuba; Col, Ven, Ec, Peru, Braz, Bol. Selected Specimens Examined. Chis (Breedlove 22733, NY, 40447, DS). Gro (Lorea 3516, IEB; Rzedowski & McVaugh 118, NY). Hgo (Sa´nchez Mejorada 747, MEXU p.p.). Oax (Mickel 4587, 5266, NY). Ver (Cha´zaro & Castillo Ch. 6791, IEB; Mu¨ller 37, K, 39, NY p.p.).

Elaphoglossum squamipes represents a distinct group with long-creeping rhizomes and broad, dull orange, rhizome and stipe scales. It is very closely related to E. peltatum, differing from E. squamipes et aff. only in blade dissection.

51. ELAPHOGLOSSUM SQUARROSUM Elaphoglossum squarrosum (Klotzsch) T. Moore, Index Fil. 15, 367. 1857. Fig. 131E. Acrostichum squarrosum Klotzsch, Linnaea 20: 424. 1847. Type. Colombia [Venezuela]. Pa´ramo de Mucute´, Moritz 319 (B; isotypes BM!, K, frag. NY!, P!). Elaphoglossum chiapense A. R. Sm., Proc. Calif. Acad. Sci, ser. 4, 40: 220, f. 4B–D. 1975. Type. Mexico. Chiapas: Mpio. Rayon, 10 km above Rayon Mezcalapa along road to Jitotol, Breedlove 32649 (DS!).

Rhizomes moderately to long-creeping, to 2 mm diam.; rhizome scales maroon, lustrous, linear, to 6 mm long, shorttoothed, especially along the elongate tips; fronds distant, 7.5–22 ⫻ 1.4–2.2 cm; phyllopodia indistinct or absent; stipes 1⁄5–1⁄2 the frond length, with linear, maroon, toothed, appressed scales ca. 2 mm long; blades narrowly elliptic, papyraceous, bases cuneate, apices acute to acuminate; veins evident, free, simple or rarely

Specimens Examined. Chis (Breedlove 32429, DS, 55547, DS, NY). Gro (Lorea 3704, FCME, IEB). Me´x (Tejero-Dı´ez 3096, IZTA). Oax (Breedlove 65919, CAS; Lorence 4042, ENCB, MEXU, NY).

Among the subulate-scaled elaphoglossums in Mexico, E. squarrosum is distinct by its long-creeping rhizomes.

52. ELAPHOGLOSSUM TECTUM Elaphoglossum tectum (Humb. & Bonpl. ex Willd.) T. Moore, Index Fil. 15. 1857. Figs. 129A–D. Acrostichum tectum Humb. & Bonpl. ex Willd., Sp. Pl., ed. 4, 5(1): 102. 1810. Type. Venezuela. Caripe, Humboldt s.n. (B-Willd. 19520!; isotypes LE!, PHumb.!). Acrostichum elongatum Kunze, Linnaea 9: 31. 1834. Elaphoglossum elongatum (Kunze) T. Moore, Index Fil. 9. 1857. Type. Peru. “Ad Pampayaco, 1829, Herb. Kunze.” Poeppig s.n. (LZ, destroyed).

Rhizomes compact, short- to moderately creeping, 2–3 mm diam.; rhizome scales linear- lanceolate, dark reddish black, sclerotic, lustrous, ca. 5 mm long, with a few pale, weak hairteeth; fronds clumped to 3 mm apart, 60–80 ⫻ 1.7–2.6 cm; phyllopodia indistinct; stipes 1⁄3–1⁄2 the frond length, with scales appressed, peltate, round with dark centers and short hairteeth; blades linear-elliptic, chartaceous, bases narrowly cuneate, apices long-acuminate; veins obscure, free, simple or onceforked, mostly 0.7–1 mm apart, at ca. 80⬚ angle to costa; hydathodes absent; blade scales on adaxial surfaces white, appressed, round, peltate, toothed, deciduous with age, scales on the abaxial surfaces reduced to stellate hairs; fertile fronds about as long as the sterile or slightly longer, but with narrower blades and longer stipes more than 1⁄2 the frond length, scales

312

ELAPHOGLOSSUM

round, peltate on abaxial costae but absent among the sporangia; 2n ⫽164 (Jam).

Distribution. Terrestrial, epipetric, or less commonly epiphytic in wet montane forests; 1350–1550 m (1750–2500 m in Guat). Mexico; Guat, Hond, CR, Pan; Jam, Hisp; Col, Ven, Ec, Peru, Braz, Bol. Selected Specimens Examined. Chis (Breedlove 23172, DS; Lloyd 4064, ENCB p.p.). Oax (Mickel 5086b, 5096, NY). Ver (Bourgeau 2538, K, 3065, NY; Mu¨ller 42, NY; Purpus 2350, 6187, US; Smith 2017, UC).

Distribution. Terrestrial or epipetric in pine-oak forests; 1450 m. Mexico; Guat, Hond, Salv, CR; Col, Ven, Ec, Peru, Braz, Bol. Specimens Examined. Oax (Anderson 13212, MICH, NY).

Elaphoglossum tenuiculum is distinct by having virtually all the stipe and blade scales stellate with no scale body.

54. ELAPHOGLOSSUM TENUIFOLIUM

This species is readily distinguished by the round, peltate scales on the stipes and adaxial blade surfaces and stellate hairs on the abaxial blade surfaces.

Elaphoglossum tenuifolium (Liebm.) T. Moore, Index Fil. 368. 1862. Figs. 127N–P.

53. ELAPHOGLOSSUM TENUICULUM

Rhizomes short-to moderately creeping, 1–2 mm diam.; rhizome scales dense to sparse, ovate, orange or tan with varying degrees of dark, shiny, sclerotic streaking, ca. 3 mm long, deeply cordate, appearing nearly peltate, entire or with occasional small, irregular teeth; fronds 1–8 mm apart, 6–23 ⫻ 0.6–1 cm; phyllopodia distinct; stipes 1⁄4–2⁄3 the frond length, glabrous or with tan, ovate scales; blades linear-elliptic, chartaceous, bases attenuate, apices acuminate; veins obscure, free, simple or onceforked, ca. 1 mm apart, at 60–70⬚ angle to costa; hydathodes absent; blade scales sparse, reduced to minute stellate trichomidia or dots on abaxial surfaces, rare adaxially; fertile fronds similar to the sterile in blade size and shape, scales absent on abaxial surfaces.

Elaphoglossum tenuiculum (Fe´e) T. Moore ex Baker, Ann. Bot. (Oxford) 5: 491. 1891. Figs. 140E–H, J. Acrostichum tenuiculum Fe´e, Me´m. Foug. 10: 6, t. 29, f. 2. 1865. Type. Venezuela. Colonia Tovar, Fendler 272 (P!). Elaphoglossum stellatum Mickel, Amer. Fern J. 69: 101. 1979. Type. Guatemala. Santa Rosa: Volca´n Tecuamburro, Steyermark 33155 (F!; isotype US!).

Rhizomes short-creeping, ca. 2 mm diam.; rhizome scales linear-lanceolate, ca. 2 mm long, castaneous to dark brown, lustrous, subentire with occasional teeth near scale bases; fronds approximate, 9–21 cm long; phyllopodia distinct; stipes 1⁄4–1⁄3 the sterile leaf length, stramineous to tan with scattered orange-tan, stellate hairs; blades linear, 5–20 cm ⫻ 4–7 mm, chartaceous, bases cuneate, apices acuminate; veins barely visible, ca. 1 mm apart, at 60–70⬚ angle to costae; hydathodes absent; blade indument abaxially of sparse, stellate hairs, especially on the costae, those on the adaxial surfaces with narrow scale body; fertile fronds ca. equal to sterile fronds in length, narrower, and longer-stiped than the sterile, stipes 1⁄3–1⁄2 the frond length, blades 5–7 mm wide, stellate hairs on abaxial costae but not among the sporangia.

Acrostichum tenuifolium Liebm., Mexic. Bregn. 163 (reprint 11). 1849. Type. Mexico. Oaxaca: Cerro Zempoalte´petl, Liebmann s.n. [Pl. Mex. 2654] (C!; isotype US!).

ELAPHOGLOSSUM VIRIDE

313

stipes 1⁄8–2⁄5 the frond length, with scales linear, tan, ca. 2 mm long, ciliolate, spreading, others smaller, stellate; blades narrowly elliptic, chartaceous, bases narrowly cuneate, apices caudate; veins evident to obscure, free, simple to once-forked, ca. 1 mm apart, at ca. 70⬚ angle to costa; hydathodes absent; blade scales orange, linear, highly skeletonized, very long-ciliolate, the hairs 3–4 times as long as the scale body width, plane or the hairs turned upward, loosely covering the abaxial surfaces, adaxial surfaces glabrous with age, with narrowly lanceolate scales with long hair-teeth or substellate with long teeth; fertile fronds slightly shorter or longer than the sterile fronds, stipes 1⁄3–1⁄2 the frond length, with a few reduced scales on abaxial costae, but not among the sporangia. Distribution. Terrestrial on banks or epipetric in pine-oak cloud forests; 1800–3100 m. Mexico; Guat. Selected Specimens Examined. Chis (Ghiesbreght 305, NY). DF (Rzedowski 19495, ENCB). Dgo (Reeder et al. 3354, ARIZ, US). Gro (Rzedowski & McVaugh 251, ENCB). Hgo (Pringle 13637, US). Jal (McVaugh 26141, MICH). Me´x (Hinton 3275, NY). Mich (Hinton 13459, NY). Mor (Lyonnet 2466, US). Oax (Pringle 4916, NY, PH, UC, US). Ver (Riba et al. 353, MEXU, NY).

Although the general habit and the dark streaking of the rhizome scales are similar to those of Elaphoglossum affine and E. alansmithii, E. tenuifolium can be distinguished by its narrow blade and generally shorter rhizome length. See comments under those species. Mickel (in Davidse et al., 1995) included E. tenuifolium in E. minutum (Fe´e) T. Moore, but the latter is a larger plant (fronds 6–38 cm long) and its the rhizomes are short-creeping to compact with the fertile fronds slightly to far exceeding the sterile in length. It occurs from Costa Rica to Bolivia, Brazil, Jamaica, and Hispaniola.

55. ELAPHOGLOSSUM VESTITUM Elaphoglossum vestitum (Schltdl. & Cham.) Schott ex T. Moore, Index Fil. 369. 1862. Fig. 128Q. Acrostichum vestitum Schltdl. & Cham., Linnaea 5: 605. 1830. Type. Mexico. Veracruz: “Prope Jalapam. Aug.,” Schiede s.n. (B!). Also at B there is Schiede 735, without data. Acrostichum fulvum M. Martens & Galeotti, Me´m. Foug. Mexique 24, pl. 3, f. 2. 1842. Type. Mexico. Veracruz: Orizaba, Galeotti 6459 (BR!). Acrostichum laminarioides Fe´e var. α longipes E. Fourn., Mexic. Pl. 1: 67. 1872. Syntypes. Mexico. Veracruz: Barranca di Agua Santa, 3000 ped, Heller 188 (P!); in regione Orizabensi pr. Santa Ana, Bourgeau 3060 bis (P!); Huatusco, Schaffner 32 “in herb. Fe´e”; in alle Cordobensi pr. La Trinidad, Bourgeau 2110 (P!–2 sheets), 2110 bis (P!–3 sheets); Jalapa, aug. sp., Hahn s.n. (P!). Acrostichum laminarioides Fe´e var. β brevipes E. Fourn., Mexic. Pl. 1: 67. 1872. Syntypes. Mexico. Veracruz: Huatusco, Mu¨ller 386; in valle Cordobensi, Schaffner 28, 31; pr. Fortı´n, Bourgeau 2199 bis; pr. La Trinidad, Bourgeau 2110 quater (P!), 3060 (P!).

Rhizomes short-creeping, ca. 5 mm diam.; rhizome scales castaneous, lustrous, linear-lanceolate, toothed, 4–6 mm long; fronds clumped, 25–60 ⫻ 1.4–3.1 cm; phyllopodia distinct, short;

Distribution. Epiphytic in wet montane forests and oak cloud forests; 300–2000 m. Mexico. Selected Specimens Examined. Chis (Breedlove 21846, NY, 38979, DS; Kress et al. 749, MEXU). Hgo (Herrera & Riba 2205, MEXU). Oax (Croat 43916, NY; Mickel 4721, NY). Pue (Riba 12, MEXU). Qro (Dı´az B. & Carranza 6556, IEB, MEXU). Ver (Nee & Taylor 29466, NY; Purpus 6425, NY).

Elaphoglossum vestitum is most closely allied to E. rubescens, resembling it in rhizome scale color and texture, ciliate blade scales lightly covering the laminae, and lax fronds with caudate tips. The latter can be distinguished by the rhizome scales not (or only weakly) toothed, the stipes often with resin dots, and the blade scales with the teeth and margins upturned. These two species may be only varietally distinct, and several specimens from Chiapas overlap in morphology. Elaphoglossum auricomum belongs to the same complex, but has more abundant, spreading stipe scales, the rhizome scales are lighter in color, and the blades have acuminate apices. Some specimens of E. paleaceum have blades only lightly covered with scales, but they have black scales on the stipes and abaxial rachises.

56. ELAPHOGLOSSUM VIRIDE Elaphoglossum viride (E. Fourn.) C. Chr., Index Filic. 318. 1905. Figs. 141A–E.

314

ELAPHOGLOSSUM

Acrostichum viride E. Fourn., Mexic. Pl. 1: 66. 1872. Type. Mexico. Veracruz: Jalapa, Hacienda de las A´nimas, Hahn 455 (P! p.p., frag. NY!).

Rhizomes compact, short- to moderately creeping, 3–5 mm diam.; rhizome scales linear-lanceolate, orange-tan to dark brown, 2–3(–5) mm long, entire, margins inrolled, the cells isodiametric and distinctly concave, appearing subclathrate; fronds approximate to slightly spaced, 15–40(–65) ⫻ 15–20(–32) cm; phyllopodia distinct, ca. 1 cm long; stipes (1⁄5–)1⁄4–1⁄3(–2⁄5) the frond length, glabrous above the phyllopodia; blades narrowly elliptic, firm, narrowly cuneate at bases, long-acuminate at apices; veins scarcely visible, free, simple or once-forked, ca. 0.8 mm apart, at 60–65⬚ angle to costa; hydathodes absent; adaxial blade surfaces glabrous; abaxial blade surfaces with scattered to sparse, minute, stellate trichomidia and occasional black scales on costal bases; fertile fronds nearly as long as, to somewhat shorter than, the sterile and about the same shape and width, stipe 1⁄2–3⁄5 the frond length, intersporangial scales absent.

scales with prominent projections or dissected scales, extending onto the proximal portion of the costae, usually dark, especially at blade bases; blades narrowly elliptic, bases narrowly to broadly cuneate, apices acute to acuminate, subcoriaceous to coriaceous; veins obscure, free, simple to 2-forked, ca. 1 mm apart, at 70– 80⬚ angle to costa; hydathodes absent; adaxial surfaces glabrous; abaxial surfaces glabrous or with sparse to scattered, minute, stellate trichomidia; fertile fronds equalling or somewhat surpassing the sterile in length, stipes (1⁄6–)1⁄3–1⁄2 the frond length, blades 1 ⁄2–2⁄3 the sterile blade length, intersporangial scales absent.

Distribution. Terrestrial or epiphytic in dry to moist woods; 300–1700 m. Mexico. Specimens Examined. Chis (Breedlove 21972, NY). Oax (Geith 95-MO74, NY; Hallberg 1671, NY; Mickel 854, 1128, 1620, 7386, NY). Pue (Sa´nchez M. 640, US). Ver (Purpus 16580, US; Riba et al. 403, NY).

Distribution. Epiphytic in montane rain forests; 1050–2070 m. Mexico.

This species belongs to the E. sartorii complex but is distinct in the long, linear, orange rhizome scales.

Selected Specimens Examined. Oax (Hallberg 1509a, 1459, NY; Hellwig 217, 218, NY; Mickel 1545, 9702, NY; Torres C. 11376, 11713, MEXU). Ver (Houck 60, MICH; Trigos 2985, MEXU).

58. ELAPHOGLOSSUM YOURKEORUM

Elaphoglossum viride closely resembles E. sartorii, but the former is distinct by the inrolled rhizome scales with isodiametric concave cells.

Elaphoglossum yourkeorum Mickel, sp. nov. Type. Mexico. Oaxaca: Distrito Tuxtepec, 8 km s of Valle Nacional on Rte 175, Mickel 6678 (holotype NY!). Figs. 125D, G.

57. ELAPHOGLOSSUM XANTHOPODUM

Ab E. sartorii laminis fertilibus angustioribus et basibus latioribus laminarum sterilium differt. (Dedicated to Gertrude and George Yourke, who have been highly supportive of the New York Botanical Garden’s fern program.)

Elaphoglossum xanthopodum Mickel, sp. nov. Type. Mexico. Veracruz: Fink 107 (holotype NY!; isotypes NY!, US!–2 sheets). Figs. 125F, E. Ab E. sartorii squamis rhizomatis longibus linearibus aurantiacis abstans. (Greek, xanthos, gold, pous, foot, referring to the golden rhizome scales.)

Rhizomes compact, short-creeping, 5–8 mm diam.; rhizome scales linear, orange, mostly ca. 10 mm long, somewhat lustrous, with irregular hair-like lateral projections, especially near the scale apices; fronds clumped, approximate to 0.5 cm distant, 15– 48(–60) ⫻ 1.8–3.8(–5.5) cm; phyllopodia distinct, 1–1.5 cm long; stipes (1⁄10–)1⁄4–2⁄5 the frond length, clothed with scattered linear

Rhizomes compact, horizontal, 4–8 mm diam.; rhizome scales linear to linear-lanceolate, 5–10 mm, concolorous orange-tan, with irregular lateral hair-like processes; fronds approximate to 0.5 cm distant, 16–58 ⫻ 3–6 cm; phyllopodia distinct, 1–1.5 cm long; stipes 1⁄6–2⁄5 the frond length, with orange-tan scales proximally; blades elliptic, broadly cuneate at bases forming a 65–95⬚ angle, apices acute to acuminate, coriaceous; veins obscure, free, simple to 2-forked, ca. 1 mm apart, at 70–85⬚ angle to costa; hydathodes absent; adaxial blade surfaces glabrous; abaxial blades surfaces with linear scales at base (2–3 mm), also with sparse orange-tan trichomidia, mostly 1–2 mm; fertile fronds about equalling to somewhat surpassing the sterile, stipes 1⁄4–1⁄3 the

ENTEROSORA

frond length, blades linear-elliptic, 10–18 mm wide, intersporangial scales absent.

315

long acuminate (stellate hairs in E. muelleri), and scales of the proximal parts of the stipes entire. Elaphoglossum petiolatum has conspicuous entire scales on the stipes proximally and ciliate scales on the blades. The name E. venustum may apply to E. petiolatum.

4 5 . E N T E R O SO R A Enterosora Baker in Thurn, Timehri 5: 218. 1886. Type: Enterosora campbellii Baker.

Distribution. Terrestrial and epipetric in disturbed dry to moist oak woods on steep slopes; 300–1300(–2600) m. Mexico; Guat, Salv, Hond. Specimens Examined. Chis (Breedlove 21840, 22196, NY). Oax (Maya J. 2182, 3934, NY; Mickel 4751, 5226, 6524, 6678, NY; Rzedowski 21125, ISC). Guat (Thieme s.n. [Donn. Sm. 5667], NY; Heyde & Lux s.n. [Donn. Sm. 4077], NY). Salv (Seiler 578, NY). Hond (Morton 7565, NY; Yuncker 5848, NY).

This species is distinct among the E. sartorii complex in having very narrow fertile blades and very broadly cuneate bases on the sterile blades. Species of Uncertain Status Elaphoglossum trichomallum Rovirosa, Pteridogr. Sur Me´xcio 252. 1909. Type. Mexico. Chiapas: L. Martı´nez 1101 “in his herb.” The combination of cited characters (creeping rhizomes, scales reddish, linear-lanceolate, crisped sparse piliform scales on both sides of blades, stipes 14–22 mm, with gold-castaneous subulate scales, blades 27–39 ⫻ 3–6 cm, elongate lanceolate, papyraceous, attenuate at both ends; fertile fronds with blades 21 ⫻ 2 cm, costal scales similar to those of stipes, epiphytic) suggests it may be E. setigerum, but we have not located a type specimen. Elaphoglossum venustum T. Moore, Index Fil. 16. 1857. Acrostichum venustum Fe´e, Me´m. Foug. 8: 68. 1857. hom. illeg., non Liebm. 1849. Type. Mexico. Mecameca, la Puebla, Schaffner 332b, c, in 1855 (P?, not found). Both Christ (1899) and Christensen (1905) considered this to be synonymous with E. araneosum D. C. Eaton [⫽ E. muelleri of this treatment]. Christ’s characterization of E. venustum certainly applies to E. muelleri, but the description by Fe´e suggests otherwise: blade scales ciliate,

Epiphytic, occasionally lithophytic; rhizomes short-creeping, dorsiventral; rhizome scales tan to golden brown, non-clathrate, at times ciliate or glandular; fronds monomorphic; stipes subarticulate, sometimes flexed at apices; blades simple to broadly lobed to 4⁄5 the way to the midrib, non-pectinate, usually thick and spongy with numerous, large, intercellular air spaces; veins usually obscure, often variously areolate, especially near midrib, if free, forked when fertile; hydathodes absent, vein tips not visible, as viewed adaxially; indument of reddish brown setae 1–3 mm on stipes, rachises, costae, and sometimes laminae, setae often more numerous around the sori, indument also of branched, sometimes glandular hairs; sori round, oval, or sometimes elongate, superficial or commonly subimpressed or deeply imbedded in the lamina; soral paraphyses nearly always absent from receptacle and sporangia, if present then reduced to weak hairs smaller than developing sporangia; spores tetrahedralglobose; x⫽37. Enterosora is a small genus of eight species in the wet Neotropics, ranging from southern Mexico to Bolivia and Surinam, and in the Antilles; one or two additional species are known from Africa (Bishop & Smith, 1992), and there may be additional species in Madagascar. The genus is closely related to Ceradenia (Ranker et al., in prep.), and may be the sister group to that. Enterosora, like Ceradenia, is anhydathodous but differs in the often anastomosing veins (free in two of the three Mexican species!), less dissected blades (often subentire to shallowly lobed), in having sori commonly imbedded in thick, spongy, highly aerenchymatous blades, and especially in the absence of whitish, globose glands in the sori. References Bishop, L. E. & A. R. Smith. 1992. Revision of the fern genus Enterosora (Grammitidaceae) in the New World. Syst. Bot. 17: 345–362; Riba, R., L. Pacheco & E. Martı´nez S. 1987. New records of pteridophytes from the state of Chiapas, Mexico. Amer. Fern J. 77: 69–71.

Key to the Mexican Species of Enterosora 1. Sori deeply immersed in the lamina (both dried and living material), the laminar pits mostly as deep or deeper than wide, with vertical sides; blades thick, with setae thinly scattered over abaxial surfaces. ............................................................................................. 1. E. ecostata. 1. Sori superficial or in shallowly concave laminar depressions; blades moderately thick, with few to numerous setae over abaxial surfaces. 2. Veins mostly 1-forked; blade margins wavy; stipes often less than 1 cm long. ................................................................................... 2. E. parietina. 2. Veins pinnate; blade margins entire; stipes evident, 1–5 cm long. ...................................................................................................... 3. E. trichosora.

316

ENTEROSORA

1. ENTEROSORA ECOSTATA Enterosora ecostata (Sodiro) L. E. Bishop, Syst. Bot. 17: 348, f. 1A–C. 1992. Figs. 142A, B. Polypodium ecostatum Sodiro, Crypt. Vasc. Quit. 308. 1893. Type. Ecuador. Pichincha: Santo Domingo de los Colorados, Sodiro s.n. (QPLS? or Q?; probable isotypes NY!, UC!). Polypodium harrisii Jenman, Gard. Chron, ser. 3, 27: 241. 1900. Grammitis harrisii (Jenman) A. R. Sm., Proc. Calif. Acad. Sci, ser. 4, 40: 230. 1975. Type. Jamaica. Mabess River, Harris 7729 (K!; isotypes B!, US!). Polypodium nesioticum Maxon, Smithsonian Misc. Collect. 47: 410. 1905. Grammitis nesiotica (Maxon) Copel., Philipp. J. Sci. 80: 127. 1951. Glyphotaenium nesioticum (Maxon) Lellinger, Amer. Fern J. 74: 57. 1984. Type. Jamaica. Portland Parish: Vicinity of Vinegar Hill, Maxon 2773 (US!).

Rhizome scales orangish tan, lustrous, to 5 ⫻ 1.5 mm, entire or irregularly dentate, eciliate or glandular-ciliate; stipes brown, 2–8 cm ⫻ 0.3–0.7 mm, setose; blades thick, spongy, (5–)10–40 ⫻ (0.5–)0.8–1.5(–2) cm, linear, acute or acuminate proximally, margins typically crenate or crenately lobed, apices acuminate, at times entire or lobed to 2⁄3 the way to the midrib; indument of dense reddish setae 1.5–2 mm long at blade margins, also with setae clustered around the soral pits and scattered rather distantly over both blade surfaces; veins usually regularly areolate along midrib, often so toward margins; sori round or somewhat elongate, to 2 ⫻ 1 mm, 2–7 per vein group, in pits with vertical sides, mixed with setae, blades often without sori proximally, normally soriferous to apices.

Distribution. Epiphytic in lowland and lower montane rain forests; 300–1000 m. Mexico; Bel, Nic, CR; Jam, Hisp; Col, Ec. Specimens Examined. Chis (Breedlove 49972, 56650, CAS; Dressler 1695, MEXU, US; Martı´nez S. 18678, MEXU, TEX, UAMIZ, 18800, 23981A, MEXU; Valdivia Q. (P.V. 2433), UAMIZ).

This differs from the other Mexican species of Enterosora by the thicker, more succulent blades and by the sori embedded in deep pits; also, the fronds are ordinarily much longer than in either of the other two species. Enterosora ecostata is probably most closely allied to E. percrassa (Baker) L. E. Bishop, from

Costa Rica and Panama, and hybridizes with it in that area (Bishop & Smith, 1992).

2. ENTEROSORA PARIETINA Enterosora parietina (Klotzsch) L. E. Bishop, Syst. Bot. 17: 357, f. 2J–L. 1992. Figs. 142D, E. Polypodium parietinum Klotzsch, Linnaea 20: 373. 1847. Grammitis parietina (Klotzsch) Fe´e, Me´m. Foug. 5: 233. 1852. Type. Venezuela. Aragua: Colonia Tovar, Moritz 253 (B!, frag. NY!; isotypes BM!, K!, P!). Polypodium rosulatum Christ in J. Bommer & Christ, Bull. Herb. Boissier, se´r. 2, 4: 662. 1896. Grammitis rosulata (Christ) Lellinger, Proc. Biol. Soc. Wash. 89: 715. 1977. Type. Costa Rica. Rı´o Naranjo, Pittier 7953 (BR, photos BM!, US!). Polypodium yarumalense Hieron., Bot. Jahrb. Syst. 34: 499. 1904. Grammitis yarumalensis (Hieron.) Proctor, Bull. Inst. Jamaica, Sci. Ser. 5: 36. 1953. Type. Colombia. Antioquia: Yarumal, Lehmann 7390 (B; isotypes BM!, K!, US!).

Rhizome scales castaneous, lustrous, linear-lanceolate, 0.5–3 ⫻ 0.2–0.3 mm, entire or sparingly glandular-ciliate, sometimes ending in a seta; stipes dark brown, 2–5(–30) ⫻ 0.2–0.4 mm, setose; blades moderately thick, spongy, 5–12(–30) ⫻ 0.3–0.8 cm, linear, attenuate proximally, margins wavy, apices acute to obtuse; indument of reddish, nodose setae 1–2 mm long, longer and sparser on abaxial surface, most concentrated near blade margins; veins free, (0–)1–2-forked, midvein slightly paler than the surrounding lamina; sori round, ca. 1 mm diam., superficial, in single rows on both sides of midvein, midway between midvein and margin, mixed with setae.

Distribution. Epiphytic in wet montane forests of Atlantic slope; 1050–1400 m. Mexico; Guat, Hond, CR, Pan; Jam, Hisp; Col, Ven, Ec, Peru. Specimens Examined. Oax (Hallberg 1515, NY, UC; Mickel 1064, NY, 5739, NY, UC).

This species, with sori in one row on each side of the midrib, is superficially similar to Ceradenia jungermannioides, which see for discussion. Enterosora parietina also resembles E. ecostata, but that species has longer stipes, larger and thicker-textured blades,

EQUISETUM

areolate venation, and deeply embedded sori in several rows on each side of the midrib.

3. ENTEROSORA TRICHOSORA Enterosora trichosora (Hook.) L. E. Bishop, Syst. Bot. 17: 357, f. 2G–I. 1992. Fig. 142C. Polypodium trichosorum Hook., Sec. Cent. Ferns, pl. 12. 1861. Grammitis trichosora (Hook.) Ching, Bull. Fan Mem. Inst. Biol. 10: 241. 1941. Type. Ecuador. Pichincha: Archidona, Jameson 349 (K!, frag. B!). Polypodium repletum Christ, Bull. Herb. Boissier, se´r. 2, 7: 260. 1907. Type. Costa Rica. 1905, Werckle´ s.n. (P!; isotype B!). Polypodium minusculum Maxon, Contr. U.S. Natl. Herb. 13: 11. 1909. Grammitis minuscula (Maxon) Copel., Philipp. J. Sci. 80: 128. 1951. Type. Guatemala. Alta Verapaz: Coba´n, Tu¨rckheim II.1987 (US!; isotypes GH!, NY!, P!, UC!, US!).

Rhizome scales tan-gold, dull to sublustrous, to 3 ⫻ 0.8 mm, entire or irregularly dentate, sparsely ciliolate with scattered, unicellular glands; stipes brown, 1–5 cm ⫻ 0.2–0.4 mm, setose; blades moderately thick, spongy, 2–10 ⫻ 0.5–1.5 cm, linear-elliptic, acute proximally, margins entire (ours) or broadly and rather deeply crenate, apices rounded to acute; indument of castaneous or dark brown setae mostly 1–2 mm on blade margins, setae scattered over the adaxial surface, abaxially the setae mostly on the midrib and around the sori; veins free, main lateral veins with 3–7 pinnately arranged branches; sori round to slightly oblong, to 2 mm ⫻ 1.5 mm on basal vein branches, 1–3 per vein group, superficial or somewhat immersed in shallow pits, absent toward blade bases.

317

Specimens Examined. Chis (Pacheco 1300, UAMIZ, UC).

In Mexico, this species is known only from the report by Riba et al. (1987). The only other simple-bladed, entire grammitids in Mexico are Ceradenia jungermannioides, Cochlidium linearifolium, and Cochlidium rostratum, all of which have much narrower, linear blades. The other two species of Enterosora in Mexico commonly have repand or shallowly lobed blades.

46 . E Q U I SE T U M Equisetum L., Sp. Pl. 2: 1061. 1753. Lectotype (chosen by Farwell, Mem. New York Bot. Gard. 6: 464. 1916): Equisetum arvense L.; E. fluviatile L. is sometimes regarded as lectotype; see discussion and additional synonymy by Pichi Sermolli, Webbia 26: 179, 1971. Terrestrial; rhizomes subterranean, perennial, long-creeping, branched; aerial stems erect, hollow, grooved, jointed, simple or with whorled branches, generally with a large central canal and smaller vallecular (internal to the stem grooves) and carinal (internal to the stem ridges) canals; leaves small, white to black, whorled, fused to form a sheath at the nodes; aerial axes perennial or annual, stems monomorphic (in ours) or dimorphic; stem ridges with silica tubercles, grooves with narrow or broad bands of stomata; sporangia borne abaxially on tightly packed peltate sporophylls, those forming the terminal strobili on the main stems and branches; spores homosporous, green, each with four elaters; x⫽108. Equisetum is terrestrial or of moist to swampy lands, forming small to extensive stands through the wide-creeping, much branched rhizomes. The genus is widespread but with only 15 species, mostly of boreal and north-temperate regions. Twelve species occur in North America, three in Mexico. Hybridization is common in the genus, some hybrids being as common as one or more of the parents. Recent molecular evidence suggests that Equisetum is most closely related to basal ferns (Pryer et al., 2001). We are grateful to R. L. Hauke for identification of Mexican specimens of Equisetum and for valuable discussions regarding their interpretation. References

Distribution. Epiphytic in montane rain forests (pine-oakLiquidambar); 1450 m. Mexico; Guat, CR; Ec.

Hauke, R. L. 1963. A taxonomic monograph of the genus Equisetum subgenus Hippochaete. Beih. Nova Hedwigia 8: 1–123; Tejero-Dı´ez, D. & S. AguilarRodrı´guez & S. Flores-Maya. 1998. El ge´nero Equisetum L. en el occidente del estado de Me´xico, Me´xico. Acta Bot. Mex. 44: 7-25.

Key to the Mexican Species of Equisetum 1. Stems producing regular whorls of branches. ........................................................................................................................................ 3. E. myriochaetum. 1. Stems normally unbranched. 2. Stem ridges conspicuously tuberculate; strobili sharply apiculate; leaf sheaths about as long as wide, at maturity stem sheaths all with black to dark brown girdles at distal edge and at the middle, ash gray between the two girdles; teeth each with distinct articulation line. ............................................................................................................................................................................. 1. E. hyemale var. affine. 2. Stem ridges flattened to convex, smooth to slightly tuberculate; strobili rounded or short-apiculate; leaf sheaths longer than wide, green or dark-girdled rarely at base of stems; teeth without articulation line. ..................................................................... 2. E. laevigatum.

318

EQUISETUM

1. EQUISETUM HYEMALE VAR. AFFINE Equisetum hyemale A. Braun ex Engelm. L. var. affine (Engelm.) A. A. Eaton, Fern. Bull. 11: 111. 1903. Figs. 143A–E, 144F, G, a. Equisetum robustum A. Braun ex Engelm. γ affine Engelm., Amer. J. Sci. Arts 46: 88. 1844. Equisetum hyemale L. subsp. affine (Engelm.) Calder & Roy Taylor, Canad. J. Bot. 43: 1387. 1965. Type. U.S.A. Grassy clayey banks of Mississippi River below Jefferson Barracks, Aug 1843, Engelmann s.n. (MO; isotype US!). For full synonymy, see Hauke (1963).

Stems erect, unbranched, bluish green, to 2.2 m tall, 5–10 mm diam., unbranched except on damaged plants; sheaths with length to width ratio of ca. 1:1 with two dark girdles, one at distal edge, the other at middle of sheath, ash gray between; stomata in single lines, one line on each side of a groove; ridges 14–50, with tubercles; stem cross sections with carinal collenchyma extending to the vascular bundles, vallecular collenchyma absent to small; endodermis of double common type; strobili terminal on stems, ca.15 mm long, each with a black apiculum; 2n⫽216 (Canada, USA).

Distribution. Usually in moist areas by streams or seeps, oak forests, pine-oak forests; 1600–3000 m. Widespread in Canada, USA; Mexico; Guat; northeasternmost Asia. Selected Specimens Examined. BCN (Thorne 58181, MEXU). BCS (Brandegee s.n., 25 Mar 1892, UC). Chih (LeSueur 1149, TEX). Chis (Little & Sharp 9911, UC). Coah (Carranza P. 331, ENCB). DF (On˜ate E. 8, FCME). Dgo (Gonza´lez & Gonza´lez 613, CHAPA, ENCB). Gro (Viveros J. L. & Casas A. 292, CAS). Gto (Dı´az Barriga 7408, IEB, MEXU). Hgo (Rzedowski 22193, ENCB, NY). Jal (Iltis et al. 1372, UC, WIS). Me´x (Pringle 3329, NY, UC, US). Mich (Rzedowski 42388, ENCB). Mor (Bonilla 523, FCME, MEXU). Nay (Gonza´lez 692, MEXU). NL (Mu¨ller 2805, UC). Oax (Mickel 6295, NY, UC). Pue (Flores C. 670, ENCB, MEXU). Qro (Carranza & Zamudio 4457, IEB). SLP (Emanuel 3104, IEB). Sin (Breedlove 18432, CAS). Son (Gentry 2108, UC, US). Tam (Gonza´lez-Medrano 2936, MEXU). Ver (Rzedowski 19046, ENCB). Unverified, Doubtful, or Mistaken Reports. Ags (Garcı´a 2388, HUAA, and three other collections, cited by Siqueiros-Delgado & Gonza´lez-Adame, 2004, but not verified).

Equisetum hyemale var. hyemale occurs in Eurasia and has stem ridges always bituberculate and teeth soon falling (vs. convex to bituberculate and teeth persistent to falling in var. affine). The dark banding of the sheaths is distinctive of E. hyemale, but often the banding is irregular or even absent. This character is quite variable in Mexico and has led some to believe Equisetum laevigatum to be more widespread than it actually is. Young E. hyemale has longer than usual sheaths that often lack the lower dark bands, resembling those of E. laevigatum. Malformed, non-green spores, evidenced by whitish powdery material in the sporangia and unopened strobili, are evidence of hybrids. Hybrids and juvenile specimens are often difficult to identify with confidence. Equisetum hyemale hybridizes with E. laevigatum (⫽ E. ⫻ferrissii Clute) in northwestern Mexico (BCN, Chih, Dgo) in areas of overlap. Stem cross sections show vallecular collenchyma prominent (as in E. laevigatum), often extending to the vallecular canal (as in E. laevigatum), and the carinal collenchyma usually extending nearly to the vascular bundles (as in E. hyemale) (Fig. 144F). Equisetum hyemale ⫻ myriochaetum (E. ⫻haukeanum Mickel) is widespread (Sin, Jal, Mich, Gro, Qro, Mor, Me´x, NL, SLP, Hgo, Ver, Pue, Oax, Chis, Guat) and nearly as common as its parents, both of which have broad distribution. Stems are tall, unbranched to irregularly and sparsely branched or much branched, to 2.5 m tall, 3.5–8.5 mm diam., with 14–36 ridges; stem sheaths with length to width ratio of 1:1.7; proximal sheaths with two dark bands and ash-gray between; distal sheaths each with single marginal band and same color as stem; teeth mostly deciduous; branch ridges with irregular to saw-toothed tubercles; strobili strongly apiculate, with malformed spores. This hybrid is extremely variable, sometimes closely resembling one or the other parent. The irregular and mixed tubercles/saw teeth can be used to determine E. ⫻ haukeanum. The nodal sheaths resemble those of E. hyemale near the base and those of E. myriochaetum distally and distinguish both branched and unbranched stems. Generally, the irregular branching of E. ⫻haukeanum is not found in E. myriochaetum. Stem cross sections (Fig. 144e) show carinal collenchyma touching the vascular bundles, the vallecular collenchyma varying from obsolete to touching the vallecular canal, and endodermis of the double common type.

2. EQUISETUM LAEVIGATUM Equisetum laevigatum A. Braun, Amer. J. Sci. Arts 46: 87. 1844. Lectotype (chosen by Hauke, 1963: 67). U.S.A. “N. Riehl No. 279. Bords immediats du Mississippi, St. Louis, Mo., Juin 1840” (NY!). Figs. 144A–E, c. Equisetum kansanum J. H. Schaffn., Ohio Naturalist 13: 21. 1912. Type. U.S.A. Kansas: Clay County, Bloom Township, Engelmann s.n. (OS; isotype US).

Stems 20–150 cm tall, 3–7 mm diam., unbranched except on damaged plants, yellowish green; sheaths with length:width ratio of 2:1 with black rim at distal edge, otherwise green; stomata in

EQUISETUM MYRIOCHAETUM

single lines, one line on each side of a groove; ridges 10–32, flattened to convex, smooth to tuberculate; sheaths green, elongate, 7–15 ⫻ 3–9 mm; teeth 10–32, each without articulation line but articulate and shedding early, leaving a dark rim on each sheath; stem cross sections (Fig. 144c) with prominent vallecular collenchyma extending to the vallecular canal; carinal collenchyma of lesser extent; endodermis of the double common type; strobili terminal on stems, ca. 10–20 mm long, apices rounded or each with a small apiculum; 2n⫽216 (USA).

319

bling saw teeth and pointing towards apices of stems; strobili terminal on branches and main stems blunt, or with slight apiculum; stem strobili to 30 mm long, 12 mm diam.; branch strobili 10 mm long, 4 mm diam.

Distribution. Terrestrial in swampy or marshy places, usually seepage areas, usually more or less exposed, in pine-oak forests, grassy flats, lower montane rain forests; (350–)700–2420 m. Mexico; Guat, Hond, Nic, CR, Pan; Col, Ven, Ec, Peru. Distribution. Terrestrial in pine-oak forests; 1130–1850 m. W USA & Can; Mexico. Specimens Examined. BCN (Chambers 524, UC; Thorne 32038, UC; Wiggins & Demaree 4860, NY, 4871, UC, US). BCS (Thomas 7897, US). Dgo (Benı´tez Paredes 887, MEXU). Son (Wolf 2510, CAS). Unverified, Doubtful, or Mistaken Reports. NL (Aguirre C. 639, UNL, cited by Aguirre Clavera´n, 1983, but this is E. hyemale).

Equisetum laevigatum is limited in Mexico to the northwestern states and is distinct by the stem ridges lacking prominent tubercles, cones rounded apically, teeth without conspicuous articulation, and prominent vallecular collenchyma. It crosses with Equisetum hyemale (E. ⫻ferrissii), which see for details. This species is an annual in Canada and the northern United States, but is perennial in Mexico.

3. EQUISETUM MYRIOCHAETUM Equisetum myriochaetum Schltdl. & Cham., Linnaea 5: 623. 1830. Type. Mexico. Veracruz: Misantla, “in sylvis humidis,” Schiede & Deppe 833 (B!). Figs. 143F–H, J, 144H, J, b. For additional synonymy, see Hauke (1963). Aerial stems to 8 m tall (mostly 2–4 m tall), with regular whorls of branches; stems 3–23 mm diam., with 16–18 ridges; stem cross sections with both carinal and vallecular collenchyma well developed, the former usually extending to the vascular bundles and the latter to the vallecular canals; endodermis commonly double; sheath length to width ratio about 1:1.5, with dark margins on distal edge, otherwise green, occasional dark sheath at base of large specimens; stomata in one line on each side of grooves; branches with 6–8 ridges, ridges bearing tubercles resem-

Selected Specimens Examined. Chis (Breedlove 21386, NY). Col (Gonza´lez s.n., 25 May 1977, IBUG). Gro (Lorea 1138, FCME). Gto (Pe´rez Calix & Carranza 2898, IEB). Hgo (Gimate 842, NY). Jal (Cochrane et al. 5186, IBUG, UC, WIS). Me´x (Hinton 2402, NY, US). Mich (Moore et al. 5739, UC). Nay (McVaugh & Koelz 762, NY). NL (Sa´nchez et al. 438, MEXU). Oax (Hallberg 1020, ENCB, MICH, US). Pue (Kelly 567, US). Qro (Servı´n 1375, IEB). Sin (Vega A. 863, IEB). SLP (Pringle 3294, BR, NY, UC, US). Tam (Gonza´lez Medrano 16961, MEXU). Ver (Copeland herb. 164, GH, MICH, UC, US).

Equisetum myriochaetum has flattened sheath segments (also found in E. ⫻haukeanum) and often has poor tubercle development. Generally, large Mexican horsetails with regular whorls of branches belong to E. myriochaetum. Equisetum myriochaetum crosses with E. giganteum to form E. ⫻schaffneri: Equisetum ⫻schaffneri Milde, Verh. Zool.-Bot. Ges. Wien 11: 345. 1861 (Fig. 144d). Type. Mexico. Veracruz: Orizaba, Schaffner 315 (B?). For synonyms, see Hauke (1963). Its stems have regular whorls of branches and are 2–4 m tall (to 7 m tall in Costa Rica), 3–22 mm diam., with 12–48 ridges; vascular bundles each with its own endodermis; carinal collenchyma extending to vascular bundles; vallecular collenchyma extending to the vallecular canal or nearly so; sheaths green to brown at bases; stomata in bands of 1–2(–3) in each groove; branches with 6–8 ridges with saw teeth or irregular tubercles; strobili terminal on branches and main stems, acute or each with a slight apiculum; spores malformed. Equisetum ⫻schaffneri rarely occurs from Mexico to Peru, in Mexico at 500–950 m elevation (Tam, Ver, Pue, Oax, Chis). Equisetum giganteum is not known to occur in Mexico, and Hauke (1963) suggested that E. ⫻schaffneri is dispersed by an occasional good spore. Equisetum ⫻schaffneri can be distinguished by its malformed, non-greens spores, intermediacy in number of rows of stomata (1–2 rows on each side of grooves); E. giganteum with broad stomatal band, 3–4 rows (vs. E. myri-

320

ERIOSORUS

ochaetum in one row on sides of each groove), branch ridges with flattened tubercles (vs. stomata in single lines and branch ridges with curved saw teeth in E. myriochaetum), and endodermis around individual bundles, as in E. giganteum (vs. continuous in E. myriochaetum).

4 7 . E R I O S O R US Eriosorus Fe´e, Me´m. Foug. 5: 152, t. 13B, f. 1. 1852. Type: Eriosorus scandens Fe´e [⫽ Eriosorus aureonitens (Hook.) Copel.]. Terrestrial; rhizomes short-creeping; rhizome scales tan to black, spreading or appressed, with long, catenate or bristle hairtips; fronds small to large, often scandent or erect, monomorphic, not articulate; stipes mostly brown to atropurpureous, hairy to glandular, rachises sometimes flexuous; blades pinnate-pinnatifid to quadripinnate; pinnae mostly at right angles to rachises, rarely retroflexed, laminae usually hairy or glandular; segments toothed or linear and each with a single vein; veins free; sori along the veins of ultimate segments, without indusia; paraphyses absent; spores tetrahedral-globose, without perispore but generally with an equatorial flange, often with ridges; x⫽87, prob. from x⫽29.

castaneous, glabrous except for scattered short hairs 0.5–0.8 mm long proximally; blades 4–5-times pinnate, ca. 10–15 cm broad; rachises flexuous, hairs sparse to abundant, often clumped in axils of blade branches; pinnule rachises retroflexed; ultimate segments narrow, 0.5 mm wide, apices emarginate, thin hairs scattered along veins on adaxial and abaxial surfaces; vein endings enlarged; spores tan; 2n⫽174 (CR). The highly dissected blades with linear segments strongly resemble those of Odontosoria schlechtendalii. Sterile material of Eriosorus flexuosus can be distinguished by having occasional blade hairs and rhizome indument of hairs, whereas Odontosoria has glabrous blades and scaly rhizomes. Key to the Mexican Varieties of Eriosorus flexuosus 1. Rhizome hairs or scales dark brown to black; fronds scandent, herbaceous or rigid. ..................................... 1a. var. flexuosus. 1. Rhizome scales light brown or tan; fronds erect and delicately herbaceous. .................................................... 1b. var. galeanus.

1a. Eriosorus flexuosus var. flexuosus

Figs. 145C, D.

Distinct by its dark rhizome indument and scandent fronds.

Eriosorus is a wholly American genus of 25 species of high elevation cloud forests in forested to open areas. It is close to Jamesonia, and in some areas resembles it and hybridizes with it. There are only two species in Mexico. Eriosorus is distinct in Mexico by its gymnogrammoid sori, 3–4-pinnate blades, and hairy rhizomes. Reference Tryon, A. F. 1970. A monograph of the fern genus Eriosorus. Contr. Gray Herb. 200: 54–174.

Key to the Merxican Species of Eriosorus 1. Blades 4–5-times pinnate, skeletonized, the ultimate segments only 0.5 mm wide; rachises flexuous, hairs sparse to abundant, often clumped in axils of blade branches. ............................................................................. 1. E. flexuosus. 1. Blades 3–4-times pinnate, the ultimate segments generally 1–1.5 mm wide; rachises subflexuous, pubescent. .............................................................................. 2. E. hirtus.

1. ERIOSORUS FLEXUOSUS Eriosorus flexuosus (Desv.) Copel., Gen. Fil. 58. 1947. Gymnogramma flexuosa Desv., Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck. Gessammten Naturk. 5: 306. 1811. Type. “America equinoctali” [from specimen]. s. coll. s.n. (P!-Desv., photo GH). Grammitis flexuosa Kunth, Nov. Gen. Sp. (Folio ed.) 1: 4. 1815 [1816]; Nov. Gen. Sp. (quarto ed.) 1: 5. 1815 [1816], hom. illeg., non Humb. & Bonpl., 1809. Type. Venezuela. Near Caracas, Humboldt & Bonpland s.n. (B-Willd. 19597!, photo GH!; isotypes P–3 sheets, photo GH!).

Rhizomes creeping, slender, 3 mm diam.; rhizome hairs scalelike, with thick, inflated cells, reddish brown, 2 ⫻ 0.5 mm; fronds scandent or scrambling, 1–3 m; stipes 1⁄3 or less the frond length,

Distribution. Rocky banks in wet montane forests, especially in open areas by roads, streams; 1650–3025 m. Mexico; Guat, Hond, Nic, Salv, CR, Pan; Hisp; Col, Ven, Guy, Ec, Peru, Braz, Bol. Selected Specimens Examined. Chis (Breedlove 22797, DS; Matuda 5241, F, GH, MEXU, US; Te´llez & Pe´rez G. 13212, MEXU). Gro (Breedlove 36022, CAS, MEXU, NY; Hinton 14309, NY, US; Lorea 2279, IEB). Oax (Mickel 4244, 6744, NY; Rzedowski 30621, 33363, NY).

1b. Eriosorus flexuosus var. galeanus A. F. Tryon, Contr. Gray Herb. 200: 164. 1970. Type. Mexico. Guerrero: Puerto Gallo, del Cerro Teotepec, Rzedowski 18594 (GH; isotype NY!) Figs. 145E, F. Distinct by its pale rhizome indument and erect fronds.

GLEICHENELLA

321

Specimens Examined. Chis (Breedlove 31948, 42747, DS, MEXU, 65750, 66059, CAS; Purpus 6722, NY, 7219, UC, US; Xolocotzi & Sharp X-470, US). Oax (Mickel 4239, NY). Unverified, Doubtful, or Mistaken Reports. Gro (Ramı´rez 21, ENCB, IEB, cited by Lorea-Herna´ndez & Vela´zquez M., 1998, but not verified).

Mexican material corresponds to var. hirtus. The broader, more pubescent segments distinguish E. hirtus from E. flexuosus.

48 . GL E I C H E N E L L A Distribution. On banks in pine and oak forests; 2400–2570 m. Mexico. Specimens Examined. Gro (Hinton 14221, 14309, GH, US; Ramı´rez Santos 21, IEB; Rzedowski & McVaugh 125, MEXU, NY; Vela´squez 752, FCME). Oax (Calzada 19459, NY).

2. ERIOSORUS HIRTUS Eriosorus hirtus (Kunth) Copel., Gen. Fil. 58. 1947. Figs. 145A, B. Grammitis hirta Kunth, Nov. Gen. Sp. (folio ed.) 1: 4. 1815 [1816]. Type. Venezuela. Silla de Caracas, Humboldt & Bonpland s.n. (P!, photo GH!; frag. B!). Psilogramme chiapensis Maxon, Bull. Torrey Bot. Club 42: 81. 1915. Type. Mexico. Chiapas: Cerro del Boquero´n, Purpus 6722 (US!; isotypes BM, F, GH!, HBG, P, US!). For additional synonyms, see A. Tryon (1970).

Rhizomes creeping, slender, 3 mm diam.; rhizome hairs bristlelike, thick, 1.5 mm long, reddish brown; fronds erect, clumped, 10–67 cm long; stipes 1⁄3 or less the frond length, castaneous, pubescent, hairs 0.5–1 mm long; blades 3–4 times pinnate, deltate, ca. 10–20 cm wide; rachises subflexuous, pubescent, hairs 0.5–1 mm long; ultimate segments 1–1.5 mm wide, with emarginate apices, thin, hairs 0.5–1 mm long on adaxial and abaxial surfaces; vein endings enlarged; spores reddish brown; 2n⫽348 (Chis).

Gleichenella Ching, Sunyatsenia 5: 276. 1940. Gleichenia Sm. sect. Acropterygium Diels, Nat. Pflanzenfam. 1(4): 353. 1900. Dicranopteris Bernh. sect. Acropterygium (Diels) Underw., Bull. Torrey Bot. Club 34: 251. 1907. Dicranopteris Bernh. subg. Acropterygium (Diels) Holttum, Reinwardtia 4: 261. 1957. Type: Gleichenella pectinata (Willd.) Ching [⬅ Mertensia pectinata Willd.]. Terrestrial; rhizomes long-creeping, cord-like, solenostelic, bearing dense pluricellular, unbranched hairs; fronds monomorphic, erect or scandent, to several meters long; blades pseudodichotomous, with pinnae consistently and repeatedly unequally branching, i.e., the larger branch alternately to one side of the rachis and then the other, the minor branches also unequally branched, lacking basiscopic accessory branches at the forks; dormant buds covered by pluricellular hairs, also often with a pair of foliaceous, stipule-like outgrowths, at least within the proximal forks; penultimate divisions pectinate, bearing narrowly lanceate to linear segments; indument absent abaxially, or of sparse stellate hairs mainly on the axes; veinlets free, 2–4-forked; sori abaxial with (6–)8–16(–25) pear-shaped sporangia; paraphyses absent; sporangia with oblique annuli; indusia absent; spores bilateral, ellipsoid, with surface rugulate and of fine rodlets; x⫽43. Most authors include Gleichenella in Dicranopteris, as subg. Acropterygium. As Østergaard Andersen and Øllgaard (2001) pointed out, differences in branching pattern, stele type, chromosome number, and spore morphology make Dicranopteris and Gleichenella at least as distinctive as the other recognized genera (Gleichenia s.str., Sticherus, Diplopterygium, and Stromatopteris) in the family. An alternative classification is to combine all genera in the family (excepting Stromatopteris) into a single genus Gleichenia, a solution not generally adopted even in conservative accounts of the family. Gleichenella is monotypic, the range that of the single species. Affinities of Gleichenella and the family Gleicheniaceae have been shown in recent molecular analyses to be with the basal leptosporangiate ferns, arising above Osmunda in phylogenetic trees, but below the schizaeoid ferns and tree ferns (Hasebe et al., 1995; Pryer et al., 1995, 2001). The fossil history of the family dates back at least as far as the Jurassic (Gandolfo et al., 1997). References

Distribution. Wet pine-oak forests; 2000–2600 m. Mexico; Guat, Hond; Cuba; Col, Ven, Ec, Peru, Bol.

Ching, R. C. 1940. On the genus Gleichenia. Sunyatsenia 5: 269–288; Gandolfo, M. A., K. C. Nixon, W. L. Crepet & G. E. Ratcliffe. 1997. A new fossil fern

322

HECISTOPTERIS

assignable to Gleicheniaceae from Late Cretaceous sediments of New Jersey. Amer. J. Bot. 84: 483–493; Holttum, R. E. 1957. Morphology, growth-habit, and classification in the family Gleicheniaceae. Phytomorphology 7: 168–184; Maxon, W. R. 1909. Gleicheniaceae. N. Amer. Fl. 16: 53–63; Nakai, T. 1950. New classification of Gleicheniales, etc. Bull. Natl. Sci. Mus., Tokyo 29: 1–71; Østergaard Andersen, E. & B. Øllgaard. 1996. A note on some morphological terms of the leaf in the Gleicheniaceae. Amer. Fern J. 86: 52–57. 1996; Østergaard Andersen, E. & B. Øllgaard. 2001. 10. Gleicheniaceae. In: G. Harling & L. Andersson (eds.), Flora of Ecuador 66: 105–170.

1. GLEICHENELLA PECTINATA Gleichenella pectinata (Willd.) Ching, Sunyatsenia 5: 276. 1940. Figs. 146A–E. Mertensia pectinata Willd., Kongl. Vetensk. Acad. Nya Handl. 25: 168, t. 4. 1804. Gleichenia pectinata (Willd.) C. Presl, Reliq. Haenk. 1(1): 71. 1825. Dicranopteris pectinata (Willd.) Underw., Bull. Torrey Bot. Club 34: 260. 1907. Type. Venezuela. [Distrito Federal:] Caracas, Bredemeyer s.n. (B-Willd. 19465!). Gleichenia nitida C. Presl, Reliq. Haenk. 1(1): 70. 1825. Type. Mexico. Haenke s.n. (PRC; frags. NY!, US!). Mertensia glaucescens Humb. & Bonpl. ex Willd. var. mexicana Fe´e, Me´m. Foug. 11: 121. 1866. Type. Mexico. Veracruz: “Sabannes de Mirador,” Galeotti 6402 (P!; isotype BR!).

Rhizomes 3–6 mm diam., with deciduous reddish brown septate hairs 2–3 mm long, surface scabrous when hairs shed; stipes stramineous to light brown, glabrous, dull; frond apices dormant (rarely continuing), pinnae forking unequally, appearing as if each pinna has one flexuous main axis with alternating, unequally branched side branches; forks containing a tuft of reddish brown or castaneous hairs, these hairs simple or usually stellate or branched at the very base, also some forks (within fork) with a pair of small, entire to pinnatifid “stipules,” lacking accessory pinnae proximal to the fork; penultimate segments narrowly elliptic; ultimate segments linear, chartaceous to coriaceous, often emarginate at the tip, usually glabrous or glabrescent, often glaucous abaxially, margins revolute at least on fertile segments; sori inframedial between segment margins and midveins; 2n⫽86 (Chis, Jam, Trin).

Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol. Selected Specimens Examined. Chis (Breedlove 42093, DS, MEXU; Calzada et al. 9159, UC, XAL). Gro (Lorea 2849, FCME, IEB, MEXU). Hgo (Moore 3022, UC). Jal (Castillo et al. 10349, MEXU). Mex (Hinton 7543, CAS, MEXU, NY, US). Oax (Mexia 9243, CAS, GH, MICH, NY, UC, US; Mickel 6137, ENCB, MEXU, NY, UC). Pue (Ma´rquez R. et al. 708, ENCB, MEXU, XAL). Tab (Gonza´lez S. 3, UAMIZ). Ver (Copeland herb. 7, DS, MEXU, UC; Orcutt 3202, DS, MEXU, NY).

Gleichenella pectinata differs from Dicranopteris flexuosa by lacking accessory pinnae proximal to the forks (not to be confused with the pair of “stipules” subtending bud, but within the branching of the axes), pinnae forking unequally to produce alternate pairs of pinnules, axes other than stipes with narrow lateral ridges (especially pronounced on axes below ultimate branching), and bilateral spores.

4 9 . H E C I ST O P T E R I S Hecistopteris J. Sm., London J. Bot. 1: 193. 1842. Type: Hecistopteris pumila (Spreng.) J. Sm. [⬅ Gymnogramma pumila Spreng.]. Epiphytic, often on rotting logs, or epipetric; rhizomes compact, short-creeping to suberect, protostelic, appearing longcreeping due to proliferous thread-like roots, apical scales clathrate; fronds distichous; stipes short to nearly absent; blades very small (1–3 ⫻ 0.2–1 cm), minutely to decidedly furcate or bifurcate at the tip, flabellate and often shallowly to deeply lacerate, lacking a distinct midrib, glabrous, epidermis with linear idioblasts adaxially; veins free, dichotomously forking; sori along the veins (gymnogrammoid) in the distal part of the blade, indusia absent; paraphyses simple or branched with a funnelform apical cell; spores pale yellowish-hyaline, tetrahedral, surfaces relatively smooth with scattered papillae; gametophytes with single gemmae; x⫽? For over 150 years of its existence, Hecistopteris was considered a monotypic genus, but a second species was recently described from Ecuador (H. pinnatifida R. C. Moran & B. Øllg.), and a third species from Guyana has been added (Kelloff & McKee, 1998). These additional taxa are all based on newly collected specimens, thus reminding us how much more collecting must be done before we have an adequate understanding of the tropical ferns. Hecistopteris is regarded by Crane (1997) as sister to Radiovittaria, and differs from that, as well as all other vittarioid ferns, by its very small, forking fronds, proliferous roots, and free veins. References

Distribution. Upright or trailing, forming dense patches on open, dry to moist slopes, montane rain forests, cloud forests, pine-oak forests, pine-oak-Liquidambar forests; 100–2150 m.

Crane, E. H. 1997 [1998]. A revised circumscription of the genera of the fern family Vittariaceae. Syst. Bot. 22: 509–517; Kelloff, C. L. & G. S. McKee. 1998. A new species of Hecistopteris from Guyana, South America. Amer. Fern J. 88: 155–157.

HEMIDICTYUM

1. HECISTOPTERIS PUMILA Hecistopteris pumila (Spreng.) J. Sm., London J. Bot. 1: 193. 1842. Figs. 147A–C. Gymnogramma pumila Spreng., Syst. Veg. Tent. Suppl. 31. 1828. Type. Surinam. Weigelt s.n. (LZ, destroyed; isotypes B!, P!; probable isotype UC!).

Rhizome scales linear, 1–1.5 ⫻ 0.1 mm; fronds clumped, 1–2 cm long, monomorphic or somewhat dimorphic, the fertile usually larger; stipes 1⁄3–2⁄3 the frond length; blades 1–2-forked at the tip; sori along the distal portions of the veins; paraphyses uniseriate, 0.3 mm long; spores whitish green.

323

ally elongate, pentagonal or hexagonal, without included veinlets, network closed at margin by a continuous intramarginal vein; sori linear, along main vein branches between costae and first anastomoses, paraphyses absent; indusia lateral, linear, very narrow, membranous; spores bilateral, with folded sometimes sparingly perforate perispore; x⫽31. Hemidictyum is a monotypic genus, the single species widespread in tropical America. Most specialists have thought it to be related to Diplazium (athyrioid ferns), but the chromosome number of x⫽31 is quite distinct from the base numbers of 40 and 41 prevalent in the athyrioid ferns. Recent unpublished molecular data by Cranfill indicate that Hemidictyum forms a clade with the diplazioid segregate genera Diplaziopsis (eastern Asia) and Homalosorus (Diplazium) pycnocarpon (Spreng.) M. Broun, in eastern United States, plus a similar Asian species D. flavoviride Alston), well apart from the core Diplazium; thus, the generic distinctness of Hemidictyum from Diplazium is strongly supported. Hemidictyum differs from Diplazium and Athyrium by the combination of large, once-pinnate fronds with subcordate-based pinnae, thin texture, netted veins, and long, indusiate sori.

1. HEMIDICTYUM MARGINATUM

Distribution. Epiphytic or on fallen logs in wet undisturbed forests of Atlantic slope; 400–600 m. Mexico; Guat, Bel, Hond, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol. Specimens Examined. Oax (Hallberg 1293, NY, US; Mickel 6440, NY, UC).

Because of its small size this species is difficult to find, and may be more widespread than current records indicate. It is colonial in nature, by virtue of proliferous thread-like roots that give rise to new plants. It is not easily confused with any other fern species.

50 . H E M I D I C T YUM Hemidictyum C. Presl, Suppl. Tent. Pterid. 110, t. 3, f. 24. 1836. Type: Hemidictyum marginatum (L.) C. Presl [⬅ Asplenium marginatum L.]. Terrestrial; rhizomes stout, decumbent to suberect; rhizome scales brownish, subclathrate, with golden brown lumina; fronds monomorphic, 1–3(–5) m long, clumped; stipes stout, green to brown, glabrous, with 2 crescent-shaped vascular bundles in cross section; blades once-pinnate (odd-pinnate), blade apices conform, the pinnae oblong, glabrous; veins unforked or 1forked near their bases, at nearly right angles to the costae, free near the costae, initially parallel and ca. 1–2 mm apart, forming a reticulate network in marginal half of the pinnae, areoles usu-

Hemidictyum marginatum (L.) C. Presl, Suppl. Tent. Pterid. 111, t. 3, f. 24. 1836. Figs. 148A, B. Asplenium marginatum L., Sp. Pl. 2: 1082. 1753. Asplenium limbatum Willd., Sp. Pl., ed. 4, 5(1): 310. 1810, nom. illeg. (based on Asplenium marginatum L.). Diplazium marginatum (L.) Diels in Engl. & Prantl, Nat. Pflanzenfam. 1(4): 229. 1899, hom. illeg., non Blume, 1828. Diplazium limbatum (Willd.) Proctor, Rhodora 68: 467. 1966. Lectotype (designated by Proctor, 1985: 390). Linnaeus cited Petiver, Pter. Amer., t. 12, f. 2, 1712, which is a redrawing of Plumier, Traite´ Foug. Ame´r., pl. 106, 1705, illustrating a plant probably from Martinique (Morne de la Calebasse), according to Proctor (1977). LINN 1250.20 is also this species.

Caudices to ca. 30 cm tall, 15 cm diam.; rhizome scales to 4 cm ⫻ 1 mm; stipes to 100 ⫻ 2 cm long, ca. 1⁄2 the frond length, bases somewhat scaly; blades 100–200 ⫻ 50–100 cm; pinnae opposite to subopposite, entire, sessile, often subcordate and overlapping the rachises, 25–55 ⫻ 6–12 cm, abruptly acuminate, bases cordate-clasping, laminae bright-green, succulent, papery and fragile on drying; indusia 15–30(–40) ⫻ 0.5–0.8 mm, entire; 2n⫽62 (Trin).

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HEMIONANTHES

1. HEMIONANTHES GRYPHUS Hemionanthes gryphus (Mickel) Mickel in McVaugh, Fl. Nov.-Gal. 17: 288. 1992. Figs. 149A–C. Cheilanthes gryphus Mickel, Amer. Fern J. 77: 112. 1987. Type. Mexico. Colima: Low mountain summits 7 mi N of Santiago, road to Durazno, Jal., via the bridge over Rı´o Cihuatla´n, McVaugh 15908 (MICH!).

Distribution. Terrestrial in lower montane and montane rain forests, especially along streams; 150–1000 m. Mexico; Guat, Hond, CR, Pan; Gr & L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, s Braz, Bol.

Rhizomes 3–7 mm diam.; rhizome scales 2–3 mm long, linear, bicolorous, with a central, lustrous, dark-castaneous to atropurpureous streak and narrow, tan margins; fronds to 46 cm long, approximate; stipes stramineous, grayish brown at bases, with very slender, mostly bicolorous scales at bases, otherwise glabrous; pinnae 5–7 pairs, each lanceolate to deltate, acuminate, with 2–4 pairs of broad obtuse lobes, the basiscopic lobes ca. twice as long as the acroscopic ones; veins mostly free but with occasional anastomoses (2–5 per pinna); sporangia along the distal 2–4 mm of the veins; 2n⫽60 (Nay).

Selected Specimens Examined. Chis (Collins & Doyle 233, US; Purpus 6764, MO, NY, UC, US; Rovirosa 857, NY; Ventura & Lo´pez 1156, ENCB, MEXU, MO, XAL). Oax (Hallberg 1613, NY, US; Martı´nezCaldero´n 392 MEXU, UC, US; Mickel 6381, ENCB, NY, UC, 7194, NY; Torres C. 11653, MEXU). Unverified, Doubtful, or Mistaken Reports. Ver (Acosta 3103-2d, CIB, XAL, cited by Palacios-Rios, 1992, but not verified).

The only large 1-pinnate, entire-margined species of Diplazium with which Hemidictyum might be confused in Mexico are D. grandifolium and D. obscurum, and these are easily distinguished by their free venation.

51 . HE M I O N A N T H E S Hemionanthes Mickel in McVaugh, Fl. Novo-Gal. 17: 286. 1992. Type: Hemionanthes gryphus (Mickel) Mickel [⬅ Cheilanthes gryphus Mickel]. Rhizomes compact, horizontal; rhizome scales linear, bicolorous; stipes stramineous, mostly glabrous, 3-grooved, about half the length of the frond; blades lanceolate to deltate, pinnatepinnatifid, to bipinnate at bases; pinna lobes obtuse, basiscopically somewhat exaggerated; adaxial surfaces glabrous, abaxial surfaces with sparse straight jointed hairs, rachises with sparse, narrow, bicolorous scales; veins mostly free, with occasional anastomoses; sporangia on the abaxial surfaces of the blades, extending along the distal 1⁄4–1⁄2 the length of the veins; indusia absent, the blade margins not reflexed or modified; spores tetrahedral, crested; x⫽30. The monospecific genus is intermediate between Cheilanthes and Hemionitis, combining characters of species of those genera, with intermediate dissection and soral disposition. Reference Mickel, J. T. 1987. A new fern from western Mexico and its bearing on the taxonomy of the cheilanthoid ferns. Amer. Fern. J. 77: 109–114.

Distribution. Terrestrial in deciduous woodlands in moist shaded gulleys; 200 m. Mexico. Specimens Examined. Col (only the type). Me´x (Vela´squez Montes 1149, MEXU). Mich (Eggler 171, US). Nay (Ranker 799, KANU; Yatskievych 85-73, CHAPA).

This species has the appearance of a hybrid between Hemionitis subcordata and Cheilanthes skinneri (Mickel, 1987). It is intermediate in blade dissection, length of pinna stalks, number of pinna pairs, blade shape, venation, pinna margin, and, most strikingly, in sorus pattern. No morphological features contradict this hypothesis. Furthermore, the two presumed parents were collected at the type locality. The spores of Hemionanthes gryphus are well formed and essentially the same size as those of the presumed parents. Thomas Ranker (pers. comm.) collected the plant in Nayarit and found it to be a diploid; it did not have the additive chemical composition that would be typical of fern hybrids. When Cheilanthes gryphus was first described (Mickel, 1987), Hemionitis subcordata was also transferred to that genus. It is possible that gryphus, skinneri, and subcordata (whatever their genera) are closely allied and form a distinct group apart from either Cheilanthes or Hemionitis, but the sorus configurations differ so dramatically that we prefer to retain skinneri, with its marginal sori and highly

HEMIONITIS

modified false indusia, in Cheilanthes, and subcordata, with its sori running nearly the entire length of the veins and its unmodified pinna margins, in Hemionitis. The latter is not a perfect fit in Hemionitis, but it would be an even more bizarre element in Cheilanthes.

52 . H E M I O N I T I S Hemionitis L., Sp. Pl. 2: 1077. 1753. Type: Hemionitis palmata L. Gymnopteris Bernh., J. Bot. (Schrader) 1799(1): 297. 1799. Type: Gymnopteris rufa (L.) Underw. [⬅ Acrostichum rufum L.] ⬅ Hemionitis rufa (L.) Sw.

Terrestrial; rhizomes short, compact, erect or ascending, scaly; fronds small, monomorphic or slightly dimorphic; stipes hairy to glabrous, castaneous to stramineous, lustrous; blades palmate to pinnate, hairy to glabrous; veins free to netted without included veinlets; laminae herbaceous, glabrous or with acicular or septate hairs on both surfaces; sori along veins for most of their length;

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indusia absent; spores tetrahedral-globose, tuberculate to cristate; x⫽30. Hemionitis is historically limited to the palmate species with netted veins (five species), but that was shown to be an artificial separation from Gymnopteris (Mickel, 1974; Giannasi & Mickel, 1979), and the two genera are now generally united. There are about 10 species, mostly of the Neotropics, with five in Mexico. However, there is still a problem with generic circumscription, since H. subcordata, with gymnogrammoid sori, hybridizes in Colima, Michoca´n, and Nayarit with Cheilanthes skinneri, which has marginal sori. It seems likely that the genera will have to be redefined without emphasis on the soral configuration. Hemionitis is distinguished from Bommeria by its sori extending nearly the entire length of the veins. References Giannasi, D. E. & J. T. Mickel. 1979. Systematic implications of flavonoid pigments in the fern genus Hemionitis (Adiantaceae). Brittonia 31: 405–412; Lellinger, D. B. 1969. The taxonomic position of Coniogramme americana. Amer. Fern J. 59: 61–65; Mickel, J. T. 1974. A redefinition of Hemionitis. Amer. Fern J. 64: 3–12; Mickel, J. T. 1987. A new fern from western Mexico and its bearing on the taxonomy of the cheilanthoid ferns. Amer. Fern J. 77: 109–114.

Key to the Mexican Species of Hemionitis 1. Fronds fully 1-pinnate, 12–60 cm long. 2. Pinnae 7–12 pairs, oblong; stipes and rachises dark reddish brown to atropurpureous; blade surfaces pubescent; veins free. ......... 4. H. rufa. 2. Pinnae 3–7 pairs, ovate-lanceolate; stipes and rachises stramineous; blade surfaces virtually glabrous, occasionally with hairs or linear scales at pinna bases; veins with occasional anastomoses near costae. ................................................................................... 5. H. subcordata. 1. Fronds palmately to pinnately lobed, 2.5–27 cm long. 3. Margins entire, without buds; hairs on pinnae stiff, not jointed, 0.5–1 mm long; basal and erect leaves fertile; spores tan. .......... 1. H. levyi. 3. Margin crenulate with buds in one or more sinuses; hairs on pinnae jointed, 1–1.5 mm long; basal leaves sterile; spores light yellow-orange. 4. Rhizome scales bicolorous with central dark brown streak; fronds deeply pinnately 5–7-lobed; blades with buds in many small sinuses. ...................................................................................................................................................................................... 3. H. pinnatifida. 4. Rhizome scales concolorous, light brown; blades 3–5-palmately lobed; blades usually with one large bud in a major sinus. ............................................................................................................................................................................................... 2. H. palmata.

1. HEMIONITIS LEVYI Hemionitis levyi E. Fourn., Bull. Soc. Bot. France 17: 237. 1870. Type. Nicaragua. ˆIle d’Omotepec [Isla de Ometepe], Le´vy [1197] (P!, frag. US!; isotype B!). Fig. 150B. Hemionitis otonis Maxon, Contr. U.S. Natl. Herb. 17: 171. 1913. Type. Costa Rica. Jime´nez [“Jiminez” in original publ.] 333 (US!).

Rhizomes erect, 1–2.5 mm diam.; rhizome scales 3–4 ⫻ 0.5 mm, bicolorous with dark castaneous streak, becoming tortuous and concolorous (tan) at bases of stipes; fronds 2.5–16 cm long, clumped; stipes 1⁄3–3⁄4 the frond length, 0.4–1.3 mm diam., castaneous, broadly grooved, with dense multicellular stiff hairs 1 mm each with bulbous base and long, lax, capitate hairs 2 mm; blades simple, undivided, 1.7–5 cm wide, orbicular to shallowly 3–5-palmately lobed, resembling Hedera, bases cordate, lobes broadly acute to obtuse, margins entire, not proliferous or recurved, both basal and erect leaves fertile; adaxial and abaxial surfaces with dense, stiff, white hairs 0.5–1 mm; veins netted, without included veinlets; sori along all veins; spores tan.

Distribution. Terrestrial, among rocks in moist lowland forests of Pacific slope; 100–750 m. Mexico; Salv, Nic, CR. Specimens Examined. Chis (Pe´rez Farrera 670, UAMIZ). Gro (Lorea 2805, IEB, 3744, IEB). Oax (Mickel 5976, 6075, 6091, NY).

Hemionitis levyi is distinct by its small frond size, entire blade margins, short laminar hairs, and lack of buds.

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HEMIONITIS

2. HEMIONITIS PALMATA Hemionitis palmata L., Sp. Pl. 2: 1077. 1753. Type. Several citations, incl. Plumier, Descr. Pl. Ame´r., pl. 33. 1693. Fig. 150C. Rhizomes erect, 2–3 mm diam.; rhizome scales linear, concolorous, light brown, 4–5 ⫻ 0.8 mm; fronds 7–27 cm long, clumped, dimorphic, the sterile much shorter and spreading with rounder lobes, the fertile fronds about twice as tall as the sterile ones; stipes 1⁄2–5⁄6 the frond length, to 1.8 mm diam., castaneous, grooved to flattened adaxially, with narrow, light brown scales, those at base grading into multicellular, acicular or capitate hairs and short glandular hairs 0.1 mm; blades palmate with 3–5 acute lobes, 4.5–10 cm wide; margins crenulate, not recurved, with buds in the larger sinuses, but usually only one developing into a new plant (under exceptionally high humidity buds sometimes also developing in many of the minor notches); veins netted without included veinlets, the five major veins dark; adaxial and abaxial surfaces pilose with jointed, somewhat curved, multicellular, acicular hairs 1–1.5 mm long; sori along all veins; spores light yellow-orange; 2n⫽60 (Oax, CR, Jam).

Hybrids are distinguished from H. palmata by the malformed spores, spottily bicolorous scales, (5–)7 pinnately arranged dark major veins, the dark portion of major veins extending into minor veins, and small buds in smaller sinuses in addition to larger bud in a major sinus. Hemionitis palmata also crosses with H. rufa in Jamaica. The hybrid should be sought where these two occur together.

3. HEMIONITIS PINNATIFIDA Hemionitis pinnatifida Baker in Hooker & Baker, Syn. Fil. 399. 1868. Syntypes. Guatemala. Bernoulli 32 (K!); Costa Rica. Wendland 438 (K!). Fig. 150A. Rhizomes erect, 2–3 mm diam.; rhizome scales linear, bicolorous, central portion dark brown with margins light brown, becoming concolorous at base of stipes; fronds 7–24 cm, clumped; stipes 1⁄2–2⁄3 the frond length, 1–2 mm diam., castaneous, grooved to flattened adaxially, with narrow scales at bases grading into multicellular, acicular to capitate hairs 2 mm long; blades deeply pinnately 5–7-lobed, 5–8 cm wide, the basal pair usually with a single basiscopic lobe; sterile lobes rounded; fertile lobes acute to acuminate; margins crenulate, not recurved, proliferous from many small buds in small sinuses; veins netted without included veinlets, dark color of 5–7 major veins extending into branches of 5–7 minor veins, minor veins pinnately arranged; adaxial and abaxial surfaces pilose with jointed, multicellular, acicular hairs 1–1.5 mm long; sori along all veins; spores light yellow-orange; 2n⫽120 (Oax).

Distribution. Terrestrial, in soil on moist rocky slopes in mesic forests; 60–1000 m. Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Cuba; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol. Selected Specimens Examined. Chis (Breedlove 27483, DS). Col (Eyerdam & Beetle 8713, UC, US). Gro (Hinton 11052, LL, MEXU, US). Oax (Mickel 5781, NY). Pue (Patro´n & Basurto 68, MEXU). Qro (Dı´az B. & Carranza 6685, IEB, MEXU). Sin (Breedlove 35580, CAS). SLP (Pringle 3972, CAS, LL, MEXU, MICH, US). Tab (Cowan 3165, CAS, MEXU). Tam (Palmer 296, UC). Ver (Herna´ndez M. 2010, MEXU). Yuc (Gaumer 1072, GH).

This species crosses with H. pinnatifida in Oaxaca (Mickel, 1974) forming a triploid hybrid with malformed spores. Apparently, the genome of H. pinnatifida includes a genome from H. palmata, since the hybrid has 30II and 30I at meiosis (Smith & Mickel, 1977). Specimens of H. palmata ⫻ pinnatifida from Oaxaca are in woods and among exposed rocks, 750–1050 m, in the districts of Juquila (Mickel 6133b, NY) and Villa Alta (Mickel 1113, NY).

Distribution. Terrestrial in soil on moist, rocky slopes in mesic forest, mostly on the Pacific slope; 50–1000 m. Mexico; Guat, Hond, Salv, Nic, CR. Selected Specimens Examined. Chis (Breedlove 38432, MEXU, 71513, CAS; Matuda 2128, GH, MEXU, US). Gro (Gonza´lez 344, FCME; Lorea 401, IEB). Jal (Rzedowski & McVaugh 1361, ENCB). Oax (Mickel 5988, 6133a, 6238, 5975, NY).

Hemionitis pinnatifida is distinct from H. palmata by having bicolorous rhizome scales and pinnately arranged lobes and major veins. Hybrids with H. palmata are known from Oaxaca (see also discussion under H. palmata).

HISTIOPTERIS

4. HEMIONITIS RUFA Hemionitis rufa (L.) Sw., J. Bot. (Schrader) 1800(2): 16. 1801 [1802]. Figs. 151A, B. Acrostichum rufum L., Syst. Nat., ed. 10, 2: 320. 1759. Gymnopteris rufa (L.) Bernh. ex Underw., Bull. Torrey Bot. Club 29: 627. 1902. Type (according to R. Tryon, Contr. Gray Herb. 194: 82. 1964). Sloane, Nat. Hist. Jam. 1: pl. 45, f. 1, 1707, illustrating a plant from Jamaica; LINN 1245.6 is also this species.

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Rhizomes ascending, 2–4 mm diam.; rhizome scales linear, bicolorous; fronds 12–45 cm, clumped; stipes ca. 1⁄2 the frond length, 1–2 mm diam., stramineous, deeply grooved, nearly glabrous; blades pinnate, 5–21 cm wide, each with conform terminal pinna; pinnae 3–7 pairs sessile, ovate-lanceolate, bases unequally rounded, occasionally with hairs or linear scales at pinna bases; blade margins entire; veins predominantly free, but with occasional anastomoses; costae dark at bases; surfaces glabrous; sori along all the veins nearly to the costae; 2n⫽60 (Oax).

Rhizomes suberect; rhizome scales bicolorous; fronds ca. 20–60 cm long, with stipes (1⁄3–) 1⁄2–2⁄3 the blade length; stipes and rachises dark reddish brown to atropurpurpeous, pubescent with short, acicular trichomes and fewer, longer, multicellular ones; blades 1-pinnate, each with a conform terminal pinna; pinnae oblong, mostly 7–12 lateral pairs, 5–8 cm long, broadly cuneate to subcordate at bases, short-stalked to 5 mm, apices acute, margins entire; veins free; laminar tissue on both sides more or less pubescent with acicular, multicellular, Ⳳ appressed trichomes; 2n⫽60 (Jam).

Distribution. Terrestrial on shaded rocky banks in woods or by roads, especially on the Pacific slope; 200–1400 m. Mexico; Guat, Nic, CR. Selected Specimens Examined. Chis (Breedlove 26995, DS). Col (McVaugh 15925, MICH). Dgo/Sin (Palmer 1416, NY, UC, US). Gro (Kruse 604, ENCB). Jal (McVaugh 19811, MICH). Me´x (Hinton 8062, ENCB, GH, US). Mich (Hinton 12213, US). Nay (McVaugh 1900, MICH). Oax (Mickel 6980, NY). Sin (Breedlove 35669, CAS). Tres Mar´ıas (Ferris 5712, DS).

Distribution. Rain forests; 300–1000 m. Mexico; Guat, Hond, Nic, CR, Pan; Jam; Col, Ven, Sur, Ec, Peru, Braz, Bol. Specimens Examined. Chis (Breedlove 33138, DS, MICH, 42848, DS; Castillo C. et al. 3842, MICH, XAL; Martı´nez S. 10493, ENCB, XAL; Matuda 2523, GH, MEXU, MICH, US, 3415, GH). Tab (Matuda 3415, US).

Hemionitis rufa hybridizes with H. palmata in Jamaica (H. (Trevis.) C. Chr.), but this hybrid has not been found in Mexico.

⫻smithii

5. HEMIONITIS SUBCORDATA Hemionitis subcordata (D. C. Eaton ex Davenp.) Mickel, Brittonia 31: 411. 1979. Figs. 151C, D. Gymnogramme subcordata D. C. Eaton ex Davenp. in Rose, Contr. U.S. Natl. Herb. 5: 138, t. 16. 1897. Gymnopteris subcordata (D. C. Eaton ex Davenp.) Underw., Bull. Torrey Bot. Club 29: 628. 1902. Coniogramme subcordata (D. C. Eaton ex Davenp.) Maxon, Contr. U.S. Natl. Herb. 17: 174. 1913, hom. illeg., non Copel., 1910. Coniogramme americana Maxon, Contr. U.S. Natl. Herb. 17: 607. 1916. Lectotype (chosen by Mickel & Beitel, 1988). Mexico. Sinaloa or Durango: Ymala and Lodiego, Palmer 1572 (US!).

Hemionitis subcordata is unique in the genus in having 1pinnate blades and laminar surfaces glabrous. In Colima this has been found to cross with Cheilanthes skinneri (see Hemionanthes). The sori are so different in these two species that it raises the question of usefulness of sorus type as a measure of relationship in this group. See Mickel (1987).

5 3. H I ST I O P T E R I S Histiopteris (J. Agardh) J. Sm., Hist. Fil. 294. 1875. Pteris L. sect. Histiopteris J. Agardh, Recens. Spec. Pter. 76. 1839. Type: Histiopteris vespertilionis (Labillardie`re) J. Sm. [⬅ Pteris vespertilionis Labillardie`re] ⫽ Histiopteris incisa (Thunb.) J. Sm. Terrestrial; rhizomes long-creeping, 2–5 mm diam., with castaneous hairs and scales; fronds large, distant, erect to subscandent; stipes stramineous to brown, lustrous, glabrous; blades bito tripinnate, often with large stipule-like pinnule at pinna base; pinnae and pinnules opposite, light pea-green, herbaceous; lamina glabrous or with crisped, multicellular hairs, often glaucous; veins netted, without included veinlets; sori marginal, continuous, blade margins reflexed and differentiated as indusia,

328

HOLODICTYUM

lacking inner indusia; spores bilateral, yellow or light brown; x⫽48. Histiopteris has one rather variable species over its pantropical range, and four other species in Southeast Asia and the South Pacific. It is closely related to Pteridium and Lonchitis, but lacks an inner indusium, has rhizome scales, and is not aggressive. It is limited mostly to higher elevations (cloud forests). References Wolf, P. G. 1995 [1996]. Phylogenetic analyses of rbcL and nuclear ribosomal RNA gene sequences in Dennstaedtiaceae. Amer. Fern J. 85: 306–327; Wolf, P. G., P. S. Soltis & D. E. Soltis. 1994. Phylogenetic relationships of dennstaedtioid ferns: evidence from rbcL sequences. Molec. Phylogen. Evol. 3: 383–392.

1. HISTIOPTERIS INCISA Histiopteris incisa (Thunb.) J. Sm., Hist. Fil. 295. 1875. Figs. 152A–C. Pteris incisa Thunb., Prodr. Pl. Cap. 171. 1800. Litobrochia incisa (Thunb.) C. Presl, Suppl. Tent. Pterid. 149. 1836. Type. South Africa. Cape of Good Hope, 1774, Thunberg s.n. (UPS-THUNB; isotype S).

Rhizomes long-creeping, 5–10 mm diam.; rhizome scales 2–3 0.8–1 mm, brown, linear-lanceolate, clathrate, along with golden brown hairs 2 mm long; fronds 1–4 m long, distant; stipes 1⁄4–1⁄2 the frond length, stramineous to castaneous; blades mostly bipinnate-pinnatifid, rarely more divided, 1–2 m wide, glabrous, glaucous; segments rounded; veins partially areolate; sporangia mixed with paraphyses; spores golden; 2n⫽96 (Jap), 192 (Jam), ca. 200 (Tristan da Cunha). ⫻

54. HOLODICTYUM Holodictyum Maxon, Contr. U.S. Natl. Herb. 10: 481. 1908. Type: Holodictyum ghiesbreghtii (E. Fourn.) Maxon [⬅ Asplenium ghiesbreghtii E. Fourn.]. Epipetric; rhizomes short, horizontal, clothed with clathrate scales; fronds 1–2 mm apart; stipes short, glabrous, dark; blades undivided, entire, oblanceolate, long-tapering to cuneate at base, glabrous, herbaceous; veins netted, without included veinlets; sori elongate, on nearly unmodified outer arcs of costal areoles, indusiate; spores bilateral, surface perforate or fenestrate; x⫽? Holodictyum is a rare, monotypic genus of Mexico. It is closely allied to Asplenium, being distinct largely on the basis of its netted veins, sori borne only on the outer arc of costal areoles, and undivided blade. Its relationship to or origin from any other extant species or species group in Asplenium is obscure; therefore, we maintain its generic status here, but without conviction that it will be found to deserve generic status once molecular data are available. The netted veins might suggest an affinity to A. purdieanum Hook., from Honduras, Colombia, and Venezuela, an equally rare species. Reference Maxon, W. R. 1908. A new genus of asplenioid ferns. Contr. U.S. Natl. Herb. 10: 481–482.

1. HOLODICTYUM GHIESBREGHTII Holodictyum ghiesbreghtii (E. Fourn.) Maxon, Contr. U.S. Natl. Herb. 10: 482. 1908. Figs. 153A–D. Asplenium ghiesbreghtii E. Fourn., Mexic. Pl. 1: 111, t. 5. 1872. Type. Mexico. Oaxaca: Barranca de Tlacolula, Ghiesbreght 16 (P–photos TEX!, US!, frag. NY!; isotype US!, frag. NY!). Asplenium finckii Baker, Ann. Bot. (Oxford) 8: 126. 1894. Holodictyum finckii (Baker) Maxon, Contr. U.S. Natl. Herb. 10: 482. 1908. Type. Mexico. Veracruz: Co´rdoba, Finck s.n. (K!; isotype B, frag. NY! “Finck 139”).

Rhizome scales 3–5 ⫻ 0.5–0.8 mm, black; stipes 0.5–1.5 cm long; blades 30–50 ⫻ 2–3 cm; sori at a narrow angle to midribs; indusia 0.8–1.5 cm ⫻ 0.8–1 mm, entire.

Distribution. Open banks and light woods in cloud forests; (1150–)2600–3050 m. Mexico; Guat, Hond, Salv, Nic, CR, Pan; Gr & L Ant; Col, Ven, Ec, Peru, s Braz, Bol, J Fdez; s Africa, e & se Asia, NZ, Tasmania. Selected Specimens Examined. Chis (Breedlove 22015, MEXU, NY, 22048, MEXU; Mickel 1237, NY). Mich (Carranza G. 458, IEB; Rzedowski 42669, MEXU). Oax (Lorence 3262, MEXU; Mickel 1065, MEXU, NY). Ver (Narave F. 975, NY; Purpus 6056, NY; Ventura 4475, MEXU, NY).

Histiopteris incisa is conspicuous by its large, light green, glabrous, glaucous fronds with partially netted veins.

HUPERZIA

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a single vein, not in distinct ranks, appressed, ascending or spreading, lanceolate to linear-lanceolate to oblanceolate, monomorphic or dimorphic, the vegetative leaves sometimes varying in size with seasonal growth patterns; margins entire or dentate; gemmiferous branchlets absent or rare (H. beiteliana), small, formed among the leaves; sporangia reniform, borne in axils of normal vegetative leaves or specialized leaves of a terminal strobilus; spores isomorphic, pitted; x⫽67, 68.

Distribution. On wet rocks and in rock crevices in shaded limestone faces, near seeps, in wet lower montane forests, tropical semi-evergreen forests; 350–1500 m. Mexico. Specimens Examined. Hgo (Cota & Gimate 7775, NY, XAL; Herna´ndez Magan˜a 5521, CAS, MEXU; Ramı´rez & Riba 636, MEXU). Oax (only the type). Pue (Go´mez-Pompa s.n., MEXU). Qro (Dı´az B. & Zamudio 4925, IEB, MEXU; Rubio 370, 648, IEB, 2069, IEB, MEXU; Servı´n 437, 874, IEB). SLP (Sharp 46289, MEXU, US). Tam (Garcı´a A. 726, ITCV; Palmer 336, NY, UC, US). Ver (Finck 139, K, NY).

Holodictyum finckii is included under H. ghiesbreghtii because it differs in only minor characters; there are few collections of either taxon.

5 5 . H UPE R Z I A Huperzia Bernh., J. Bot. (Schrader) 1800(2): 126. 1801. Urostachys Herter, Beih. Bot. Centralbl. 39(2): 249. 1922, nom. superfl. Type: Huperzia selago (L.) Schrank & Mart. [⬅ Lycopodium selago L.]. Epiphytic or terrestrial; stems dichotomously branched, erect or pendent, lacking horizontal stems; leaves microphyllous with

Huperzia is a cosmopolitan genus of about 400 species, mostly of the wet tropics in middle to higher elevations and with several temperate species. There are 15 species in Mexico. Huperzia is the largest lycopod genus, related to Lycopodium s.str. and Lycopodiella. We thank Benjamin Øllgaard for comments and suggestions. References Herter, W. 1949. Index Lycopodiorum. Estudios Bot. Regio´n Uruguaya, Montevideo 20: 1–120; Nessel, H. 1939. Die Ba¨rlappgewa¨chse (Lycopodiaceae). Gustav Fischer, Jena. 404 pp.; Øllgaard, B. 1975. Studies in Lycopodiaceae. I. Observations on the structure of the sporangium wall. Amer. Fern J. 65: 19–27; Øllgaard, B. 1979. Studies in Lycopodiaceae, II. The branching patterns and infrageneric groups of Lycopodium s.l. Amer. Fern J. 69: 49–61; Øllgaard, B. 1983. Lycopodiaceae. In: Stolze, R., Ferns and fern allies of Guatemala, Part III. Fieldiana, Bot., n.s. 12: 20–44; Øllgaard, B. 1989. Index of the Lycopodiaceae. Biol. Skr. 34: 1–135; Underwood, L. M. & F. E. Lloyd. 1906. The species of Lycopodium of the American tropics. Bull. Torrey Bot. Club 33: 101–124; Wikstro¨m, N. 2001. Diversification and relationships of extant homosporous lycopods. Amer. Fern J. 91: 150–165; Wikstro¨m, N. & P. Kenrick. 1997. Phylogeny of Lycopodiaceae (Lycopsida) and the relationships of Phylloglossum drummondii Kunze based on rbcL sequences. Int. J. Pl. Sci. 158: 862–871; Wikstro¨m, N. & P. Kenrick. 1999a. Phylogeny of epiphytic Huperzia (Lycopodiaceae): paleotropical and neotropical clades corroborated by plastid rbcL sequence. Nordic J. Bot. 20: 165–171; Wikstro¨m, N. & P. Kenrick. 1999b. Diversification and relationships of extant homosporous lycopods. XVI International Botanical Congress, Abstracts. p. 91; Wikstro¨m, N. & P. Kenrick. 2001. Evolution of Lycopodiaceae (Lycopsida): estimating divergence times from rbcL gene sequences using nonparametric rate smoothing. Mol. Phylogenet. Evol. 19: 177–186; Wikstro¨m, N., P. Kenrick & M. Chase. 1999. Epiphytism and terrestrialization in tropical Huperzia (Lycopodiaceae). Pl. Syst. Evol. 218: 221–243.

Key to the Mexican Species of Huperzia 1. Plants terrestrial, stems generally erect. 2. Leaves deeply serrate, narrowed to petiole-like bases. ............................................................................................................................. 13. H. serrata. 2. Leaves entire or only minutely cilio-denticulate, broadest at bases or slightly above. 3. Leaves tightly appressed, coriaceous, entire; sporangia hidden by the strongly appressed, imbricate leaves. .............................. 3. H. crassa. 3. Leaves spreading to deflexed, firm but not coriaceous, entire or denticulate; sporangia evident among the spreading leaves. 4. Plants bearing gemmae among the sporophylls in the recent year’s growth. ....................................................................... 1. H. beiteliana. 4. Plants lacking gemmae. 5. Leaves mostly 5–7 mm long, minutely cilio-denticulate; stems stramineous to light brown; 300–2400 m. ............. 12. H. reflexa. 5. Leaves mostly 11–20 mm long, entire; stems green to dark red; 2100–3000 m. ....................................................... 6. H. hippuridea. 1. Plants epiphytic; stems generally pendent, rarely stiffly erect. 6. Basal leaves ovate to oblong-lanceolate, 2.5–3.5 mm wide. ................................................................................................................. 8. H. myrsinites. 6. Basal leaves linear to linear-lanceolate, 0.4–0.6 mm wide. 7. Leafy branches very slender, 1–3(–6 at base) mm wide (including leaves). .................................................................................. 11. H. pringlei. 7. Leafy branches thicker, (8–)12–35 mm broad (including leaves). 8. Leaves ascending, widely spreading only at base of plants. 9. Leaves tightly appressed, coriaceous; sporangia completely hidden among the tightly imbricate leaves; sporophylls not distinctly smaller than other leaves; leaves of distal divisions in 10–12 ranks. ............................................... 4. H. cuernavacensis. 9. Leaves slightly spreading, subcoriaceous; sporangia evident among the more loosely arranged sporophylls, which are somewhat smaller than other leaves; leaves of distal divisions in 6 ranks. .............................................................. 14. H. taxifolia.

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HUPERZIA 8. Leaves widely spreading (sporophylls more ascending). 10. Stems pendent, flaccid, 0.5–1.2 mm diam. (without leaves); leaves scattered, distant, slightly petiolulate, often upwardly falcate. 11. Leaves 1(–3) mm wide; stems stramineous to greenish or light brown. .................................................................. 7. H. linifolia. 11. Leaves 0.6–1 mm wide; stems reddish. ....................................................................................................................... 2. H. capillaris. 10. Stems erect or pendent, usually stiff, (1.5–)2–4 mm diam. (without leaves); leaves abundant, sessile or slightly petiolulate, not upwardly falcate. 12. Stems dark red; leaves mostly (2–)3–4 cm long. ..................................................................................................... 10. H. pithyoides. 12. Stems stramineous to light brown; leaves mostly 0.7–2 cm long. 13. Leaves somewhat distant, 1–2 mm wide, narrowed to petiolulate bases, mostly widely spreading. ............ 9. H. orizabae. 13. Leaves rather crowded, 0.2–1 mm wide, widely spreading to ascending. 14. Leaves 0.8–1 mm wide, linear-subulate. ..................................................................................................... 5. H. dichotoma. 14. Leaves 0.2–0.3 mm wide, filiform. ................................................................................................................ 15. H. wilsonii.

1. HUPERZIA BEITELIANA Huperzia beiteliana Mickel, Brittonia 44: 312. 1992. Type. Mexico. Oaxaca: Distrito Ixtla´n, 46 km N of Ixtla´n de Jua´rez on Rte 175, 1 km N of high point at Cerro Pelo´n, Mickel 9650 (NY!). Figs. 157D–H. Shoots erect, indeterminate, 12–45 cm tall, becoming longdecumbent, evergreen, with long-trailing senescent portions turning brown, juvenile and mature portions similar without strong annual constrictions; stems (without leaves) 2–3 mm diam.; juvenile buds erect; gemmiphores and gemmae produced mostly among the sporophylls in recent year’s growth (mostly in the distal 1⁄3 of the fertile zone), less frequently in the 1–8 mm breaks between fertile regions; gemmae 5–6 ⫻ 3–4 mm, lateral lobes broadly obtuse, 1.7 mm wide; leaves spreading to deflexed, dark green, lustrous, firm but not coriaceous, all more or less linear, 5–7 ⫻ 1.3–1.6 mm with entire margins, stomates on both abaxial and adaxial surfaces; spores 31–42 ␮m diam. (avg. 37).

northern North America and Eurasia in the general characters of leaves, gemmae, and spores, but is distinct in the much greater plant size (H. selago generally less than 15 cm tall), lack of annual constrictions, gemmiphores and gemmae located mostly among the sporophylls (not restricted to a pseudowhorl between the fertile zones), gemmae 5–6 mm long (vs. 4–5 mm), and spores 31–42 (avg. 37 ␮m diam.) vs. 29–37 (avg. 31 ␮m). Huperzia beiteliana occurs nearly 3000 km from the nearest populations of H. selago.

2. HUPERZIA CAPILLARIS Huperzia capillaris (Sodiro) Holub, Folia Geobot. Phytotax. 20: 71. 1985. Figs. 158F–H. Lycopodium capillare Sodiro, Recens. Crypt. Vasc. Quit. 90. 1883. Urostachys capillaris (Sodiro) Herter, Index Lyc. 54. 1949. Type. Ecuador. Hac. Guanaxilla, Sodiro s.n. (K). Lycopodium guatemalense Maxon, Contr. U.S. Natl. Herb. 17: 177, t. 9. 1913. Urostachys guatemalensis (Maxon) Herter, Repert. Spec. Nov. Regni Veg. 19: 165. 1923. Type. Guatemala. Pansamala´, Tu¨rckheim s.n. [Donn. Sm. 957] (US, frag. BM; isotype UC!).

Similar to H. linifolia in the scattered (not imbricate), spreading, linear-falcate leaves, but distinct in the leaves being narrower (mostly 8–12 ⫻ 0.6–1 mm vs. 15–25 ⫻ 2–3 mm, 1 mm only near stem apices), and with reddish stems.

Distribution. Terrestrial on wet rocky banks in pine-oak forests; 2575–2700 m. Mexico. Specimens Examined. SLP (Schaffner s.n., Aug 1879, NY). Oax (Taylor 2442, DUKE, ENCB).

The specimen from San Luis Potosı´ (Schaffner s.n., NY, Aug 1879, near Santa Marı´a del Rı´o) was labelled as “Lycopodium selago var. recurvatum mihi” [Schaffner], but that name was apparently never published. Huperzia beiteliana closely resembles H. selago (L.) Bernh. of

Distribution. Epiphyte in wet forests; 1170–1300 m. Mexico; Guat, CR, Pan; Col, Ven, Ec, Braz. Specimens Examined. Chis (Breedlove 29630, DS, 53567, CAS).

HUPERZIA DICHOTOMA

3. HUPERZIA CRASSA Huperzia crassa (Humb. & Bonpl. ex Willd.) Rothm., Repert. Spec. Nov. Regni Veg. 54: 60. 1944. Fig. 154B. Lycopodium crassum Humb. & Bonpl. ex Willd., Sp. Pl., ed. 4, 5(1): 50. 1810. Urostachys crassus (Humb. & Bonpl. ex Willd.) Herter ex Nessel, Ba¨rlappgewa¨chse 75. 1939. Type. Ecuador. Antisana, Humboldt & Bonpland 2263 (BWilld. 19417; isotypes BM, P-Humb.)

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leaves isomorphic, coriaceous, lustrous, spirally arranged in many ranks, crowded and strongly imbricate, ascending and tightly appressed to somewhat spreading only at base of plant, the basal ones more spreading, linear-lanceolate, 1–2 cm ⫻ 1.3–2 mm, usually zoned, signifying seasons of growth; sporophylls similar to other leaves, not distinctly smaller; sporangia hidden by the crowded sporophylls.

Plants 6–9(–15) cm tall, erect from short, anisodichotomous, creeping bases, erect stems simple to twice dichotomous, the divisions subparallel; stems ca. 2 mm diam. (without the leaves, 7–9 mm with the leaves), stramineous; leaves spirally arranged in 8–10 ranks, tightly appressed and imbricate, coriaceous, lustrous, linear-lanceolate, 0.7–1 cm ⫻ 1.5–2 mm above bases, 1 mm wide at bases, entire; sporophylls like vegetative leaves; sporangia hidden by the strongly appressed imbricate sporophylls.

Distribution. Pendent epiphyte in evergreen cloud forests, wooded slopes with oak, fir, and Podocarpus, and epipetric in volcanic flows; 1850–3000 m. Mexico; Guat, Salv. Selected Specimens Examined. Chis (Breedlove 22819, DS, MEXU, NY). Gro (Croat 45516, MO). Jal (Iltis et al. 7899, MEXU, NY). Me´x (Hinton 9966, LL, NY). Mich (Garay & Nava 3077, IEB). Mor (Pringle 7613, GH, MO, NY, US). Oax (Camp 2756, NY; Mickel 3841, NY). Sin (Gentry & Arguelles 18188, NY, US). Ver (Matuda 28781, MEXU).

Distribution. Terrestrial in paramos and open cloud forests; 2700–2900 m. Mexico; Guat, CR, Pan; Hisp; Col, Ven, Ec, Peru, Bol. Specimens Examined. Oax (Mickel 9651, NY; Sharp s.n., 26 Dec 1970, MEXU, NY).

This is the northernmost locality for this species. It was found with Jamesonia alstonii, also of paramos and reaching its northern limit here. Among the terrestrial club mosses, H. crassa is distinct by its tightly appressed, coriaceous leaves, and hidden sporangia.

4. HUPERZIA CUERNAVACENSIS Huperzia cuernavacensis (Underw. & F. E. Lloyd) Holub, Folia Geobot. Phytotax. 20: 72. 1985. Fig. 155C. Lycopodium cuernavacense Underw. & F. E. Lloyd, Bull. Torrey Bot. Club 33: 110. 1906. Urostachys cuernavacensis (Underw. & F. E. Lloyd) Herter ex Nessel, Ba¨rlappgewa¨chse 92. 1939 [“cuernavensis”]. Type. Mexico. Morelos: mountains above Cuernavaca, Pringle 7613 (NY!; isotypes GH!, K, MO!, P, US!, Z).

Plants mostly 25–70 cm long, stems arching to pendent, forked 2–4(–6) times; stems 3–7 mm diam. (without leaves);

Huperzia cuernavacensis is very closely allied to H. taxifolia and difficult to distinguish from it. It is generally strongly coriaceous with leaves tightly appressed and at higher elevations and more westerly distribution, but H. taxifolia appears to be quite variable in its leaf texture and degree of leaf appression. Øllgaard (1983) stated that a few collections tend to intergrade in leaf shape and texture with H. taxifolia, but the number of ranks of leaves in the distal portion (6 ranks in H. taxifolia vs. 10–12 in H. cuernavacensis) serves to distinguish them. The collection from the state of Me´xico is narrower and resembles H. hartwegiana (Spring) Trevis., a related species of Colombia, which has eight ranks of leaves in the distal portion and a slight cone structure, whereas H. cuernavacensis has the sporangia in the axils of undifferentiated leaves. Much more work needs to be done, especially as to the effects of elevation and of differences in seasonal rainfall.

5. HUPERZIA DICHOTOMA Huperzia dichotoma (Jacq.) Trevis., Atti Soc. Ital. Sci. Nat. 17: 248. 1874. Figs. 158J, K. Lycopodium dichotomum Jacq., Enum. Stirp. Vindob. 314. 1762. Urostachys dichotomus (Jacq.) Herter, Beih. Bot. Centralbl. 39: 249. 1922. Lectotype (Proctor 1977: 28). Jacquin, Hort. Bot. Vindob. 3: t. 45. 1776, illustrating a plant from Martinique.

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HUPERZIA

Plants 15–55 cm long, pendent from arching bases, 2–6 times dichotomous, the primary divisions often widely divaricate (45– 90⬚), the distal divisions usually divergent at smaller angles; stems 1.5–3.5 mm diam. (without leaves), stramineous to light brown; leaves spirally arranged in many ranks, spreading (especially the basal ones) to ascending, linear-subulate, proximal leaves 1.5–2 cm ⫻ 0.8–1 mm, entire, papyraceous, often subfalcate, midribs raised abaxially; sporophylls reduced to 8–15 ⫻ 0.5 mm, otherwise similar to vegetative leaves, not expanded at bases; sporangia visible; 2n⫽ca. 264 (PR).

Distribution. Terrestrial in cloud forests, pine-oak cloud forests; 2100–3000 m. Mexico; Guat, Hond, Salv, CR, Pan; Gr Ant; Col, Ven, Guy, Ec, Peru, Bol. Selected Specimens Examined. Chis (Breedlove 15097, US, 22800, DS, MEXU, NY, 30146, DS, MEXU, MO, NY, 34423, MEXU, MO; Martı´nez S. & Garcia 17512, UAMIZ, MEXU; Mu¨nch 112, DS). Oax (Geith 95-M054, NY; Mickel 4898, NY, UC, US; Torres C. 10268, MEXU). Pue (Arse`ne 1683, MEXU).

Distribution. Pendent epiphyte in wet montane forests; 200– 2250 m. USA (Fla); Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Fr Gui, Ec, Galapagos, Peru, Braz, Bol, Parag. Selected Specimens Examined. Chis (Breedlove 29782, MEXU, NY). DF (Schaffner 3, in 1875, NY). Gro (Rzedowski & McVaugh 97, ENCB). Oax (Conzatti 3818, MEXU; Mexia 9141a, B, UC, US). Pue (Flores Castorena 660, ENCB, MEXU). Rev (Reyes Bonilla 41, CHAPA, INIF). SLP (Pringle 3976, B, CAS, MEXU, NY, UC, US). Tam (Herna´ndez et al. 1912, NY). Ver (Purpus 1990, NY, UC).

This species is very similar to H. wilsonii, but that species has much narrower leaves that are more widely spreading.

6. HUPERZIA HIPPURIDEA Huperzia hippuridea (Christ) Holub, Folia Geobot. Phytotax. 20: 73. 1985. Fig. 154A. Lycopodium hippurideum Christ in Pittier, Prim. Fl. Costaric. 3(1): 56. 1901. Urostachys hippurideus (Christ) Herter ex Nessel, Ba¨rlappgewa¨chse 88. 1939. Type. Costa Rica. Ca´rtago: Pa´ramo, massif facing east [de Buena Vista], Pittier 10619 (P, not found; isotypes US!).

Plants erect from decumbent bases; stems 15–37 cm ⫻ 2–3 mm. (without leaves), green to dark red, simple to 2(–3) times forked, the divisions subparallel; leaves spirally arranged in several ranks, widely spreading, entire, somewhat deflexed at base of plants, ascending at stem tips, linear-lanceolate, mostly 11–20 ⫻ 0.7–1 mm, occasionally with zones of shorter leaves (5 mm), midribs not evident; sporophylls similar to the vegetative leaves; sporangia evident in leaf axils.

The stems are typically green to stramineous in Costa Rica, but in Oaxaca, El Salvador (Seiler 158), and Chiapas (Breedlove 30146) the stems are red. Breedlove 25410, DS (Chiapas), though, has pale stems. This tendency toward red color is also well developed in the Greater Antilles. Huperzia hippuridea looks like a small-leaved H. pithyoides, but is much taller, terrestrial, has somewhat shorter leaves without thickened margins, and branches not as widely diverging. The identity of H. mexicana is unclear, as no type was designated. [Huperzia mexicana (Herter) Holub, Folia Geobot. Phytotax. 20: 73. 1985. Lycopodium mexicanum Herter, Bot. Jahrb. Syst. 43(Beibl. 98): 35, 49. 1909. Type. Mexico. “San Luis Potosı´, Oaxaca”] The name was later (Herter, 1949) synonymized under H. pithyoides. Smith (1981) separated H. mexicana (pendent epiphyte) from H. hippuridea (erect, terrestrial) on the basis of habit and habitat, though Seiler (158, NY) reported that H. hippuridea may occupy both habitats, at least rarely, in El Salvador. Øllgaard (1983; in Davidse et al., 1995) separated H. mexicana (erect, epiphytic, but frequently terrestrial with seasonal zones of shorter leaves) from H. hippuridea (erect, terrestrial with regular leaf length) on the basis of habitat and presence or absence of regular constrictions. It is also closely related or identical to H. montana Underw. & F. E. Lloyd of the West Indies (Cuba, Jamaica, Hispaniola). Huperzia lechleri (Hieron.) Holub of Bolivia and H. mandioccana (Raddi) Trevis. of Brazil belong to the same complex.

7. HUPERZIA LINIFOLIA Huperzia linifolia (L.) Trevis., Atti Soc. Ital. Sci. Nat. 17: 248. 1874. Fig. 156C.

HUPERZIA ORIZABAE Lycopodium linifolium L., Sp. Pl. 2: 1100. 1753. Urostachys linifolius (L.) Herter, Repert. Spec. Nov. Regni Veg. 19: 165. 1923. Lectotype (chosen by Proctor, Fl. L. Antill. 2: 26. 1977). Plumier, Traite´ Foug. Ame´r., pl. 166, f. C, 1705, illustrating a plant from Martinique or Hispaniola.

Plants pendent, flaccid, 13–40 cm long, 2–5 times forked; stems 0.5–1.2 mm diam. (without leaves), stramineous to greenish or light brown; leaves spreading to oblique (somewhat ascending), distant, mostly 1–2 mm apart, spiral in few (4–6) ranks, twisted (with branches appearing somewhat flattened), linearlanceolate, narrowed to bases or short-petiolate, straight or usually subfalcate, usually pale or yellowish green, papyraceous to chartaceous, entire, midveins prominent abaxially, leaves near base of plants 1.5–2.5 cm ⫻ 2–3 mm, abruptly narrowed to 0.5 mm at leaf bases, smaller toward tips of stems where they are 1–2 cm long, 1 mm wide above narrowed bases; sporophylls similar to vegetative leaves but slightly smaller; 2n⫽260–280 (PR).

Distribution. Epiphytic in wet forests at low to middle elevations; 25–2000 m. Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Sur, Ec, Braz, Bol. Selected Specimens Examined. Chis (Breedlove 30858, DS, MEXU, NY, 31518, DS). Gro (Lorea 2408, 3026, FCME). Oax (Mickel 1038, ENCB, MEXU, NY, US; Wendt et al. 3630, CAS, CHAPA, ENCB, MEXU, MO, NY). Pue (Tejero & Castilla 152, ENCB, IZTA). Tab (Herna´ndez X. 208, CHAPA). Ver (Pringle 5929, MEXU; Purpus 1991, UC).

Mexican material of this species is fairly uniform. Nessel’s varieties (1939) are not sufficiently distinct to be used with confidence. Narrowed leaf bases and leaves not in pseudowhorls distinguish this species from slender specimens of the H. taxifolia alliance. In Mexico, Huperzia linifolia is closest to H. capillaris, which see for discussion.

8. HUPERZIA MYRSINITES Huperzia myrsinites (Lam.) Trevis., Atti Soc. Ital. Sci. Nat. 17: 248. 1874. Fig. 155A.

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Lycopodium myrsinites Lam., Encycl. 3: 654. 1789 [1792]. Plananthus myrsinites (Lam.) P.-Beauv., Prodr. Aethe´ogam. 111. 1805. Urostachys myrsinites (Lam.) Herter, Repert. Spec. Nov. Regni Veg. 19: 166. 1923. Type. “S. Domingue” [Hispaniola], Martin s.n. (P–Lam.; isotype P!). Lycopodium skutchii Maxon, Proc. Biol. Soc. Wash. 46: 159. 1933. Type. Guatemala. Chimaltenango: Chichavac, Skutch 243 (US!).

Stems mostly 20–60 cm long, 1 mm diam (without leaves), forked mostly 4–8 times; leaves dimorphic, forming distinct strobili; vegetative leaves spreading, mostly not imbricate, spirally arranged in several ranks with bases twisted to give branches a flattened appearance, leaves near base of stems 7–11 ⫻ 2.5–3.5 mm, narrowly ovate to oblong-lanceolate, entire, bases rounded, abruptly narrowed to 0.75 mm wide at attachment, subsessile, apices acuminate, midribs prominent abaxially; sporophylls much smaller than other leaves, pseudodecussate, narrowly to broadly ovate, 2 ⫻ 1.5–2 mm, in 4 ranks; sporangia evident.

Distribution. Pendent epiphytes at middle elevations wet montane forests on Atlantic slope, pine-oak forests with fir and Podocarpus; 1150–3000 m. Mexico; Guat, Hond, Salv, Nic, CR; Hisp; Col, Ven, Trin, Guy, Ec, Peru. Selected Specimens Examined. Chis (Anderson & Anderson 5469, NY; Breedlove 28198, DS, MO, NY, 31853, CAS, ENCB, NY; Matuda 5244, DS, MEXU, US). Gro (Lorea 2351, 2358, ENCB, FCME). Oax (Mickel 5625, 6622, 7093, NY). Ver (Cha´zaro & Herna´ndez 4050, XAL).

Lycopodium myrsinites is conspicuous by its broad, short sterile leaves and much-reduced sporophylls.

9. HUPERZIA ORIZABAE Huperzia orizabae (Underw. & F. E. Lloyd) Holub, Folia Geobot. Phytotax. 20: 75. 1985. Figs. 157J–L. Huperzia orizabae Underw. & F. E. Lloyd, Bull. Torrey Bot. Club 33: 110. 1906. Urostachys orizabae (Underw. & F. E. Lloyd) Herter ex Nessel, Ba¨rlappgewa¨chse 175. 1939. Type. Mexico. Veracruz: Orizaba, Sierra de Agua, Mu¨ller 690 (NY!; isotypes BR, P, US!).

Plants pendent, 10–32 cm long, with stems forking 1–4(–6) times, 2–3 mm diam. (without leaves), stramineous to brown; leaves in only 4–6 ranks, somewhat distant, chartaceous, linearlanceolate, ca. 0.7–1.8 cm ⫻ 1–2 mm, widest near the centers,

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HUPERZIA

midribs prominent, mostly widely spreading, deflexed at base of plants, somewhat ascending and becoming smaller towards plant apices; sporophylls like the regular leaves but slightly smaller; sporangia evident among the spreading sporophylls.

Distribution. Epiphytic in wet montane rain forests, pine-oak forests, Liquidambar forests, lower montane rain forests; 900– 2250 m. Mexico; Guat, Hond, Salv, Nic, CR; Gr Ant; Col, Ven. Distribution. Epiphytic in wet montane forests of Atlantic slope; 1300–2200 m. Mexico; Guat, Hond, Salv. Specimens Examined. Chis (Breedlove 15233, US, 28201, DS, NY, 32643, DS; Tillett 636-38, US). Gro (Campos 1519, FCME). Oax (Hallberg 1404a, NY; Mickel 3771, NY). Ver (Mu¨ller 690, NY, US).

Leaf shape in Huperzia orizabae is like that of H. linifolia, but the stems are much thicker (2–3.5 mm) and widely branching (spreading). It is related to Huperzia bradeorum Christ of Guatemala to Ecuador, which is an erect epiphyte and appears to have smaller leaves (less than 1 cm) at the base of the plants.

Selected Specimens Examined. Chis (Breedlove 31419, DS, MEXU, NY, 53544, ENCB, NY). Gro (Lorea 3024, FCME). Me´x (Schaffner 1, NY). Oax (Conzatti 3547, MEXU, US). Qro (Servı´n 973, CAS, IEB). SLP (Pringle 3975, DS, UC). Tam (Sharp et al. 52289, US). Ver (Copeland herb. 162, MEXU, MICH, UC; Nee et al. 26380, F, NY, XAL).

Huperzia pithyoides is close to H. hippuridea, but stouter and with longer leaves. Since it grows high in trees, it is probably more common than the few collections would suggest.

11. HUPERZIA PRINGLEI Huperzia pringlei (Underw. & F. E. Lloyd) Holub, Folia Geobot. Phytotax. 20: 76. 1985. Figs. 155B, 158A–E.

10. HUPERZIA PITHYOIDES Huperzia pithyoides (Schldtl. & Cham.) Holub, Folia Geobot. Phytotax. 20: 76. 1985. Figs. 154C, D. Lycopodium pithyoides Schltdl. & Cham., Linnaea 5: 623. 1830. Urostachys pithyoides (Schltdl. & Cham.) Herter, Repert. Spec. Nov. Regni Veg. 19: 164. 1923. Type. Mexico. Veracruz: prope Jalapa, Schiede & Deppe s.n. (B!). Deppe 832 (BM, HBG), annotated as Lycopodium pithyoides, n. sp., possible isotypes. Lycopodium gigas Herter, Bot. Jahrb. Syst. 43(Beibl. 98): 50. 1909. Urostachys gigas (Herter) Herter, Repert. Spec. Nov. Regni Veg. 19: 164. 1923. Lectotype (chosen by Herter, 1923: 164). “Su¨dmexiko. Epiphyt auf Ba¨umen, besonders Eichen, Ha¨nge der Cordilleren, 2000 m,” Haarstro¨m s.n. (P).

Plants erect to suberect; stems mostly 20–30 cm long, 4–5 mm diam. (without the leaves), dark red (rarely light brown), 2–4 times forked, the divisions often widely divergent; leaves chartaceous, spirally arranged in many ranks, green, basal 3 mm red on abaxial surface with green margins, widely spreading (deflexed at base of plants, ascending at stem tips), linear, mostly (2–)3–4 cm ⫻ 0.7–1 mm at bases, midribs immersed, not visible, margin entire, revolute (when dry); sporophylls similar to vegetative leaves in form, size and habit.

Lycopodium pringlei Underw. & F. E. Lloyd, Bull. Torrey Bot. Club 33: 109. 1906. Urostachys pringlei (Underw. & F. E. Lloyd) Herter ex Nessel, Ba¨rlappgewa¨chse 131. 1939. Type. Mexico. Oaxaca: Sierra de Clavelinas, Pringle 4994 (NY!; isotypes DS!, GH!, JE, K!, MEXU!, MO!, NY!, UC!, US!). Lycopodium tuerckheimii Maxon, Contr. U.S. Natl. Herb. 13: 23, t. 1. 1909. Huperzia tuerckheimii (Maxon) Holub, Folia Geobot. Phytotax. 20: 77. 1985. Urostachys. tuerckheimii (Maxon) Herter, Repert. Spec. Nov. Regni Veg. 19: 163. 1923. Type. Guatemala. Coba´n, Tu¨rckheim II. 1864 (US!, photo B!).

Plants pendent, mostly 20–70 cm long; stems 4–8 times forking, the divisions narrowly to widely divergent (40–140⬚), 0.5–1 mm diam. (without leaves); leaves linear to linear-lanceolate, spirally arranged in several (ca. 6) ranks, spreading and somewhat recurved, to ascending or even appressed, leaves mostly 4–8 ⫻ 0.4–0.6 mm; sporophylls roughly 4-ranked, broadly lanceolate to acuminate, 2–3 ⫻ 0.75–1 mm; sporangia evident among the sporophylls.

HUPERZIA SERRATA

335

Lycopodium reflexum Lam., Encycl. 3: 653. 1789 [1792]. Urostachys reflexus (Lam.) Herter, Beih. Bot. Centralbl. 39: 249. 1922. Type. Martinique. Martin s.n. (P-LA 442!).

Plants erect, tufted or shortly decumbent; stems erect, mostly 9–30 cm tall, 1–2 mm diam. (without leaves), stramineous to light brown, simple to 3 times forked, the isodichotomous divisions subparallel or weakly diverging; leaves widely spreading, spirally arranged in many ranks, strongly deflexed toward the base of plants (ascending only at stem tips), linear-lanceolate, mostly 5–7 ⫻ 0.5–1 mm at base, margins minutely ciliatedenticulate, adaxial surfaces lustrous, midribs immersed, not evident; sporophylls similar to other leaves; sporangia evident; 2n⫽ca. 256 (Jam). Distribution. Pendent epiphyte in wet montane forests/cloud forests, oak forests, pine-oak forests, oak-Drimys forests; 1150– 2750 m. Mexico; Guat, Hond, Salv. Selected Specimens Examined. Chis (Breedlove 50083, CAS, ENCB, MO, NY). Gro (Lorea 2974, FCME, NY). Hgo (Gimate 976, CHAPA, ENCB, MEXU, NY). Jal (McVaugh 13908, MEXU, NY). Me´x (Hinton 8936, NY). Mich (McVaugh 22809, ENCB, MEXU, MICH, NY). Mor (Rose 4416, NY, US). Oax (Mickel 5365, ENCB, NY, UC, 7051, NY). Ver (Mu¨ller 361, NY).

The type of H. pringlei (Pringle 4994, NY) has appressed juvenile leaves 4–5 mm long, widely divergent dichotomies (40– 65⬚) and narrow (1–2 mm), almost uninterrupted strobili with appressed ovate sporophylls. In describing H. tuerckheimii, Maxon cited its divisions as being widely divergent, very slender (1–1.5 mm) and scarcely interrupted, and the sporophylls deltateovate. The distinctions between H. pringlei and H. tuerckheimii (Smith, 1981), based on elevation (2000–3000 m vs. 1300 m), sporophyll shape (ovate vs. lanceolate), the aspect of oldest leaves (spreading vs. appressed), and width of strobilus (interrupted, greater than 1.5 mm diam. vs. uninterrupted, less than 1.5 mm) do not appear to separate the two species. Huperzia pringlei is also close to H. filiformis (Sw.) Holub of Central and South America to Bolivia and Hawaii (type from Hawaii), and is only marginally, if at all, different from it. Øllgaard (1983) considered the material from southern Central America, northern South America, and the West Indies, commonly referred to as Huperzia verticillata (L.f.) Trevis. or H. setacea Lam. (both based on Old World material), to differ from the Old World material and used the name H. acerosa Sw. for neotropical material. Rare in Guatemala but common in the West Indies, H. acerosa can be distinguished from H. pringlei by the leaves of H. acerosa being of almost uniform length, only slightly shorter and narrower toward the fertile apices (juvenile leaves 4–6 ⫻ 0.2–0.3 mm, mature sporophylls 3–4 ⫻ 0.5–0.7 mm). It is rare at lower elevations in Guatemala (900–1500 m) and not found in Mexico.

Distribution. Terrestrial, rarely epiphytic on open banks at lower to upper middle elevation wet forests, montane rain forests, oak forests; 300–2400 m. Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Ec, Peru, Braz, Bol, Parag. Selected Specimens Examined. Chis (Breedlove 23137, MO, NY, 31608, DS, MEXU, NY). Gro (Mexia 9038, B, UC, NY). Hgo (Breedlove 59573, CAS, MO, NY). Jal (Mexia 1517, CAS, DS, MO, NY, UC, US). Me´x (Hinton 3378, MEXU, US). Oax (Mickel 4803, NY; Rzedowski 32823, ENCB, MO). Pue (Fraile et al. 76, ENCB, UAMIZ). Ver (Pringle 8146, BM, K, MO, MEXU, NY, UC).

Huperzia reflexa is distinct by its erect, dichotomously forking stems with spreading leaves and conspicuous sporangia. There are many splinter names for slight variations in Costa Rica and South America, but Mexican material of this very common species is uniform and compares favorably with the West Indian material of the type variety. It is reported by Smith (1981) to be rarely epiphytic; further observations are needed to confirm the epiphytic habitat.

13. HUPERZIA SERRATA

12. HUPERZIA REFLEXA

Huperzia serrata (Thunb. ex Murray) Trevis., Atti Soc. Ital. Sci. Nat. 17: 248. 1874. Figs. 157A–C.

Huperzia reflexa (Lam.) Trevis., Atti Soc. Ital. Sci. Nat. 17: 248. 1874 [1875]. Figs. 154E, F.

Lycopodium serratum Thunb. ex Murray, Syst. Veg., ed. 14, 944. 1784. Urostachys serratus (Thunb.) Herter, Bot. Arch. 3: 13. 1923. Type. Japan. Thunberg s.n. (UPS-Thunb. 25333B; microfiche NY!, UC!).

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HUPERZIA

Lycopodium sargassifolium Liebm., Overs. Kongel. Danske Vidensk. Selsk. Forh. Medlemmers Arbeider 4: 47. 1847 [1848]. Type. Mexico. Oaxaca: Dto. Villa Alta, Tepitongo, Liebmann s.n. [Pl. Mex 2027] (C!; isotypes K, frag. NY!, UPS!, US!). Urostachys serratus (Thunb.) Herter var. japonica-neotropicus Herter ex Nessel, Revista Sudamer. Bot. 6: 159. 1940. Lectotype (chosen by Mickel & Beitel, 1988: 241). Cuba. Summos monte Pico, Ekman 5498 (S!; isolectotypes B!, BONN, F!, G).

Plants erect, long-decumbent at bases, 3–20 cm; stems mostly 12–16 cm tall, 1.5–2 mm diam. (without leaves), green, divisions isodichotomous, older portion of stems yellow, prominent annual constrictions from formation of shorter leaves at end of growing season, slight reduction of leaf size from juvenile portions to mature portions; leaves deflexed near stem bases, spreading toward stem tips, spirally arranged in many ranks, no stomata on adaxial surfaces, largest leaves narrowly obtrullate with apices abruptly acuminate (7–16 ⫻ 1–3.5 mm above middle), narrowed to petiole-like bases (0.8–1 mm wide), smallest leaves (at annual constrictions) obtrullate to lanceolate (3–6 ⫻ 0.75– 1.25 mm above middle), narrowed to petiole-like bases (0.5–1 mm), deeply serrate, sometimes doubly so, with 5–15 large teeth curving towards tips; sporangia evident in zones, asexual 3-lobed propagules (gemmae) 4–4.5 ⫻ 3.5–4 mm produced at end of year’s growth on gemmiphores (short stems 3.5–5 mm with 6 pairs of leaves), spirally placed among leaves; 2n⫽528 (India).

Distribution. Terrestrial in wet montane forests on Atlantic slope, pine-oak forest; 1200–2130 m. Mexico; Cuba, Haiti; Hawaii, Japan, China. Specimens Examined. Hgo (Breedlove 59467, CAS, MEXU, UC; Guevara F. s.n., FCME; Nun˜ez F. s.n., 28 Nov 1980, FCME; Reyes R. s.n., FCME). Oax (Liebmann s.n. [Pl. Mex. 2027], C, NY, UPS, US).

The major portion of this species’ range is in the Old World with disjunct, montane populations in wet forests of Mexico, Cuba, and Hispaniola. The type of Huperzia sargassifolia corresponds to Old World material of H. serrata. Large, serrate leaves exhibiting annual cycles of shorter leaves, and asexual gemmae (and gemmiphores that stay on the plant once the gemmae have been dispersed by rain or wind) are distinguishing characters of H. serrata.

14. HUPERZIA TAXIFOLIA Huperzia taxifolia (Sw.) Trevis., Atti Soc. Ital. Sci. Nat. 17: 248. 1874. Fig. 156A. Lycopodium taxifolium Sw., Prodr. 138. 1788. Urostachys taxifolius (Sw.) Herter, Repert. Spec. Nov. Regni Veg. 19: 162. 1923. Type. Jamaica. Swartz s.n. (S). Lycopodium nitens Schldtl. & Cham., Linnaea 5: 623. 1830. Type. Mexico. Veracruz: Jalapa, Schiede 831 (B!; isotypes BM, K!, US! frag. ex B, NY! frag. ex K).

Stems 20–60 cm long, 1–3 mm diam. (without leaves), arching to pendent, (2–)3–5(–7) times forked; leaves spirally arranged in six ranks, strongly to weakly imbricate, linear-lanceolate, 1.2– 1.5 cm ⫻ 1.5–2 mm at bases, only slightly narrowed at bases, slightly spreading (more so at base of plants) to ascending, subcoriaceous, entire, midribs immersed but distinct, branches more distinctly zoned at higher elevations, where leaves more imbricate; sporophylls similar to other leaves but somewhat smaller, linear-lanceolate with expanded bases (1 mm), but gradually reduced toward stem apices to 7 mm long; sporangia evident among loosely arranged sporophylls.

Distribution. Pendent or arching epiphytes in wet montane forests or cloud forests, often with Liquidambar; 100–1800(– 2250) m. Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Ec, Peru, Bol. Selected Specimens Examined. Chis (Dressler 1580, UC, US). DF (Arse`ne 10988, US). Gro (Fonseca 186, FCME). Hgo (Herna´ndez M. & Herna´ndez V. 4767, MEXU). Jal (Iltis et al. 7899, UC, WIS). Me´x (Tejero-Dı´ez 2987, IZTA). Oax (Hallberg 1417, NY, US). Pue (Rzedowski 29966, ENCB, NY). Sin (Vega A. et al. 3134, ENCB). SLP (Schaffner 2, NY). Tam (Sharp et al. 52268, MEXU, US). Ver (Purpus 7763, UC).

Huperzia taxifolia is highly variable, with some specimens having strongly coriaceous, spreading leaves. Several robust specimens of Huperzia taxifolia occur at 1900–2900 m, and approach H. cuernavacensis: their leaves are spreading, not lustrous, and they have loose strobili (see comments under H. cuernavacensis). A collection by Gentry & Argu¨elles (18188, NY) from a cliff face in Sinaloa has tightly appressed leaves with revolute margins and

HYMENOPHYLLUM

may be another environmental variant of H. taxifolia or H. cuernavacensis. Lycopodium schwendeneri Herter may be synonymous with Huperzia taxifolia, though no type was designated and Herter indicated only “Mexico” for type locality.

15. HUPERZIA WILSONII

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Øllgaard (1983) reported that plants become pendent with age. It is rare in Guatemala at 1300–1500 m. Huperzia wilsonii closely resembles H. dichotoma but is distinct in its very narrow leaves.

5 6 . H Y M E N O P H Y L L UM

Huperzia wilsonii (Underw. & F. E. Lloyd) B. Øllg., Opera Bot. 92: 170. 1987. Fig. 156B.

Hymenophyllum Sm., Me´m. Acad. Roy. Sci. (Turin) 5: 418. 1793 [1794]. Lectotype (chosen by C. Presl, Hymenophyllaceae 31. 1843): Hymenophyllum tunbrigense (L.) Sm.

Lycopodium wilsonii Underw. & F. E. Lloyd, Bull. Torrey Bot. Club 33: 111. 1906. Urostachys wilsonii (Underw. & F. E. Lloyd) Herter, Repert. Spec. Nov. Regni Veg. 19: 163. 1923. Type. Puerto Rico. Luquillo Mts., Wilson 271 (NY!). Lycopodium trichodendron Herter, Bot. Jahrb. Syst. 43(Beibl. 98): 49. 1909. Urostachys trichodendron (Herter) Herter, Repert. Spec. Nov. Regni Veg. 19: 163. 1923. Lectotype (chosen by Herter, 1923: 163). Guadeloupe. Bory 103 (P). Lycopodium arcanum Maxon ex Yuncker, Publ. Field Mus. Nat. Hist., Bot. Ser. 17: 310, t. 3. 1938. Type. Honduras. Comayagua: Above El Achote, Yuncker, Dawson & Youse 6149 (US!).

Sphaerocionium C. Presl, Hymenophyllaceae 33. 1843. Lectotype (chosen by Copel., Philipp. J. Sci. 64: 10. 1937): Sphaerocionium hirsutum (L.) C. Presl [⬅ Hymenophyllum hirsutum (L.) Sw.]. Hymenophyllum subg. Mecodium C. Presl ex Copel., Philipp. J. Sci. 64: 93. 1937. Mecodium (C. Presl ex Copel.) Copel., Philipp. J. Sci. 67: 17. 1938. Lectotype (chosen by Copel., Phillip. J. Sci. 64: 10. 1937): Hymenophyllum polyanthos (Sw.) Sw. [⬅ Trichomanes polyanthos Sw.]. For more complete synonymy, see Morton, 1968.

Plants ca. 20 cm long, erect when young, becoming pendent, 4–6 times dichotomous; stems ca. 1–1.5 mm diam. (not including leaves), stramineous to light brown; leaves spirally arranged in several ranks, sparsely covering the stems, spreading to somewhat ascending, linear, 1–1.3 cm ⫻ ca. 0.2–0.3 mm, leaf bases often tinged with red; sporophylls resembling vegetative leaves, not expanded at bases; sporangia conspicuous among the sparse narrow leaves.

Mostly epiphytic, less commonly epipetric or even terrestrial; rhizomes long-creeping, thread-like, hairy, the hairs basifixed or centrally attached; fronds small; stipes slender to stout, glabrous to hairy; blades pinnatifid to several times pinnate; laminae generally one cell thick, often completely glabrous, some with hairs on leaf tissue, veins, and/or margins; sori marginal, in bilabiate involucral cups; sporangia borne on short to long receptacles; spores tetrahedral-globose, green; gametophytes of branched ribbons. In a broad sense, the genus contains over 300 species, about one third of them neotropical, with a few species into wet temperate areas. Some authors, such as Copeland (1937), have split the genus into several genera. Hymenophyllum is distinct in its thin blades and bilabiate involucres with receptacles usually not exserted; x⫽11, 13, 18, 21, 22, 28, 36. References

Distribution. Pendent epiphytes of wet montane forests on Atlantic slope; 1400–1600 m. Mexico; Guat, Nic, CR, Pan; Jam, PR, L Ant; Col, Ven, Trin, Ec, Peru. Specimens Examined. Oax (Hallberg 1416, NY; Rzedowski 34051, ENCB).

Copeland, E. B. 1937. Hymenophyllum. Philipp. J. Sci. 64: 1–188; Hennequin, S., A. Ebihara, M. Ito, K. Iwatsuki & J.-Y. Dubuisson. 2003. Molecular systematics of the fern genus Hymenophyllum s.l. (Hymenophyllaceae) based on chloroplastic coding and noncoding regions. Molec. Phylogen. Evol. 27: 283– 301; Morton, C. V. 1947. The American species of Hymenophyllum section Sphaerocionium. Contr. U.S. Natl. Herb. 29: 139–199; Morton, C. V. 1968. The genera, subgenera, and sections of the Hymenophyllaceae. Contr. U.S. Natl. Herb. 38: 153–214; Pacheco, L. 1994. Pteridofitas: Familia Hymenophyllaceae. Flora de Mexico 6(2): 1–56. Consejo Nacional de la Flora de Me´xico, Me´xico, D. F.; Pryer, K. M., A., R. Smith, J. S. Hunt, & J.-Y. Dubuisson. 2001. rbcL data reveal two monophyletic groups of filmy ferns (Filicopsida: Hymenophyllaceae). Amer. J. Bot. 88: 1118–1130.

Key to the Mexican Species of Hymenophyllum 1. Blade margins toothed. 2. Sori perpendicular to the laminae; stipes and rachises essentially glabrous; ultimate segments 0.7–1 mm wide; fronds commonly 4–12 cm long. ..................................................................................................................................................................... 21. H. tunbrigense. 2. Sori in plane of laminae; stipes and rachises with sparse hairs abaxially; ultimate segments 1–2 mm wide; fronds commonly 8– 20 cm long.

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HYMENOPHYLLUM

3. Indusial margins entire to serrulate; stipes usually more than 2 cm long. ................................................................................... 9. H. fucoides. 3. Indusial margins deeply laciniate; stipes mostly less than 2 cm long. ...................................................................................... 6. H. ectocarpon. 1. Blade margins entire. 4. Blades glabrous; rhizome hairs centrally attached, sessile. 5. Fronds 0.8–2.5(–3) cm long; basal half of involucres immersed; rhizomes 0.1–0.2 mm diam.; rhizome hairs reddish brown, 0.5–0.8 mm long. 6. Blades mostly 2–4 cm long; involucres in a dense cluster at tips of blades. ............................................................... 15. H. paucicarpum. 6. Blades mostly less than 2 cm long; involucres one to several at blade tips, not tightly clustered. 7. Stipes alate to bases; ultimate segments 0.5–0.8(–1) mm wide. ................................................................................... 3. H. brevistipes. 7. Stipes not alate; ultimate segments 1.5–2 mm wide. ..................................................................................................... 1. H. abruptum. 5. Fronds 3–25 cm long; basal 1⁄4–1⁄3 of involucres immersed, sessile or appearing stalked; rhizomes 0.3–0.5 mm diam.; rhizome hairs reddish brown to whitish tan, 0.8–2 mm long. 8. Blades essentially pinnate or pinnatifid, primary segments simple to bifid; ultimate segments 2.5–3 mm wide; rhizome hairs whitish cream, 1 mm long (tip to tip); stipes non-alate. .......................................................................................... 2. H. asplenioides. 8. Blades bipinnatifid to tripinnate; ultimate segments 0.8–1.3 mm wide; rhizome hairs reddish brown; stipes alate or not. 9. Ultimate segments 1–1.3 mm wide, narrowed near forks to 0.5 mm; involucres 0.8–1 mm ⫻ 1–1.5 mm, greatly constricted at bases to appear stalked (0.5 mm wide); rhizome hairs 1.5–2 mm long (tip to tip); stipes narrowly alate to bases. ................................................................................................................................................................ 14. H. myriocarpum. 9. Ultimate segments 0.8–1 mm wide; involucres 1–1.5 ⫻ 0.8–1 mm wide, basal 1⁄4–1⁄3 immersed; rhizome hairs 0.8–1 mm long (tip to tip); stipes alate to bases or non-alate. 10. Blade margins plane; stipes alate to non-alate; rachises alate throughout; pinnae adnate or the basal pinna pair short-stalked. ..................................................................................................................................................... 16. H. polyanthos. 10. Blade margins undulate or crisped; stipes non-alate; proximal half of rachises non-alate or essentially so; proximal 2–4 pinna pairs short-stalked. 11. Segments strongly crisped; pinnae imbricate. ........................................................................................... 7. H. fendlerianum. 11. Segments plane to only slightly crisped; pinnae not imbricate. ............................................................... 22. H. undulatum. 4. Blades hairy; rhizome hairs simple. 12. Hairs present on laminar surfaces, margins, and veins. 13. Stipes stout, 0.4–1 mm or more diam. 14. Blades oblanceolate, 2.5–8 cm wide; fronds pendent; pinnae reduced at base of blades, narrow, pinnate-pinnatifid; rachises and stipes broadly alate. .......................................................................................................................................... 18. H. sieberi. 14. Blades ovate to ovate-lanceolate, 5–10 cm wide; fronds arching; basal pair of pinnae the longest or nearly so, pinnate-bipinnatifid; stipes not alate. ..................................................................................................................... 4. H. crassipetiolatum. 13. Stipes filiform, 0.1–0.25 mm diam. 15. Pinnae mostly simple and linear, the blades occasionally bipinnatifid or pinnate-bipinnatifid, segments 1–1.3 mm wide; stellate hairs of leaf surfaces and margins stalked, stalks 0.1 mm long; rhizome hairs 0.5 mm (see also H. fragile) long; stipe hairs with 4–5 rays. ............................................................................................................................. 11. H. lanatum. 15. Pinnae mostly pinnatifid, segments 1–2 mm wide; stellate hairs of leaf surfaces and margins sessile; rhizome hairs 0.5–1 mm long; stipe hairs simple, forked, or stellate. 16. Stipe hairs mostly simple; rachises not alate to basal pinnae; fronds 8–25 cm long; rhizome hairs 1 mm long; segments 1.8–2 mm wide. ...........................................................................................................................................19. H. tegularis. 16. Stipe hairs mostly forked or stellate; rachises alate throughout, stipes alate at distal end; fronds 3–6 cm long; rhizome hairs 0.5 mm long; segments 1–1.3 mm wide. .............................................................................................. 8. H. fragile. 12. Hairs restricted to blade margins and veins. 17. Stipes not at all alate; rachises not alate to basal pinnae. 18. Marginal hairs sessile, stellate; vein hairs abundant, stellate, subsessile and stalked, pubescence dense, stellate, arms long (at first glance blade appears hairy); indusial margins with stellate, subsessile hairs. ......................... 17. H. pulchellum. 18. Marginal hairs with stalks 0.1 mm long, forked or stellate; vein hairs scattered, forked to stellate, stalked, pubescence obviously confined to veins; indusial margins with simple hairs. ..................................................... 20. H. trapezoidale. 17. Stipes alate, at least in the distal 1⁄4; rachises alate throughout. 19. Stipes alate one half to full length. ................................................................................................................................. 10. H. hirsutum. 19. Stipes alate only in the distal 1⁄4 to 1⁄2. 20. Stipes 0.2–0.3 mm diam.; blades pinnate-pinnatifid with only 1–2 segment pairs per pinna. .......................... 5. H. crispum. 20. Stipes 0.2–1 mm diam.; blades pinnate-bipinnatifid. 21. Blade margins with stalked forked or stellate hairs; blades lanceolate. ........................................................ 12. H. maxonii. 21. Blade margins with simple or sessile forked hairs; blades ovate-deltate. ............................................... 13. H. microcarpum.

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1. HYMENOPHYLLUM ABRUPTUM Hymenophyllum abruptum Hook., Sp. Fil. 1: 88, t. 31B. 1844. Figs. 163A, B. Mecodium abruptum (Hook.) Copel., Philipp. J. Sci. 67: 26. 1938. Syntypes. Jamaica. Menzies s.n., Purdie s.n. (K! isosyntype of Purdie s.n. at BM!).

Similar to H. brevistipes in its small size, but distinct in the non-alate stipes, less divided (pinnatifid) blades with wider segments (1.5–2 mm vs. less than 1 mm); 2n⫽56 (Jam.).

Distribution. Epiphytic in wet montane forests; 300–1800 m. Mexico; Guat, Hond, Nic, CR, Pan; Cuba, Jam; Ven, Guy, s Braz. Selected Specimens Examined. Chis (Breedlove 49947, 53564, ENCB, 56216, ENCB, NY). Oax (Hallberg 1435, ENCB; Mickel 981, 995, ENCB, 5756, ENCB, NY, 7115, ENCB). Pue (Vera Santos 3352, US). Ver (Mu¨ller 432, NY).

Hymenophyllum asplenioides is distinct by its glabrous, pinnatifid to pinnatisect blades. Distribution. Epiphytic in montane rain forests; 1300 m. Mexico; Guat, Bel, Hond, CR, Pan; Cuba, Jam, Hisp; Fr Gui. Specimens Examined. Chis (Breedlove 29591, 32205, DS).

2. HYMENOPHYLLUM ASPLENIOIDES Hymenophyllum asplenioides (Sw.) Sw., J. Bot. (Schrader) 1800(2): 98. 1801. Figs. 161A, B. Trichomanes asplenioides Sw., Prodr. 136. 1788. Mecodium asplenioides (Sw.) Copel., Philipp. J. Sci. 67: 26. 1938. Type. Jamaica. Swartz s.n. (S).

Rhizomes long-creeping, 0.3 mm diam.; rhizome hairs centrally attached, sessile, whitish cream, dense, 1 mm long; fronds 4–20 cm long, distant; stipes 0.1–0.2 mm diam., non-alate, glabrous; blades oblong, pinnatisect to pinnatifid, 1–3 cm wide; pinnae 4–15 pairs, decurrent, shallowly pinnatifid, oblong, 2.5–3 mm wide; margins entire, glabrous; rachises alate; involucres sessile, not immersed nor stalked, nearly round, entire, 1–1.5 mm wide, receptacles not exserted; 2n⫽72 (Jam).

3. HYMENOPHYLLUM BREVISTIPES Hymenophyllum brevistipes Liebm., Mexic. Bregn. 290 (reprint 138). 1849. Lectotype (chosen by Smith, 1981: 130). Mexico. Veracruz: San Antonio Huatusco, Liebmann s.n. [Pl. Mex. 2127, Fl. Mex. 557] (C!). Figs. 161H, J, 163C–E. Rhizomes creeping, 0.1–0.2 mm diam.; rhizome hairs common, centrally attached, sessile, reddish brown, 0.5–0.8 mm long; fronds 1.5–2.5 cm long, distant; stipes 1–1.2 cm long, 1⁄2 the frond length, 0.1 mm diam., alate to bases; blades pinnatifid, subflabellate, 1–2 cm wide, glabrous; rachises alate; pinnae 1–3 pairs; segments simple to 1–2-bifid, 0.5–0.8(–1) mm wide, entire, glabrous; margins plane, entire; sori terminating distal segments, involucres immersed in basal 1⁄2, 1.5–2 ⫻ 1.5–2 mm, entire.

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HYMENOPHYLLUM

Distribution. Epiphytic, or less commonly terrestrial in wet montane forests; 750–1900 m. Mexico; Guat, CR, Pan; Cuba, Jam; Col, Ven, Braz. Selected Specimens Examined. Chis (Breedlove 21968, NY). Oax (Mexia 9205, NY; Mickel 1591, 4718a, 5753, 6532, 6671, 7209, NY). Ver (Liebmann 2124, 2127, C).

Hymenophyllum brevifrons Kunze (type from French Guiana) differs in its non-alate stipes and slightly wider segments, but is very closely allied. Some workers (e.g, Mickel & Beitel, 1988) have recognized it as occurring in Mexico, but the stipe of H. brevistipes is often extremely narrow and inconspicuous to essentially absent. We place specimens previously identified as H. brevifrons under H. brevistipes. Further work is needed to resolve the systematics of this group.

4. HYMENOPHYLLUM CRASSIPETIOLATUM Hymenophyllum crassipetiolatum Stolze, Amer. Fern J. 66: 15, f. 1. 1976. Type. Guatemala. Zacapa: Volca´n Gemelos, Steyermark 43302 (US!; isotypes F, GH!). Figs. 164A–D. Rhizomes long-creeping, ca. 0.6 mm diam.; rhizome hairs simple, orange-tan, 1.5–2 mm long; fronds 22–35 ⫻ 5–10 cm, distant; stipes 1⁄3–2⁄5 the frond length, 3–16 cm long, (0.5–)0.8–1 mm diam., nonalate, with numerous, stalked, stellate hairs; blades pinnate-pinnatifid or pinnate-bipinnatifid, ovate-lanceolate, attenuate at apices, proximal pinnae slightly reduced; rachises nonalate in proximal third, increasingly alate in distal third because of decurrent pinnae; pinnae to ca. 25 pairs, ascending at tips, to ca. 8 cm long, attenuate; ultimate segments 0.6–1 mm wide, subacute, plane; hairs of veins and leaf tissue scattered, mostly sessile, stellate (3–4-rayed), those of margins bistellate, located on small teeth; involucres orbicular, immersed, with valves ca. 1 mm diam.

Distribution. Epiphytic in montane rain forests; (200–)1400– 2800 m. Mexico; Guat, Hond, Salv. Specimens Examined. Chis (Calzada et al. 8773, IBUG, NY, UC, XAL; Matuda 2910, MEXU, MICH, NY, UC, US, 5233, DS, F, MEXU, MO).

This is the largest species of Hymenophyllum in Mexico and is most closely related to H. lindenii Hook., known from Colombia, Venezuela, and Ecuador. It also somewhat resembles H. microcarpum, but the latter has more widely deltate blades and lacks hairs on the laminar tissue, though hairs are present on veins and margins.

5. HYMENOPHYLLUM CRISPUM Hymenophyllum crispum Kunth, Nov. Gen. Sp. 1: 26. 1815. Figs. 159C, D. Sphaerocionium crispum (Kunth) Klotzsch, Linnaea 18: 537. 1844. Type. Venezuela. Silla de Caracas, Humboldt s.n. (P-Humb.!; B!). Sphaerocionium schiedeanum C. Presl, Hymenophyllaceae 60. 1843. Hymenophyllum schiedeanum (C. Presl) Bosch, Ned. Kruidk. Arch. 5: 191. 1863. Type. Mexico. Schiede s.n. (PRC?).

Rhizomes long-creeping, 0.3 mm diam.; rhizome hairs 1 mm long; fronds 4–20 ⫻ 1.5–5 cm, distant; stipes 0.2–0.3 mm diam., narrowly alate in distal 1⁄4, lightly pilose, hairs mostly simple; blades linear, pinnate-pinnatifid, proximal pinnae slightly reduced; rachises alate, undulate; segments 1–1.2 mm wide; margins undulate to crisped, entire; hairs on blade margins simple, hairs of costae rare, mostly stalked, stellate; indusial margins with simple hairs, receptacles not exserted.

HYMENOPHYLLUM FENDLERIANUM

341

Hymenophyllum fucoides and H. ectocarpon belong to a complex in need of further study. The latter is distinguished by the deeply laciniate indusial margins and stipes less than 2 cm long. Although specimens from Oaxaca fit the description for H. ectocarpon, both Smtih (1981) and Stolze (1976) reported the taxon from low elevations (300–1200 m), whereas in Oaxaca it occurs at high elevations. Pacheco (1994) included H. ectocarpon in H. fucoides.

7. HYMENOPHYLLUM FENDLERIANUM Hymenophyllum fendlerianum J.W. Sturm in Martius, Fl. Bras. 1(2): 291. 1859. Figs. 161E–G. Distribution. Epiphytic or terrestrial in cloud forests; 1500– 2700 m. Mexico; Guat, Hond, CR; Jam; Col, Ven, Ec, Peru, s Braz, Bol. Selected Specimens Examined. Chis (Breedlove 29433, MEXU, NY, 33668, DS, MEXU, 57071, CAS, ENCB). Gro (Hinton 14306, NY). Hgo (Gonza´lez G. 56, MEXU; Herna´ndez X. 873, US). Oax (Mickel 4147, 4911, 5288, NY). Ver (Heitz 292, XAL).

6. HYMENOPHYLLUM ECTOCARPON Hymenophyllum ectocarpon Fe´e, Me´m. Foug. 11: 115. 1866. Syntypes. Guadeloupe. “Matouba, Ravine Flore, Rivie`re Goyave,” 1861, L’Herminier s.n.; Martinique. Rivoire s.n. (P?). Fig. 162G. Similar to H. fucoides except: rhizome hairs 0.5 mm long; fronds 2.5–3.5 cm long; stipes mostly less than 2 cm long, sparsely hairy; segments 1–1.3 mm wide; marginal teeth 0.3–0.5 mm long; sori 1.8 mm long; involucral margins laciniate.

Distribution. Epiphytic in cloud forests; 800–2700 m. Mexico; Guat, Bel, Nic, CR, Pan; L Ant.

Mecodium fendlerianum (J.W. Sturm) Copel., Philipp. J. Sci. 67: 26. 1938. Hymenophyllum undulatum (Sw.) Sw. var. fendlerianum (J. W. Sturm) Stolze, Fieldiana, Bot., n.s., 20: 64. 1989. Lectotype (chosen by Lellinger, Me´m. New York Bot. Gard. 38: 12. 1984). Venezuela. [Aragua:] Colonia Tovar, Fendler 35 (isolectotypes BM!, BR–photo BM!, US!), Moritz 450 (isosyntype BM!). Hymenophyllum contortum Bosch, Ned. Kruidk. Arch. 5: 170. 1863. Mecodium contortum (Bosch) Copel., Philipp. J. Sci. 67: 26. 1938. Type. Costa Rica. Aguacata, Hoffman s.n. (B).

Rhizomes creeping, 0.3 mm diam.; rhizome hairs centrally attached, sessile, reddish brown, 0.8–1 mm; fronds 4–10 ⫻ 1.2–2 cm, distant; stipes ca. 1⁄4–1⁄3 the frond length, 0.1–0.2 mm diam., non-alate, a few hairs at bases or glabrescent; blades 2–3-pinnate, glabrous, proximal half of rachises mostly non-alate or essentially so; pinnae many, reduced at blade bases, proximal 2–4 pairs short-stalked; segments 0.8–1 mm wide, entire, crisped, glabrous; sori terminal or lateral on pinnae, not immersed, 1 ⫻ 1 mm, suborbicular, receptacles not exserted.

Distribution. Epiphytic, less commonly terrestrial, in wet forests; 1150–1700 m. Mexico; Guat, CR; Gr Ant; Col, Ven, Sur, Ec, Peru, s Braz, Bol.

Specimens Examined. Chis (Breedlove 33010a, DS). Oax (Mickel 4146, 4173, 5287, NY).

Selected Specimens Examined. Chis (Breedlove 22382, DS, 32441, DS, MICH, NY). Oax (Breckon & Breckon 2418, NY; Conzatti 3535, US; Mexia 9265a, ARIZ, GH, MICH, NY, UC, US; Mickel 1557c, 4830, 5754c, NY). Ver (Mu¨ller 399, 687, both in 1855, NY).

Unverified, Doubtful, or Mistaken Reports. Qro (Rubio 1490, IEB, cited by Dı´az-Barriga & Palacios-Rios, 1992, but not verified, doubtful). Ver (Purpus 4368, E p.p., cited by Palacios-Rios, 1992, but not verified).

Hymenophyllum fendlerianum is distinguished from H. undulatum by being more crisped and having imbricate pinnae. Proctor (1985) suggested they may not be distinct.

342

HYMENOPHYLLUM

8. HYMENOPHYLLUM FRAGILE Hymenophyllum fragile (Hedw.) C. V. Morton, Contr. U.S. Natl. Herb. 29: 172. 1947. Figs. 160C, D. Trichomanes fragile Hedw., Fil. Gen. Sp., t. 18. 1802. Type. “America meridionalis.” (G). Hymenophyllum intercalatum Christ, Bull. Herb. Boissier, se´r. 2, 4: 942. 1904. Type. Costa Rica. Turrialba, Pittier 13259 (P?).

chises narrowly alate, with reddish black hairs 0.8–1 mm on abaxial surfaces; pinnae adnate; segments 1–2 mm wide, glabrous, apices obtuse; margins plane, coarsely serrate, teeth 0.1–0.3 mm long; involucres not immersed, 2.5–3 ⫻ 1.5–2 mm, constricted at bases, margins entire to serrulate, receptacles slightly to longexserted; 2n⫽112 (Jam).

Rhizomes long-creeping, 0.2 mm diam.; rhizome hairs simple, light brown, 0.5 mm long; fronds 3–6 ⫻ 1–2 cm, distant; stipes 0–1 cm long, 0.1–0.2 mm diam., with forked or stellate hairs, alate only at apices (proximal pinnae decurrent); blades pinnate to pinnate-pinnatifid, pinnae bifid or more forked; segments 1– 1.3 mm wide; margins plane to undulate, entire; rachises alate, with 2–3-stellate hairs; sessile stellate hairs on veins, laminae, and especially on blade margins; involucres not or slightly immersed, as long as wide, margins with sessile stellate hairs, receptacles not exserted. Distribution. Epiphytic in dense wet forests; 1850–2700 m. Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Guy, Sur, Ec, Peru, s Braz, Bol. Selected Specimens Examined. Chis (Breedlove 33586, NY; Ghiesbreght 610, NY; Matuda 2909, NY). Gro (Lorea 1998, FCME, ENCB; Reveal et al. 4224, MEXU). Oax (Mickel 3781, 4141, 6759, NY). Ver (Mu¨ller 7, NY; Ventura A. 7582, NY).

Distribution. Epiphytic or epipetric, rarely terrestrial; 2000– 2700 m. Mexico; Guat, Hond, CR, Pan; Gr Ant; Col, Ven, Ec, Peru, Braz, Bol.

Among the taxa with toothed margins, H. tunbrigense differs by being much smaller in size and segment width, with involucres often oriented at right angles to the plane of the blades, indusial margins entire to scarcely serrate, and plants often terrestrial. Hymenophyllum fucoides is part of a complex including H. ectocarpon (q.v.), the latter differing from H. fucoides in having deeply laciniate involucral margins.

Selected Specimens Examined. Chis (Breedlove 21457, 29432, 31622, 32402, 33688, NY; Martı´nez S. 19827, NY; Somers 283, NY). Gro (Lorea 2365, ENCB p.p.). Oax (Mickel 4247, 4252a, NY).

10. HYMENOPHYLLUM HIRSUTUM

Hymenophyllum fragile is distinguished by its small size, pinnatifid pinnae, sessile stellate hairs on the blade surfaces, and forked or stellate hairs on the stipes.

Hymenophyllum hirsutum (L.) Sw., J. Bot. (Schrader) 1800(2): 99. 1801. Figs. 159A, B.

Trichomanes fucoides Sw., Prodr. 136. 1788. Meringium fucoides (Sw.) Copel., Philipp. J. Sci. 67: 45. 1938. Type. Jamaica. Swartz s.n. (S; photo UC).

Trichomanes hirsutum L., Sp. Pl. 2: 1098. 1753. Sphaerocionium hirsutum (L.) C. Presl, Hymenophyllaceae 34. 1843. Type. Based on Filicula digitata Plumier, Traite´ Foug. Ame´r. 73, pl. 50B, 1705, most likely illustrating a plant from the West Indies. Trichomanes ciliatum Sw., Prodr. 136. 1788. Hymenophyllum ciliatum (Sw.) Sw., J. Bot. (Schrader) 1800(2): 100. 1801. Sphaerocionium ciliatum (Sw.) C. Presl, Hymenophyllaceae 34. 1843. Type. Jamaica. Swartz s.n. (S?). Hymenophyllum ciliatum (Sw.) Sw. var. nudipes Kunze, Linnaea 18: 351. 1844. Type. Mexico. Leibold 72 p.p. (LZ, destroyed; isotype B). See Morton (1947) for more synonymy.

Rhizomes creeping, 0.3–0.5 mm diam.; rhizome hairs reddish, 0.8–1 mm long; fronds 8–20 ⫻1.5–5 cm, somewhat distant; stipes usually more than 2 cm long, 0.4–0.7 mm diam., proximal 1⁄3 of stipes not alate, hairs sparse, simple; blades tripinnatifid, ovate to linear-lanceolate, reduced slightly towards base, glabrous; ra-

Rhizomes long-creeping, 0.3–0.5 mm diam.; rhizome hairs simple, 1 mm long; fronds 3–12 ⫻ 1.5–3 cm, pendent or subdistant; stipes 0.2–0.4 mm diam., alate ca. 1⁄2 or more their length, hairs forked or stellate; blades pinnate-pinnatifid, ovate; rachises alate; pinna margins plane to slightly crispate, entire; hairs on

9. HYMENOPHYLLUM FUCOIDES Hymenophyllum fucoides (Sw.) Sw., J. Bot. (Schrader) 1800(2): 99. 1801. Figs. 162E, F.

HYMENOPHYLLUM MAXONII

343

veins stellate, some stalked, some sessile; hairs on margins 2forked, bistellate or basally twice-stellate; involucres nearly round, margins with stalked, stellate hairs, receptacles not exserted; 2n⫽72 (Jam).

Distribution. Mostly epiphytic in wet montane forests; 1200– 2450 m. Mexico; Guat, CR, Pan; Gr & L Ant; Col, Ven, Sur.

Distribution. Epiphytic or terrestrial in wet montane forests; 600–1900 m. Mexico; Guat, Bel, Hond, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol. Selected Specimens Examined. Chis (Breedlove 57768, NY; Martı´nez S. 13504, NY). Gro (Lorea 3035, F, FCME). Oax (Hallberg 1304a, NY; Mickel 1004, ISC, MICH, NY, 5760, 7116, NY). Pue (Orcutt 4004, DS p.p.; Barnes & Land 497a, US). Ver (Nee 24947, XAL).

Among the Mexican hymenophyllums with hairs on only the blade margins and veins, H. hirsutum is distinct by stipes alate at least in distal half.

11. HYMENOPHYLLUM LANATUM Hymenophyllum lanatum Fe´e, Me´m. Foug. 11: 116, t. 31, f. 3. 1866. Figs. 160G, H. Sphaerocionium lanatum (Fe´e) Copel., Philipp. J. Sci. 67: 31. 1938. Type. Guadeloupe. Sainte-Rose, 1861, L’Herminier s.n. (P; isotype US!).

Rhizomes long-creeping, 0.2 mm diam.; rhizome hairs simple, 0.5 mm long; fronds 2–5 ⫻ 1–1.5 cm, subdistant; stipes 1⁄3–1⁄2 the frond length, 0.1–0.2 mm diam., not alate, with hairs sparse, stalked, stellate; blades pinnate-bipinnatifid; segments 1–1.3 mm wide, ovate to lanceolate; rachises alate with stalked, stellate hairs; margins plane, entire; hairs of veins, margins and lamina surfaces stellate with stalks 0.1 mm long and 4–5-rayed; involucres round, margins with stalked, stellate hairs, receptacles not exserted; 2n⫽72 (Jam).

Selected Specimens Examined. Chis (Breedlove 32223, NY). Gro (Lorea 2365, ENCB p.p.). Oax (Hellwig 499, NY; Mickel 4277, 5370, 5743, 5748, 5758, 5948a, 6634, NY).

Of the Mexican species of Hymenophyllum with hairs on the laminar surfaces, H. lanatum is distinguished by its mostly simple, linear pinnae and stellate stipe hairs.

12. HYMENOPHYLLUM MAXONII Hymenophyllum maxonii Christ ex C. V. Morton, Contr. U.S. Natl. Herb. 29: 165. 1947. Type. Guatemala. Alta Verapaz: trail from Senahu´ to Actala´, Maxon & Hay 3326 (US!). Figs. 164E–H. Rhizomes long-creeping, 0.3–0.5 mm diam.; rhizome hairs orange-tan, ca. 1 mm; fronds 4.5–20 ⫻ 2–4.5 cm, distant; stipes 1 ⁄4–1⁄3 the frond length, 0.2–0.5 mm diam., alate 1⁄8–1⁄2 their length distally, with stalked, forked or stellate hairs; blades 2–3pinnatifid, lanceolate; rachises alate; pinna margins entire, plane; hairs on margins 2-forked or stalked, forked or stellate; hairs on veins stellate to forked, stalked or not; involucres semiorbicular, margins with mostly 2-forked hairs.

Distribution. Epiphytic in montane rain forests; 1450–2830 m. Mexico; Guat.

344

HYMENOPHYLLUM

Selected Specimens Examined. Chis (Breedlove 32245, F, NY). Oax (Arreguı´n 1694, ENCB; Ishiki 1666, MEXU). Pue (Herrera & Riba s.n., MEXU; Riba et al. 920A, MEXU). Ver (Beaman 5447, MEXU, NY; Beaman & Alvarez 5773, ARIZ, MEXU; Riba 978, ENCB; Ventura A. 205, IEB).

Hymenophyllum maxonii can be distinguished from H. microcarpum by the former having lanceolate (vs. ovate-deltate) blades and blade margins with stalked, forked or stellate (vs. single or sessile, forked) hairs.

13. HYMENOPHYLLUM MICROCARPUM Hymenophyllum microcarpum Desv., Me´m. Soc. Linn. Paris 6: 333. 1827. Figs. 159E, F. Sphaerocionium microcarpum (Desv.) Copel., Philipp. J. Sci. 67: 34. 1938. Type. Hispaniola. s. coll. s.n. (P!; photo US!). For additional synonymy, see Morton, 1947.

Rhizomes long-creeping, 0.8–1 mm diam.; rhizome hairs 1.5– 2 mm long; fronds 10–30 ⫻ 4–10 cm, distant; stipes 0.5–1 mm diam., alate ca. 1⁄4–1⁄2 the length, with hairs stellate; blades pinnate-pinnatifid or pinnate-bipinnatifid, ovate-deltate; rachises alate with mixed stalked, stellate hairs and simple hairs; margins plane or undulate, entire; segments 0.8–1 mm wide; hairs on veins stalked, stellate and simple, sessile on margins, forked; involucres ovate-elliptic, 1 ⫻ 0.8–1 mm, margins with simple hairs, receptacles not exserted.

Mecodium myriocarpum (Hook.) Copel., Philipp. J. Sci. 67: 25. 1938. Type. Colombia. Hartweg 1530 (K!; isotypes LE!, P!). Hymenophyllum nigrescens Liebm., Mexic. Bregn. 292 (reprint 140). 1849. Type. Mexico. Puebla: Barrancas around Chinautla, Liebmann s.n. [Pl. Mex. 2139, Fl. Mex. 537, 538] (C!). Sphaerocionium nigricans C. Presl ex Klotzsch, Linnaea 18: 536. 1844. Hymenophyllum nigricans (C. Presl ex Klotzsch) Kunze, Bot. Zeitung (Berlin) 5: 244. 1847, hom. illeg., non Colla, 1836. Mecodium nigricans (C. Presl ex Klotzsch) Copel., Philipp. J. Sci. 67: 25. 1938. Syntypes. Venezuela. [Aragua:] Colonia Tovar, Moritz 268, 268b; Peru. Dombey 87 (P!).

Rhizomes creeping, ca. 0.5 mm diam.; rhizome hairs centrally attached, sessile, reddish brown, 1.5–2 mm long; fronds subdistant, 10–25 ⫻ 1.5–3.5 cm; stipes 1⁄8–1⁄5 the frond length, 0.5–0.8 mm diam., alate to bases, glabrous; blades 2–3-pinnatifid, ovate to linear-lanceolate, glabrous; rachises alate; pinnae many, adnate; segments 1–1.3 mm wide, narrowed near forks to 0.5 mm; margins plane, entire, glabrous; involucres round to broadly ovate, 0.8–1 ⫻ 1–1.5 mm, greatly constricted at bases (0.5 mm wide) and appearing stalked, margins entire, receptacles not exserted.

Distribution. Epiphytic in wet montane forests; 1200–3000 m. Mexico; Guat, Hond, Salv, Nic, CR, Pan; Col, Ven, Guy, Sur, Fr Gui, Ec, Peru, Bol. Selected Specimens Examined. Chis (Breedlove 22039, 32056, NY). Gro (Martı´nez S. 1113, MEXU; Rzedowski & McVaugh 181, MICH). Jal (Iltis 2229a, NY). Oax (Mickel 4396, NY). Pue (Herna´ndez Xolocotzi & Sharp X-130, MEXU; Rzedowski 29971, MICH, NY). Ver (Mu¨ller 1490, in 1853, NY; Sharp 45570, MEXU, US).

Distribution. Mostly epiphytic in wet montane forests; 1900– 2200 m. Mexico; Guat, Hond, Salv, Nic, CR, Pan; Gr Ant, Dominica; Col, Ven, Guy, Ec, Peru, s Braz, Bol. Selected Specimens Examined. Chis (Breedlove 32211, DS, 32215, NY; Mu¨nch s.n., DS; Rovirosa 962, PH). Oax (Hallberg 1520a, NY; Mickel 921b, 4289, 5577, 6539, 6758, NY).

Hymenophyllum microcarpum is distinct by its large, deltate blades and stout, distally winged stipes.

14. HYMENOPHYLLUM MYRIOCARPUM Hymenophyllum myriocarpum Hook., Sp. Fil. 1: 106, t. 37D. 1844. Figs. 161C, D, 163M.

The type specimen of Hymenophyllum nigrescens is a very dark, poorly pressed plant with sori wider than long and constricted proximally, appearing as merely a bad specimen of H. myriocarpum. Among the glabrous hymenophyllums, H. myriocarpum is distinct by the involucres being broader than long and the laminae being greatly constricted below the sori making them appear stalked. Hymenophyllum axillare Sw. of the Greater Antilles (type Jamaica) closely resembles H. myriocarpum in the involucres being wider than long and somewhat contricted below. Further study may show the two to be the same. Fronds occasionally appear indeterminate with irregular development of some distal pinnae.

HYMENOPHYLLUM PULCHELLUM

15. HYMENOPHYLLUM PAUCICARPUM Hymenophyllum paucicarpum Jenman, Bull. Bot. Dept. Jamaica 18: 4. 1890. Type. Jamaica. Jenman s.n. (NY!). Figs. 163F–H. Hymenophyllum siliquosum Christ, Bull. Herb. Boissier, se´r. 2, 4: 938. 1904. Mecodium siliquosum (Christ) Copel., Philipp. J. Sci. 67: 26. 1938. Lectotype (chosen by Lellinger, Proc. Biol. Soc. Wash. 89: 716. 1977). Costa Rica. Werckle´ 307 (P! isolectotype CR).

345

Rhizomes creeping, 0.3–0.5 mm diam.; rhizome hairs centrally attached, sessile, reddish brown, 0.8–1 mm long; fronds 3–17 ⫻ 2–6 cm, distant; stipes 1.5–8 cm long, ca. 1⁄4–1⁄3 the frond length, 0.3 mm diam., non-alate; blades 2–3(–4)-pinnatifid, deltateovate, glabrous; rachises narrowly alate, except occasionally proximally; pinnae several to many, adnate to short-stalked at bases; segments 0.8–1 mm wide; margins entire, plane, glabrous; involucres immersed for their basal 1⁄3, ovate, 1–1.5 ⫻ 0.8–1 mm, entire; 2n⫽56 (Braz, Jam, Jap, Fiji, New Hebrides).

Similar to H. polyanthos, but distinct by its clustered sori at blade apices, involucral lips narrowly elliptic (vs. suborbicular), and small frond size (2–4 cm).

Distribution. Epiphytic, rarely epipetric, in dense wet forests; 850–2700 m. Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, s Braz, Bol; Africa, e Asia. Distribution. Epiphytic in montane rain forests, pine-oakLiquidambar forests; 800–1300 m. Mexico; Hond, CR, Pan; Gr Ant.

Selected Specimens Examined. Chis (Breedlove 29869, NY; Ghiesbreght 609, NY). Gro (Rzedowski 18609, NY). Hgo (Ramı´rez & Riba 635, XAL; Rzedowski 28690, NY). Oax (Lorence 3220, NY; Mickel 4873, NY). Pue (Sa´nchez M. 629, US). Ver (Mu¨ller 1799, NY; Purpus 1987, NY).

Specimens Examined. Chis (Breedlove 22385, DS, MEXU, 29908, 36969, NY; Espejo et al. 4353, UAMIZ; Martı´nez S. 14983, MEXU, UAMIZ). Oax (Ishiki 1622, MEXU; Mickel 6706, NY).

This is an extremely widespread and polymorphic species, varying considerably in frond size, shape, degree of rachis and stipe wings, and involucral shape (orbiculate to lanceolate).

The sori are localized at the blade apices and have thick, exserted receptacles; pinna segments are narrower than in other species of subg. Mecodium and are arranged in a more flabellate manner. This seems to be a very distinct species, related to H. brevistipes.

16. HYMENOPHYLLUM POLYANTHOS Hymenophyllum polyanthos (Sw.) Sw., J. Bot. (Schrader) 1800(2): 102. 1801. Figs. 162A, B, H, J. Trichomanes polyanthos Sw., Prodr. 137. 1788. Mecodium polyanthos (Sw.) Copel., Philipp. J. Sci. 67: 19. 1938. Type. Jamaica. Swartz s.n. (S?; isotype BM, photo UC). Hymenophyllum jalapense Schltdl. & Cham., Linnaea 5: 619. 1830. Type. Mexico. [Veracruz:] Near Jalapa, Schiede & Deppe [812] (B!; isotypes BM, Lphoto NY!, LE!). Hymenophyllum millefolium Schltdl. & Cham., Linnaea 5: 620. 1830. Type. Mexico. Veracruz: Jalacingo, Schiede & Deppe [813] (B!; isotype US!). Hymenophyllum botryoides Bosch, Ned. Kruidk. Arch. 5: 160. 1863. Type. Mexico. “Gelcotti 5394” [Galeotti 6394] (K?; isotype US!).

17. HYMENOPHYLLUM PULCHELLUM Hymenophyllum pulchellum Schltdl. & Cham., Linnaea 5: 618. 1830. Figs. 159G, H. Sphaerocionium pulchellum (Schltdl. & Cham.) C. Presl, Hymenophyllaceae 34. 1843. Type. Mexico. Veracruz: Near Jalacingo, Schiede & Deppe [807] (B!; isotypes LE!–3 sheets). For additional synonymy, see Morton (1947: 150).

Rhizomes long-creeping, 0.2–0.3 mm diam.; rhizome hairs simple, sparse, 1 mm long; fronds 6–30 ⫻ 1.5–4 cm, pendent, subdistant; stipes 0.2–0.3 mm diam., non-alate, with hairs sparse to dense, subsessile, stellate or forked; blades pinnate-pinnatifid, linear to oblong, pinnae reduced proximally; rachises not alate; pinna segments 0.8–1 mm wide; margins entire, plane to undulate; hairs on veins abundant, stellate, stalked or subsessile; hairs on margins subsessile, stellate; involucres slightly immersed, round to semicircular, margins with subsessile stellate hairs, receptacles not exserted.

346

HYMENOPHYLLUM

Distribution. Epiphytic or terrestrial or epipetric in wet montane forests; 750–2700 m. Mexico; Guat, Bel, Hond, Salv, Nic, CR; s Braz.

Distribution. Epiphytic (or on rocks), in montane rain forests; 800 m. Mexico; Guat, Bel, Hond, Nic, CR, Pan; PR, L Ant; Ven.

Selected Specimens Examined. Chis (Breedlove 22384, 56646, NY; Dressler 1694, NY). Oax (Hallberg 1517, NY; Mickel 1047, ISC, MEXU, NY, 4831, 5587, 6702, NY). Ver (Mu¨ller 8, in 1858, NY; Purpus 1986, NY).

Specimens Examined. Chis (Breedlove 32954, DS; Mu¨nch 11, DS, US).

Among the species with hairs restricted to blade margins and veins, H. pulchellum is distinct by its non-alate stipes and sessile, stellate hairs on laminar and indusial margins.

18. HYMENOPHYLLUM SIEBERI Hymenophyllum sieberi (C. Presl) Bosch, Ned. Kruidk. Arch. 4: 414. 1859. Figs. 164J–L. Sphaerocionium sieberi C. Presl, Hymenophyllaceae. 58. 1843. Type. Martinique, Kohaut s.n. (Sieber exsicc. 71).

Rhizomes long-creeping, 0.3–0.5 mm diam.; rhizome hairs reddish brown, simple, 1.5–2 mm long; fronds 15–40(–55) ⫻ 2.5–8 cm, distant, pendent; stipes 1⁄10–1⁄5(–1⁄3) the frond length, 1.5–8 cm long, 0.4–0.5 mm diam., conspicuously alate in distal half, with stellate (3-rayed) hairs, hairs uncommonly simple or forked; blades pinnate-pinnatifid, oblanceolate; rachises alate throughout; pinnae 15–30 pairs, close, subimbricate, 2–3(–5) cm long, to 1.8 cm wide at bases, sessile, rather shallowly pinnatifid; primary segments 5–13 pairs, these mostly minutely bifid at their apices, ultimate segments very short, 2 mm long or less, less than 1 mm wide; margins entire; hairs on leaf surfaces and veins few, stellate, 3–5-rayed, subsessile, marginal ones forked or stellate (3-rayed), rather long-stalked; involucres slightly immersed at bases, valves suborbicular to slightly oblong, ca. 1.5 mm long, entire.

The linear blades, pendent fronds, reduced basal pinnae, and winged stipes distinguish this species from H. crassipetiolatum.

19. HYMENOPHYLLUM TEGULARIS Hymenophyllum tegularis (Desv.) Proctor & Lourteig, Bradea 5: 385. 1990. Figs. 160E, F. Davallia tegularis Desv., Me´m. Soc. Linn. Paris 6: 316. 1827. Type. Plumier, Traite´ Foug. Ame´r., pl. 99A. 1705, s. loc. Hymenophyllum elegantulum Bosch, Ned. Kruidk. Arch. 4: 408. 1859. Sphaerocionium elegantulum (Bosch) Copel., Philipp. J. Sci. 67: 32. 1938. Type. Based on Hook., Sp. Fil. 1: 91, t. 33A. 1846, in turn based on Ecuador, Pillzhum, Jameson s.n. (K!; isotype NY!).

Rhizomes long-creeping, 0.1–0.2 mm diam.; rhizome hairs simple, light brown, 1 mm long; fronds pendent, 8–25 ⫻ 2.5–5 cm, distant; stipes ca. 1⁄6 the frond length, ca. 0.2 mm diam., not alate (except for decurrent basal pinnae), hairs dense, simple or forked; blades pinnate-pinnatifid, linear to lanceolate, proximal pinnae greatly reduced; rachises alate only distally, not-alate proximally, densely stellate-hairy; pinnae pinnatifid, segments often forked, 1.8–2 mm wide; margins plane or undulate, entire; sessile stellate hairs on veins, laminae and margins; involucres not or slightly immersed, longer than broad, 2 mm long, margins with stellate hairs, receptacles not exserted.

HYMENOPHYLLUM TUNBRIGENSE

Distribution. Mostly epiphytic in cloud forests; 2000–2800 m. Mexico; Guat, Hond, Salv, Nic, CR, Pan; Cuba, Hisp; Col, Ven, Ec, Peru, Bol. Selected Specimens Examined. Chis (Breedlove 25405, 53592, NY). Gro (Hinton 14306, NY p.p., US). Hgo (Breedlove 59486, NY; Pringle 13255, CAS, F, MICH, MO). Oax (Diggs 2127, ENCB, NY; Mickel 4143b, NY, 4892, ENCB, MEXU, NY, US). Ver (Heitz 419, XAL, Pringle 6079, NY, US).

Among the species with hairs on laminar surfaces as well as on margins and veins, H. tegularis is distinct by its long fronds (8–25 cm) and simple stipe hairs.

347

Distribution. Epiphytic in wet montane forests; 750–2830 m. Mexico; Guat, Hond, CR, Pan; Col, Ven, Sur. Selected Specimens Examined. Chis (Breedlove 30899, 31478, NY). Gro (Lorea 3710, FCME, MEXU). Me´x (Hinton 3268, MEXU, NY). Mor (Lyonnet 2858, MEXU, US). Oax (Mickel 5947, NY; Torres C. 11333, MEXU, NY). Pue (Vera Santos 3677, NY). Ver (Nee et al. 24977, NY; Pringle 8157, NY).

Hymenophyllum trapezoidale is distinguished by blade hairs restricted to margins and veins only, marginal hairs stalked, stipes non-alate, and simple hairs on indusial margins.

21. HYMENOPHYLLUM TUNBRIGENSE 20. HYMENOPHYLLUM TRAPEZOIDALE Hymenophyllum trapezoidale Liebm., Mexic. Bregn. 293 (reprint 141). 1849. Syntypes. Mexico. Puebla: Chinautla, Liebmann s.n. [Pl. Mex. 2151, Fl. Mex. 527, 528] (C!). Lectotype (chosen by Smith, 1981: 135). Liebmann s.n. [Fl. Mex. 528] (C!, frag. US!; isotype GH). Figs. 160A, B. Hymenophyllum apterum Bosch, Ned. Kruidk. Arch. 5: 194. 1863. Lectotype (chosen by Morton, 1947: 167). Mexico. Veracruz: Cuesta Grande de Jalacingo, Schiede [810 p.p.] (L?).

Rhizomes creeping, 0.3 mm diam.; rhizome hairs 1 mm long; fronds pendent, 1.5–8 ⫻ 2–6 cm, distant; stipes 0.3–0.5 mm diam., non-alate, with hairs sparse, simple or occasionally forked; blades pinnate-pinnatifid, linear or linear-lanceolate; rachises alate in distal half, non-alate proximally; pinnae 10–25 pairs, ascending, pinnatisect; segments 0.8–1 mm wide, margins entire; hairs of veins scattered, stalked, stellate; hairs of margins stalked (stalks 0.1 mm), mostly forked or stellate; laminar surfaces glabrous; involucres immersed, margins with some simple hairs, receptacles not exserted.

Hymenophyllum tunbrigense (L.) Sm., Me´m. Acad. Roy. Sci. (Turin) 5: 418. 1793 [1794]. Figs. 162C, D. Trichomanes tunbrigense L., Sp. Pl. 2: 1098. 1753. Type. Uncertain, LINN 1253.4 and 1253.5 are this species. Hymenophyllum fucoides (Sw.) Sw. var. frigidum Liebm., Mexic. Bregn. 291 (reprint 139). 1849. Type. Mexico. Oaxaca: “Cerro de Sempoaltepec,” Liebmann s.n. [Pl. Mex. 2152, Fl. Mex. 554] (C!).

Rhizomes creeping, 0.2–0.3 mm diam.; rhizome hairs 0.3–0.5 mm long, concentrated at base of fronds; fronds arching or erect, 4–12 ⫻ 1.2–2.5 cm; stipes 1⁄3 the frond length, 0.3 mm diam., very narrowly alate distally, glabrous except for occasional short hairs; blades pinnate-bipinnatifid, ovate-lanceolate, glabrous, proximal pinnae not much reduced; rachises barely alate; segments 0.7–1 mm wide, glabrous; margins remotely or distantly serrulate; involucres not immersed, turned nearly at right angles to plane of blades, margins entire to denticulate, 2–2.3 ⫻ 1–1.3 mm, receptacles not exserted; 2n⫽26 (England, Tristan da Cunha).

348

HYPOLEPIS

Distribution. Epiphytic or epipetric (or on soil) in cloud forests or montane rain forests; 1850–2750 m. USA (SC); Mexico; Guat, Hond, Salv, CR, Pan; Jam, Hisp; Ven, Ec, Bol, Chile, n Arg; w Europe, Africa, Austr, NZ. Selected Specimens Examined. Chih (Correll & Gentry 23102, NY). Chis (Breedlove 10019, MICH, 25005, NY). Hgo (Breedlove 59485, CAS). Oax (Hallberg 869, MICH, NY; Mickel 3827, NY). Pue (Riba et al. 338, MEXU). Qro (Zamudio et al. 6605, IEB, 7368, IEB). Ver (Ventura A. 18468, ENCB, XAL).

Hymenophyllum tunbrigense is related to H. fucoides and H. ectocarpon but differs in narrower (0.7–1 mm wide) segments, smaller size, glabrous blades, and subentire indusial segments. There is considerable variation in involucral shape and margin over this species’ range. Involucres are oblong and toothed in Oaxaca and Europe, subentire and nearly round in Chihuahua.

22. HYMENOPHYLLUM UNDULATUM Hymenophyllum undulatum (Sw.) Sw., J. Bot. (Schrader) 1800(2): 101. 1801 [1802]. Figs. 163J–L. Trichomanes undulatum Sw., Prodr. 137. 1788. Mecodium undulatum (Sw.) Copel., Philipp. J. Sci. 67: 26. 1938. Type. Jamaica, presumably Swartz s.n. (S, photo NY; isotypes BM, L–photo UC).

Similar to H. myriocarpum and H. fendlerianum in the irregular development of distal pinnae, giving a somewhat indeterminate appearance to the plants, but distinct from the former in the sori not constricted proximally and stipes and proximal part of rachises non-alate (vs. rachises and most of stipe length alate), and from the latter in the segments plane or slightly undulate (vs. very strongly undulate) and sori broader than long (vs. suborbicular); 2n⫽56 (Jam.).

Distribution. Epiphytic in montane rain forests; 1150–3000 m. Mexico; Guat, Hond, Nic, CR, Pan; Cuba, Jam, Guadeloupe; Col, Ven, Sur, Ec, Peru, se Braz, Bol. Selected Specimens Examined. Chis (Breedlove 10016, MICH, US, 22058, MICH, 22383, DS p.p., 22756, MICH, 22939, 31806, NY). Oax (Mickel 1041, ENCB, ISC, NY, 1107, ENCB, ISC, MEXU p.p., NY, 5742, 5754a, NY). Unverified, Doubtful, or Mistaken Reports. Ver (Barnes et al. 72, F, cited by Pacheco, 1994, but not verified).

The closest relatives of H. undulatum are H. myriocarpum and H. fendlerianum. Hymenophyllum myriocarpum in Chiapas occurs at higher elevations than H. undulatum; Hymenophyllum fendlerianum, which has more crispate fronds, has been collected with H. undulatum (Breedlove 22383, DS). There is some confusion of H. fendlerianum and H. undulatum in the Antilles. A photograph of the holotype of H. undulatum shows the lamina plane rather than strongly crisped. It remains to be seen whether H. undulatum is indeed a distinct taxon, a form of H. myriocarpum, or perhaps the same as H. fendlerianum.

57. HYPOLEPIS Hypolepis Bernh., Neues J. Bot. 1(2): 34. 1806 [1805]. Type: Hypolepis tenuifolia (G. Forst.) Bernh. [⬅ Lonchitis tenuifolia G. Forst.]. Terrestrial, thicket-forming; rhizomes long-creeping, much branched from the leaf bases, hairy; fronds mostly large, often scandent, 2–4-pinnate, generally hairy, rarely glabrous; stipes stramineous, atropurpureous or castaneous, hairy to glabrous, spiny or not; laminae herbaceous to coriaceous, glabrous to hairy; veins free; sori marginal (to rarely medial), round to oblong, generally protected by differentiated stramineous to tan indusial flaps or recurved teeth, without paraphyses, inner indusia absent; spores bilateral, tuberculate to spinulose, perispore absent; x⫽29, 52. Hypolepis is a pantropical genus of about 45 species, with 15– 20 in America. It is most closely allied to Paesia (x⫽26) and Pteridium (x⫽52). Counts of n⫽29 in H. nigrescens perhaps reflect an ancient link to a dysploid series from 26 to 29 and the

HYPOLEPIS BLEPHAROCHLAENA

doubling of 26 to make the more common n⫽52. The species with n⫽29 form a natural group (emarginate segments, recurved spines) and may deserve taxonomic segregation. Hypolepis is distinguished by the surficial, long-creeping, hairy rhizome, discrete sori, and bilateral spores.

349 References

Brownsey, P. J. 1983. Polyploidy and aneuploidy in Hypolepis, and the evolution of the Dennstaedtiales. Amer. Fern J. 73: 97–108; Mickel, J. T. 1973. The phylogenetic position of and the classification within the Dennstaedtiaceae. J. Linn. Soc., Bot. 67: 135–144.

Key to the Mexican Species of Hypolepis 1. Frond axes glabrous or hairy, lacking spines. 2. Frond axes reddish purple; indusia vestigial, not protecting the sori, marginal, not protected by recurved teeth. ................ 6. H. pulcherrima. 2. Frond axes stramineous to castaneous; sori indusiate. 3. Indusia wide (0.5–0.8 mm), with subentire to erose margins. ..................................................................................................... 4. H. muenchii. 3. Indusia narrow (0.1–0.3 mm), with ciliate margins. 4. Adaxial blade surfaces with hairs along veins; abaxial surfaces with numerous glandular hairs; axes with dense hairs. ........................................................................................................................................................................................ 9. H. thysanochlaena. 4. Adaxial blade surfaces glabrous with hairs mostly along costae; abaxial surfaces with scattered hairs; axes with rare to scattered hairs. ........................................................................................................................................................................ 10. H. trichochlaena. 1. Frond axes armed with spines. 5. Spines mostly recurved, often dark-tipped; teeth emarginate; laminae chartaceous, often drying dark (nearly black). ............ 5. H. nigrescens. 5. Spines straight, stramineous; teeth not emarginate; laminae firm to subcoriaceous, drying greenish to blackish. 6. Indusia blackish; fronds drying greenish brown to blackish; rachises very sparsely spiny. .............................................. 2. H. melanochlaena. 6. Indusia stramineous to tan; fronds drying greenish; rachises moderately spiny. 7. Adaxial blade surfaces with glandular hairs scattered over surfaces and dense on costae; indusia short (0.1–0.3 mm), narrow (0.1–0.3 mm), with ciliate margins. 8. Erect acicular hairs absent. ............................................................................................................................................... 3. H. microchlaena. 8. Erect acicular hairs present on blade surfaces and indusia among the jointed glandular hairs. ................................... 8. H. stuebelii. 7. Adaxial blade surfaces with rare to scattered jointed hairs, mostly along costae; indusia long (0.8–1 mm), narrow (0.1– 0.3 mm) to wide (0.5–0.8 mm), with erose to long-ciliate margins. 9. Indusia with erose to ciliate margins; laminae dull beneath; segments perpendicular to the costae; (1780–)2150–2450 m. ..................................................................................................................................................................................... 1. H. blepharochlaena. 9. Indusia with irregularly toothed margins; laminae lustrous beneath; segments slightly ascending; 450–1800 m. ........ 7. H. repens.

1. HYPOLEPIS BLEPHAROCHLAENA Hypolepis blepharochlaena Mickel & Beitel, Mem. New York Bot. Gard. 46: 219. 1988. Type. Mexico. Oaxaca: Distrito Teotitla´n, 26–29 km NE of Teotitla´n del Camino, vicinity of pass at Puerto Soledad, Mickel 4151 (NY!; isotype US!). Figs. 165Q–T. Rhizomes long-creeping, much branched, 5–8 mm diam., lustrous dark brown, with brown ctenitoid hairs, leaving persistent bases; fronds 2–3 m long, distant; stipes stramineous to reddish brown, nearly black at bases, muricate, rachis and pinna axes yellow-brown to castaneous, spiny with stramineous, straight spines to scabrous with enlarged hairs and persistent hair bases; blades tripinnate-pinnatifid, deltate to ovate-deltate, subcoriaceous; segments pinnatifid, the lobes perpendicular to costae, with jointed hairs scattered in grooves and sparse on veins adaxially; abaxial surfaces dull, glabrous except for sparse hairs on costae, 0.5–1 mm long; sori marginal, with marginal flaps oblong and deeply ciliate, 1–1.3 ⫻ 0.8–1 mm.

Distribution. Clearings and forest margins at higher elevations; (1780–)2150–2450 m. Mexico; Guat, Hond, Salv, CR. Selected Specimens Examined. Chis (Breedlove 32362, NY; Purpus 6710, UC, US). Hgo (Gimate Leyva s.n., 30 Jul 1967, NY; Martı´nez S. 28463, MEXU). Oax (Mickel 4131, NY; Smith et al. 4138, US). Pue (Riba et al. 291, NY; Rzedowski 23407, NY). Ver (Purpus 6057, NY; Ventura A. 5017, NY).

Besides the dull abaxial surfaces and the wide, erose to ciliate indusia, which distinguish it from Hypolepis repens, this species generally occurs at higher elevations, (1780–)2150–2450 m, whereas H. repens occurs between 450 and 1250(–1550) m. The non-glandular, almost glabrous adaxial blade surfaces and long

350

HYPOLEPIS

(1–1.3 mm), wide (0.8–1 mm) indusia separate H. blepharochlaena from H. microchlaena. Specimens of H. blepharochlaena have often been mistakenly named H. mexicana (here regarded as synonymous with H. repens), but the type material of the latter has indusia not noticeably broad nor ciliate and was collected at 4000–5000 ft (1250–1550 m).

2. HYPOLEPIS MELANOCHLAENA Hypolepis melanochlaena A. R. Sm., Proc. Calif. Acad. Sci., ser. 4, 40: 221, f. 7D–E. 1975. Type. Mexico. Chiapas: Mu¨nch 35 (DS!, probable isotypes DS!, US!). Figs. 166A–C.

Rhizomes long-creeping, 3–7 mm diam., clothed with brown hairs; fronds large, distant; stipes stramineous to light brown, castaneous at bases with short, stramineous spines and many spreading hairs 1–2 mm long (the spines onto tertiary axes); blades long-deltate, tripinnate-pinnatifid, basal pinnae 40 ⫻ 20 cm; segments pinnatifid, lobes oblong, rounded; laminae thin, herbaceous; adaxial laminar surfaces with numerous glandular hairs, dense on veins; abaxial surfaces and veins with abundant short glandular hairs; veins running to margins, with slightly clavate tips; sori submarginal, indusia short and narrow, 0.3 ⫻ 0.1–0.3 mm, with ciliate margins.

Complete fronds not seen, probably subscandent; rachises tan to brownish, glabrous, sparingly aculeate, spines patent, straight, stramineous; blades 3-pinnate-pinnatifid, dissected as in H. repens; pinnae alternate, equilateral, largest more than 35 ⫻ 10 cm; penultimate axes not alate, straight, without adaxial wings, with hairs along edges of grooves; costae and costules below completely glabrous; leaf tissue firm to subcoriaceous, drying greenish brown to blackish, glabrous abaxially; sori 1–2 pairs per ultimate segment; indusia flap-like, entire, drying blackish.

Distribution. Wet forests; 1175–1400 m. Known only from the type collection. The abundant glandular hairs, especially on the adaxial laminar surfaces, and short (0.1–0.3 mm), narrow (0.1–0.3 mm), ciliate indusia distinguish this species from other species with spiny axes (e.g., H. repens).

4. HYPOLEPIS MUENCHII

Distribution. Known in Mexico only from the type, without exact locality or habitat data. Smith (1985) reported it from western Venezuela, which is a remarkably disjunct distribution. Hypolepis melanochlaena seems to be closely related to H. repens, but is distinct by the dark color and glabrous blade surfaces. A specimen from Veracruz (Go´mez-Pompa et al. 5460, XAL) has dark blade color and is virtually glabrous, but has minute catenate hairs (to 0.2 mm) at the base of the penultimate segments.

3. HYPOLEPIS MICROCHLAENA Hypolepis microchlaena Mickel & Beitel, Mem. New York Bot. Gard. 46: 220. 1988. Type. Mexico. Oaxaca: Distrito Ixtla´n, 79 km NW of Ixtla´n on Rte 175, 1 km S of Campamento Vista Hermosa, ca. 1 hour hike on trail toward Tarabundı´, Mickel 5657 (NY!; isotypes ENCB!, MEXU!, UC!). Figs. 166M, N.

Hypolepis muenchii (Christ) Mickel, comb. nov. Figs. 166K, L. Dennstaedtia muenchii Christ, Bull. Herb. Boissier, se´r. 2. 5: 732. 1905 [“munchii”]. Type. Mexico. Chiapas: San Pablo, Mu¨nch 137 (P?, not found; isotypes DS, US!, frag. NY!). R. M. Tryon (Rhodora 63: 71. 1961) suggested that Mu¨nch 76 (DS, US) is probably also incorrectly numbered type material; the locality and elevation are the same. Hypolepis eurychlaena Mickel & Beitel, Mem. New York Bot. Gard. 46: 220. 1988. Type. Mexico. Oaxaca: Ixtla´n, 5 km W of Ixtla´n de Jua´rez on Rte 175, Mickel 4632 (NY!; isotypes MEXU!, UC!, US!).

Similar to H. thysanochlaena and H. trichochlaena in the nonspinose frond axes, but distinct in its indusia broader (0.5–0.8 vs. 0.1–0.3 mm) and subentire to erose (vs. ciliate).

HYPOLEPIS PULCHERRIMA

351

Distribution. Roadsides and other disturbed open areas in wet forest, pine-oak woods with subtropical hardwoods; 1050–2700 m. Mexico.

Distribution. Forest margins or in light woods in wet montane forests on Atlantic slope; 250–2200 m. Mexico; Guat, Hond, Salv, CR, Pan; Gr Ant; Col, Ven, Ec, Peru, Braz, Bol.

Selected Specimens Examined. Chis (only the type). Oax (Mickel 7231, NY).

Selected Specimens Examined. Chis (Breedlove 21374, DS, 33616, NY; Purpus 6710, PH). Gro (Lorea 3174, FCME). Oax (Hellwig 324, NY; Mickel 1029, 5594, NY; Rzedowski 32984, NY). Ver (Riba 982, MEXU, UAMIZ; Ventura A. 13874, XAL).

Among the unarmed species of Hypolepis, H. muenchii is distinct in its 0.5–0.8 mm wide indusia with subentire to erose margins (vs. 0.1–0.3 mm wide with ciliate margins).

5. HYPOLEPIS NIGRESCENS Hypolepis nigrescens Hook., Sp. Fil. 2: 66, t. 90C. 1852. Lectotype (chosen by Proctor, 1985: 179). Jamaica. Wilson s.n. (K; isolectotypes BM, P!). Figs. 165E–G. Dennstaedtia rubicaulis Christ, Bull. Herb. Boissier, se´r. 2, 5: 258. 1905. Type. Costa Rica. Navarro, Werckle´ s.n. (P!; isotype US, frag NY!).

Rhizomes long-creeping, branched, dark brown, with sparse ctenitoid hairs; fronds to 6 m long; stipes reddish to yellowish brown, with spines numerous, straight to often slightly recurved, usually dark-tipped, with hairs often stiff and dark-tipped, also with minute, appressed, multicellular ctenitoid hairs; blades narrowly oblong-lanceolate, usually 4-pinnate-pinnatifid; segments pinnatifid, emarginate at vein tips; laminae usually drying blackish green, chartaceous; hairs 0.5 mm long, sparse on costae and major veins abaxially, glabrous adaxially, costae broadly grooved and naked; sori round, indusial flaps hardly more than a tooth 0.8–1 ⫻ 0.1–0.3 mm, subentire to slightly lacerate. 2n⫽58 (Jam, Oax).

Of the New World species, this is the most unusual and differs from the others in its chromosome number (n⫽29 vs. 52), the recurved dark spines, dark-drying laminae, and emarginate teeth. It is the most scandent with long fronds that often unroll slowly with the distal pair of pinnae entirely expanded before the next pair is produced. Plants in Hispaniola are often placed in a different species, Hypolepis hispaniolica Maxon, but that is probably only varietally distinct at best. Hypolepis nigrescens is most closely related to Hypolepis brooksiae Alderw. of Southeast Asia, the type of which Hooker included as a syntype of H. nigrescens..

6. HYPOLEPIS PULCHERRIMA Hypolepis pulcherrima Underw. & Maxon, Proc. Biol. Soc. Wash. 43: 84. 1930. Lectotype (chosen by Lellinger, Proc. Biol. Soc. Wash. 89: 716. 1977). Jamaica. Blue Mt. Peak, Maxon 9912 (US!). Figs. 166D–F. Rhizomes long-creeping, 2–4 mm diam., with brown hairs; fronds 0.7–1 m, distant; stipes ca. 1⁄2 the frond length, castaneous, glabrescent, slightly bumpy from fallen hair bases; blades deltate, tripinnate to tripinnate-pinnatifid, ca. 35–40 cm wide; pinnae narrowly deltate, equilateral, short-stalked ca. 1 cm; segments coarsely toothed to pinnatifid, teeth/lobes ascending, acute; laminae dull abaxially, membranous; hairs catenate, most common on major axes adaxially, sparse on veins, scattered or reduced (0.1–0.3 mm) abaxially on costules and major veins; veins ending short of margins, darker than laminae; indusia vestigial, ciliate, not protecting the sori.

352

HYPOLEPIS

well developed, subentire or irregularly toothed scarious flaps, oblong, 0.8–1 ⫻ 0.1–0.3 mm, 1 per lobe; 2n⫽104 (CR), 208 (Fla).

Distribution. Lacking habitat data; oak woods; 2100–2900 m in Costa Rica and Panama (Moran in Davidse et al., 1995). Mexico; CR, Pan; Jam, Hisp. Specimens Examined. Chis (Mu¨nch 111, DS, US).

Ghiesbreght 366 (BM, K) is possibly this species, but the specimens are too incomplete to be certain. These specimens completely lack flexuous, catenate hairs on the axes below, but otherwise agree with H. pulcherrima in the nearly naked sori and dark axes (though castaneous rather than atropurpureous). Chiapas material lacks acicular hairs and is generally less hairy than specimens from Jamaica. Antillean material has atropurpureous axes, is densely pubescent with catenate hairs and a few acicular hairs intermixed, and the indusia are poorly differentiated and ciliate with catenate, glandular hairs. Specimens from Costa Rica and Panama are similar to Antillean material.

7. HYPOLEPIS REPENS Hypolepis repens (L.) C. Presl, Suppl. Tent. Pterid. 162. 1836. Figs. 165N–P. Lonchitis repens L., Sp. Pl. 2: 1078. 1753. Cheilanthes repens (L.) Kaulf., Enum. Filic. 215. 1824. Lectotype (chosen by Proctor, Fl. L. Antill. 2: 130. 1977). Plumier, Traite´ Foug. Ame´r., pl. 12, 1705, based on a plant from Martinique (“le mont Noe¨l, & montant le Morne la Calebasse”). Hypolepis mexicana Liebm., Mexic. Bregn. 260 (reprint 108). 1849. Lectotype (chosen by Smith, 1981: 137). Mexico. Oaxaca: “Hacienda de Sta. Gertrudes ved Talea og Mina de Dolores,” Liebmann [Pl. Mex. 2751, Fl. Mex. 803] (C!; isolectotypes BM, US!–frag. NY!).

Rhizomes long-creeping, 5–10 mm diam., with brown hairs; fronds 2–4 m long, distant; stipes ca. 1⁄2 the frond length, dark at bases, stramineous to brown adaxially, with short, straight, stramineous spines and many spreading hairs 1–2 mm long (with the spines onto the tertiary axes); blades deltate to long-deltate, 3–4-pinnate-pinnatifid, to 1.5 m wide; pinnae deltate, equilateral, stalked; segments pinnatifid, lobes oblong, rounded, slightly ascending; laminae lustrous beneath, firm to coriaceous, jointed hairs adaxially sparse on axes and major veins, jointed hairs abaxially common on laminae; veins ending at blade margins, with slender tips, lighter than the laminae; sori submarginal, indusia

Distribution. Margins of wet forests, roadsides and clearings at mid-elevation on Atlantic slope; 450–1800 m. USA (Fla); Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Cuba, PR, L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol, Arg. Selected Specimens Examined. Chis (Ghiesbreght 230, NY; Purpus 7228, NY). Gro (Martı´nez S. 4234, ARIZ, XAL). Hgo (Herrera & Riba 2228, MEXU; Sa´nchez M. 648, MEX, US). Oax (Mickel 4743, 7241, NY). Pue (Rzedowski 32435, CAS, IEB, MEXU). Ver (Copeland herb. 74, NY; Ventura A. 3674, ENCB, MEXU, XAL).

This is the most common and widespread species of the genus in America. For comparison with taxa having spiny axes, see discussion under H. blepharochlaena and H. microchlaena.

8. HYPOLEPIS STUEBELII Hypolepis stuebelii Hieron., Hedwigia 48: 230, t. 10, f. 8. 1909. Type. Ecuador. Manabı´: prope San Florencio in itinere ab urbe Quito, Stu¨bel 796 (B). Figs. 166G, H, J. Rhizomes long-creeping, 3–5 mm diam., with brown hairs; fronds 1(–2) m, distant; stipes ca. 1⁄2 the frond length, dark at bases, orange-brown distally, with abundant short hairs 0.1–0.2 mm and scattered spines on primary and secondary axes; blades deltate, 3-pinnate, ca. 1.5 mm wide; pinnae deltate, slightly inequilateral, stalked; segments pinnatifid, lobes oblong, rounded; laminae dull abaxially, firm to coriaceous; hairs throughout both catenate, gland-tipped, and hyaline-acicular, adaxially sparse on veins, abaxially abundant; veins ending at the margins, obscure or darker than laminae; indusia well developed, oblong, margins with acicular hairs ca. 1 ⫻ 0.5 mm, 1 per lobe.

HYPOLEPIS TRICHOCHLAENA

Distribution. Pine-oak forests; 1150 m. Mexico; CR, Pan; Jam, Hisp; Col, Ven, Ec, Peru. Specimen Examined. Gro (Vela´squez 281, FCME).

This species is distinct by its spiny axes and a mixture of acicular and catenate blade hairs.

9. HYPOLEPIS THYSANOCHLAENA Hypolepis thysanochlaena Mickel & Beitel, Mem. New York Bot. Gard. 46: 221. 1988. Type. Mexico. Oaxaca: Distrito Ixtla´n, trail from San Pedro Nolasco N to Llano Verde, N slope past the Llano Verde and Las Cruces, Mickel 5360 (NY!; isotypes ENCB!, UC!). Figs. 165H, J–M. Rhizomes long-creeping, branched, ca 2.5 mm diam., with brown, lustrous, ctenitoid or mostly bristle-like hairs; fronds 60– 100 cm long; stipes and axes yellow-brown to castaneous, unarmed, densely clothed with ctenitoid (and some bristle-like) hairs 1–2 mm long; blades deltate to ovate-deltate to lanceolate, 3–4-pinnate-pinnatifid; rachises somewhat flexuous to straight; costae adaxially broadly grooved with abundant ctenitoid hairs on margins, glandular hairs numerous abaxially on costae, veins, and surface; veins ending short of margins; sori round, indusial flaps thin, narrow, 0.5 ⫻ 0.1–0.3 mm, with ciliate margins.

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Distribution. Roadsides and other disturbed open areas in wet forests, pine-oak woods with subtropical hardwoods; 1050– 2700 m. Mexico. Selected Specimens Examined. Chis (Breedlove 22056, DS, MEXU, 31732, DS, NY; Gonza´lez Espinosa et al. 1677, MEXU). Oax (Hellwig 414, NY; King 2123, MICH; Mickel 1061, 1150, 4679, 5360, NY).

The New World taxa with non-spiny axes are in need of revision. This species and H. muenchii differ in blade hairiness, indusial width, and indusial margin from well known species of South America, such as H. bogotensis H. Karst. (with abaxial surfaces felted, hairy) and H. viscosa (H. Karst.) Mett. (with abundant adaxial glandular hairs and narrow (0.1–0.3 mm wide), ciliate indusia with glandular hairs on the surfaces). Collections from Chiapas cited as H. bogotensis H. Karst. by Smith (1981) belong to this species. One collection (Mickel 1150B, Oaxaca, NY) with malformed spores, resembling this species in its glandular hairy lamina, seems to be a hybrid with H. muenchii, judging from the strongly flexuous stipes and erose indusial margins. The indusia are of intermediate width (0.3–0.5 mm wide). Breedlove 31732 (DS, NY) from Chiapas also has malformed spores and a chromosome number of 2n⫽ca. 50II ⫹ 54I at meiosis.

10. HYPOLEPIS TRICHOCHLAENA Hypolepis trichochlaena Mickel & Beitel, Mem. New York Bot. Gard. 46: 222. 1988. Type. Mexico. Oaxaca: Distrito Ixtla´n, 65–68 km N of Ixtla´n, 41–44 km S of Valle Nacional, Mickel 4322 (NY!; isotypes ENCB!, NY!, US!). Figs. 165A–D. Similar to H. thysanochlaena except: abaxial surfaces with only scattered hairs; adaxial surfaces with hairs only along pinna axes; indusia narrow, 0.1–0.3 mm wide, with ciliate margins.

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Distribution. Terrestrial in wet woods; 1600–2700 m. Mexico. Specimens examined. Mich (Rzedowski 42649, ENCB, IEB, MEXU, XAL). Oax (Hallberg 825, MICH, frag. US; Mickel 4311, 5282, 5319, 6554, 6590, NY).

The narrow, ciliate indusia and non-glandular, almost glabrous axes and adaxial blade surfaces distinguish this species. One collection (Mickel 4837, NY, from Oaxaca, 825–1225 m) with malformed spores, resembling H. trichochlaena in having few hairs and narrow, ciliate indusia, seems to be a hybrid with H. thysanochlaena, as evidenced by narrower indusia and a scattering of glandular hairs.

58. I S O E¨ T E S by W. Carl Taylor and R. James Hickey

Isoe¨tes L., Sp. Pl. 2: 1100. 1753. Type: Isoe¨tes lacustris L. Aquatic, amphibious, or seasonally wet-terrestrial, grass-like, tufted; rootstocks buried, subglobose, usually 1–2 cm in diam., 2to 3-lobed, with sclerified scales present or absent; roots arising along grooves between rootstock lobes, simple or dichotomously branched, in transection containing an eccentric vascular strand and surrounding lacuna; leaves linear, mostly 5–50 cm long, stiff to pliant, erect to reclining or recurved, mostly 1–1.5 mm wide at mid-length, bright green to olive green, attenuate to apices, in transection terete to triangular and containing a central vascular strand surrounded by 4 longitudinal and transversely septate lacunae; ligules triangular, membranous, inserted above sporangia; velum partly or entirely covering adaxial wall of each sporangium; sporangia ovoid, solitary, embedded in leaf bases; spores of two kinds (plants heterosporous); megaspores white to dark gray, tetrahedral-globose, mostly 250–750 μm in diam., tens to hundreds per megasporangium; microspores light gray or brown in mass, ovoid, monolete, mostly 25–50 μm long, thousands per microsporangium; megagametophytes multicellular, white, endosporic, exposed when megaspores open along trilete ridges, with archegonia located by quartets of brownish neck cells; microgametophytes 9-celled, endosporic, each with single antheridium releasing 4 multiflagellate spermatozoids; x⫽11. Isoe¨tes is a worldwide genus of more than 200 species. Species

are difficult to identify because their morphology yields few characters that easily distinguish species. Spore size, ornamentation, and color, velum cover, and sporangial wall coloration provide character states that will usually distinguish species. A microscope fitted with an ocular micrometer has been used to measure spore size. At least 10 spores were measured to determine their average diameter. Megaspores were measured dry on a glass slide. Microspores were hydrated and measured on a slide in a drop of water under a cover slip. A 10⫻ to 20⫻ hand lens or dissecting microscope is needed to resolve megaspore texture. Normally megaspores are globose and boldly marked with an equatorial ridge encircling the spore and three ridges converging at the spore’s proximal pole. Between these ridges spores may be tuberculate, pustulate, rugate, cristate, verrucate, echinate, or smooth. The girdle, a zone along the distal side of the equatorial ridge, may be textured differently than the rest of the spore. Plants collected at the end of their growing season with mature spores still within sporangia are easiest to identify. Plants collected early in their growing season with small, fragile, immature megaspores are difficult to identify. Mature spores from the previous growing season may be found in decaying leaf bases or in soil around the rootstocks. The velum is a thin flap of tissue that entirely or partly covers the adaxial wall of each sporangium. Isoe¨tes orcuttii has a velum that completely covers the adaxial wall of the sporangium. The velum of the other Mexican species covers less than 50% of the adaxial wall of each sporangium. Sporangial wall coloration appears as spots or streaks of brown colored cells on the semi-transparent sporangial wall. The sporangial walls of Isoe¨tes howellii and I. montezumae usually have brown spots or streaks. The sporangial wall cells of other Mexican species are clear, without brown spots or streaks. Chromosome counts and the production of malformed spores in interspecific hybrids indicate that Isoe¨tes species have evolved in two ways, either by ecological isolation and genetic divergence without increase in chromosome number, or by interspecific hybridization and chromosome doubling. Recognition of basic diploid species with uniform spores and chromosome counts of 2n⫽22, sterile interspecific hybrids with irregular spores, and allopolyploids with uniform spores and chromosome counts that are multiples of 22, helps in distinguishing Isoe¨tes species. Many Mexican collections of Isoe¨tes show some irregularity in spore size and shape, indicating they could be interspecific hybrids. Interspecific hybrids are recognized by the production of irregular spores that vary in size, shape, and texture. They are usually found where two species occur together. In addition, basic diploid (2n⫽22) and tetraploid (2n⫽44) chromosome counts have been reported for Mexican species. Basic diploid species generally have a smaller average spore size than tetraploid species. References Pfeiffer, N. E. 1922. Monograph of the Isoe¨taceae. Ann. Missouri Bot. Gard. 9: 79-232; Taylor, W. C., N. T. Luebke, D. M. Britton, R. J. Hickey & D. F. Brunton. 1993. Isoe¨taceae. In: FNA Ed. Comm. 2: 64–75.

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Key to the Mexican Species of Isoe¨tes 1. Megaspores averaging less than 375 μm diam. 2. Velum covering 100% of each sporangium; leaves less than 10 cm long; microspores brown in mass. ............................................ 4. I. orcuttii. 2. Velum covering less than 25% of each sporangium; leaves more than 10 cm long; microspores light gray in mass. .................. 2. I. mexicana. 1. Megaspores averaging more than 375 μm diam. 3. Sporangial walls usually brown-spotted or streaked to completely brown. ........................................................................................... 4. I. orcuttii. 4. Megaspores rugate; microspores brown in mass, averaging 25–35 μm long. .................................................................................... 1. I. howellii. 4. Megaspores smooth to tuberculate-verrucate; microspores light gray in mass, averaging 35–45 μm long. .......................... 3. I. montezumae. 3. Sporangial walls unspotted, unstreaked, clear. 5. Megaspores pustulate-tuberculate, averaging 400–475 μm diam.; leaves triangular in mid-length transection. ......................... 5. I. pallida. 5. Megaspores echinate-cristate, averaging 475–650 μm diam.; leaves half-terete in mid-length transection. ................................ 6. I. pringlei.

1. ISOE¨TES HOWELLII

2. ISOE¨TES MEXICANA

Isoe¨tes howellii Engelm., Trans. Acad. Sci. St. Louis 4: 385. 1882. Type. U.S.A. Oregon: near The Dalles, June & August 1880, Howell s.n. (MO!; isotypes NY!, US). Figs. 168A, H.

Isoe¨tes mexicana Underw., Bot. Gaz. 13: 93. 1888. Type. Mexico. Chihuahua: base of the Sierra Madre, Pringle 1447 (NY!; isotypes BM, G, M, MO!, NY!, PH, UC, US, UPS, VT). Figs. 167A-C, 168B.

Rootstock bilobed; leaves ca. 20, ca. 25 cm long, pale to sclerified lustrous black toward bases, half-terete in mid-length transection; velum covering less than 50% of each sporangium; sporangial walls brown-streaked to completely brown; megaspores white, average 375–450 μm in diam., rugate with girdle smooth; microspores brown in mass, average 25–35 μm long, echinate; 2n⫽22 (USA).

Rootstocks bilobed; leaves ca. 20, ca. 25 cm long, pale toward bases, half-terete in mid-length transection; velum covering less than 25% of each sporangium; sporangial walls unspotted; megaspores white, average 275–375 μm in diam., smooth or nearly so with girdle smooth; microspores light gray in mass, average 25– 35 μm long, echinate-tuberculate; 2n⫽22 (Dgo, Jal).

Distribution. Temporary pools and streams in seasonally wet, open woodlands; 1000–1500 m. November–January. Canada, USA; Mexico.

Distribution. Ponds, streams, grassy meadows in seasonally wet, oak-pine woodlands; 1750–2500 m. September–November. Mexico.

Specimens Examined. BCS (Brandegee s.n., 24 Jan 1890, NY, s.n., Nov. 1902, UC; Breedlove & Axelrod 43344, CAS; Carter et al. 2358, UC).

Selected Specimens Examined. Chih (Pringle 1713, G, MO, UC). Dgo (Breedlove 18848, NY; McVaugh 21725, MICH, NY). Gto (McVaugh 24072, MEXU, MICH, NY). Hgo (Pringle 8796, BM, C, E, G, M, MO, MEXU, NY, PH, UC, US, VT; Rzedowski 36110, ENCB). Jal McVaugh 17600, CAS). Mich (Gonza´lez 260, ENCB, MEXU, NY). Qro (Rzedowski & McVaugh 483, MEXU, MICH, NY).

Isoe¨tes howellii reaches the southern extent of its range in the Sierra La Laguna region of southern Baja California. In this region I. howellii has megaspores with an obscure rugate texture of low, meandering ridges and microspores averaging less than 35 μm in length. Isoe¨tes montezumae, which generally resembles I. howellii, has tuberculate megaspores, and microspores averaging more than 35 μm in length.

Isoe¨tes mexicana and I. montezumae are similar in general appearance. However, most collections of I. mexicana can by distinguished by their smooth, shiny, porcelain-like megaspores averaging less than 375 μm in diameter, each contained by an unspotted sporangial wall. Occasionally, megaspores of I. mexicana are obscurely tuberculate. In contrast, I. montezumae has roughened, usually lusterless, obscurely to boldly tuberculate

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megaspores that average more than 375 μm in diameter, each contained by a spotted sporangial wall. Sometimes, megaspores of I. montezumae lack apparent tubercles, but spore diameter will usually distinguish these smooth-spored individuals from I. mexicana. In addition, according to their original descriptions, microspores of I. mexicana average less than 35 μm in length, whereas those of I. montezumae will average more than 35 μm in length. When making these distinctions, it is important to use mature spores. Study of herbarium specimens indicates that Isoe¨tes mexicana and I. montezumae could be treated as a single variable species or two intergrading species. There are collections of these two taxa that are easily assignable to either I. mexicana or I. montezumae, but there are also many collections of specimens with intergrading characters that are difficult to place. However, within the overlapping ranges of I. mexicana and I. montezumae there are specimens bearing irregular spores that vary in size and shape (Fig. 168 G). Production of irregular spores is a characteristic of interspecific hybrids. Collections of what appear to be interspecific hybrids within the overlapping ranges of these taxa are evidence that I. mexicana and I. montezumae are genetically divergent to an extent that they can be treated as two species.

3. ISOE¨TES MONTEZUMAE Isoe¨tes montezumae A. A. Eaton, Fern Bull. 5: 25. 1897. Type. Mexico. Me´xico: Plains near Flor de Marı´a, Pringle 3459 (MO!; isotypes BM, E, JE, M, NY!, PH, UC, US, VT). Fig. 168C. Rootstock bilobed; leaves ca. 20, ca. 30 cm long, pale toward bases, triangular in mid-length transection; velum covering less than 25% of each sporangium; sporangial walls usually brownspotted; megaspores white, average 400–550 μm in diam., smooth or obscurely to boldly tuberculate-verrucate with girdle smooth; microspores light gray in mass, average 35–45 μm long, echinatetuberculate.

DF (Rzedowski 34261, ENCB). Jal (Pringle 4003, GH, L, MO, NY, VT). Me´x (Pringle 3459, BM, E, F, G, GH, JE, M, MEXU, MO, NY, PH, US, VT). Mich (Gonzalez 280, ENCB, MEXU, NY). Mor (Pringle 6660, BM, E, F, G, JE, LL, MO, NY, PH, UC, US, VT). Nay (McVaugh 18672, CAS, NY). Qro (Rzedowski 51180, IEB). Zac (Rzedowski & McVaugh 984, ENCB, IEB, MEXU, MICH, NY).

Isoe¨tes montezumae and I. mexicana appear to be closely related species that could be treated as a one variable species. Differences in spore size and ornamentation, in addition to what appear to be interspecific hybrids, support recognizing them as genetically divergent species until additional evidence proves otherwise. See discussion under I. mexicana. Most collections of I. montezumae can be identified by their larger spores, tuberculate megaspores, and sporangial walls that are spotted or streaked with patches of brownish cells. In contrast, I. mexicana has smaller spores and more often shiny, smooth, porcelain-like megaspores that are contained within unspotted sporangial walls.

4. ISOE¨TES ORCUTTII Isoe¨tes orcuttii A. A. Eaton, Fern Bull. 8: 13. 1900. Type. U.S.A. California: South Chollas Valley, San Diego, Orcutt 1242 (MO!; isotypes GH, US). Figs. 168D, H. Rootstock bi- or trilobed; leaves ca. 10, ca. 5 cm long, pale toward bases, triangular in mid-length transection; velum covering 100% of each sporangium; sporangial walls unspotted; megaspores white to dark gray, average 250–350 μm in diameter, smooth to obscurely tuberculate with girdle smooth; microspores brown in mass, average 25–35 μm long, tuberculate; 2n⫽22 (USA).

Distribution. Vernal pools; ca. 300 m. March–April. USA (Calif); Mexico. Distribution. Ponds, streams, grassy meadows, roadside ditches in seasonally wet, oak-pine woodlands; 2000–3500 m. August–November. Mexico; Guat. Selected Specimens Examined. Ags (McVaugh 818, ENCB, MICH). Chis (Breedlove 15070, DS, ENCB, F, LL, MEXU, MICH, NY, WIS).

Specimens Examined. BCN (Orcutt s.n., 12 Apr 1886, NY; Moran 26667, CAS, MICH).

Isoe¨tes orcuttii reaches the southern extent of its range in northern Baja, California. This species is distinguished from all other Mexican species by its complete velum. Occasionally, its megaspores vary in color from white to gray. Comparisons of

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DNA nucleotide sequences indicate that I. orcuttii and two other western North American species, I. nuttallii A. Braun and I. minima A. A. Eaton, are more closely related to Isoe¨tes species in the Mediterranean region than they are to other North American species.

5. ISOE¨TES PALLIDA Isoe¨tes pallida Hickey, Amer. Fern J. 78: 35. 1988. Type. Mexico. Oaxaca: Rain ponds along Hwy 180, 4 mi W of Costa Rica, Kral 25320 (MO!; isotypes MICH!, VDB!). Fig. 168E. Rootstock bi- or trilobed; leaves ca. 30, ca. 25 cm long, pale toward bases, triangular in mid-length transection; velum covering less than 25% of each sporangium; sporangial walls unspotted; megaspores white, average 400–475 μm in diam., pustulate-tuberculate with girdle smooth; microspores light gray in mass, average 25–35 μm long, echinate; 2n⫽44 (Oax).

Distribution. Temporary pools and saturated soil in seasonally wet savanna; 2800–3000 m. August–November. Mexico. Specimens Examined. Chis (Breedlove 20424, DS, MEXU). Jal (Pringle 3333, BM, E, G, GH, JE, M, MO, MICH, MU, NY, PH, UC, US, VT). Me´x (Arreguı´n 711, IEB, XAL; Pringle 13261, C, GH, L, MO, UC, US, VT, s.n., 11 August 1897, NY).

Isoe¨tes pringlei is the only Mexican species with echinatecristate megaspores. Megaspore ornamentation is easily observed with a 10⫻ hand lens.

5 9 . J A M E SO N I A Jamesonia Hook. & Grev., Icon. Filic. 2: t. 178. 1830. Type: Jamesonia pulchra Hook. & Grev.

Distribution. Temporary pools and saturated soil in seasonally wet savanna; ca. 100 m. August–November. Mexico; Bel. Specimens Examined. Oax (Hickey & Hickey 962, UC). Camp (Novelo 598, MEXU).

Specimens from the Campeche (Novelo 598), originally annotated as Isoe¨tes cubana Engelm. ex Baker and I. jamaicensis Hickey, bear megaspores averaging 400 μm in diameter with the pustulate-tuberculate ornamentation of I. pallida.

6. ISOE¨TES PRINGLEI Isoe¨tes pringlei Underw., Zoe 1: 98. 1890. Type. Mexico. Jalisco: near Guadalajara, Pringle 2633 (NY!; isotype MO!). Figs. 167D, 168F. Rootstock bilobed; leaves ca. 15, ca. 25 cm long, pale toward bases, half-terete at mid-length transection; velum covering less than 25% of each sporangium; sporangial walls unspotted; megaspores white, average 475–650 μm in diam., echinate-cristate with girdle granulate; microspores light gray in mass, average 35– 45 μm long, smooth.

Terrestrial; rhizomes short to long-creeping, with dark, bristlelike hairs; fronds linear, erect, small to medium-sized, generally indeterminate, the apices slow to fully unroll; stipes short, dark; blades 1-pinnate, the pinnae small, mostly round, imbricate, hairy or glandular, at least abaxially, herbaceous to coriaceous, margins generally inrolled; veins free, branching; sporangia along the veins, often obscured due to the dense indument and the curved pinna margins; false indusia absent or present; spores tetrahedral-globose, each with equatorial flange; x⫽87, prob. from x⫽29. Jamesonia has nineteen species largely of high elevation in tropical America from Mexico to Bolivia and southeastern Brazil. It is closely allied to Eriosorus, with which it hybridizes in Central and South America (A. Tryon, 1970; Sa´nchez B., 2000). The Oaxaca locality is the northernmost record for the genus. Jamesonia is distinct in the linear, 1-pinnate blades that are Ⳳ permanently circinate at apices. References Sa´nchez Baracaldo, P. 2000. Phylogenetics and biogeography of an Andean fern radiation, the Jamesonia-Eriosorus complex. Ph.D. thesis, Univ. California, Berkeley; Tryon, A. F. 1962. A monograph of the fern genus Jamesonia. Contr. Gray Herb. 191: 109–197; Tryon, A. F. 1970. A monograph of the fern genus Eriosorus. Contr. Gray Herb. 200: 54–174.

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LASTREOPSIS

1. JAMESONIA ALSTONII

trea recedens J. Sm. ex T. Moore] [⫽ Lastreopsis tenera (R. Br.) Tindale].

Jamesonia alstonii A. F. Tryon, Contr. Gray Herb. 191: 168. 1962. Type. Colombia. Valle: Cordillera Occidental, Los Farallones, bajo el filo de la Cordillera en el Cerro de La Torre, La Laguna, Cuatrecasas 21884 (GH!; isotypes COL, US!). Figs. 169A–C.

Polystichum sect. Parapolystichum Keyserl., Polyp. Herb. Bunge. 11, 45. 1873. Parapolystichum (Keyserl.) Ching, Sunyatsenia 5: 239. 1940. Type: Parapolystichum effusum (Sw.) Ching [⬅ Polypodium effusum Sw.] ⬅ Lastreopsis effusa (Sw.) Tindale.

Rhizomes long-creeping, 1 mm diam.; rhizome hairs concolorous, brown, 1–1.5 mm long; fronds 15–40 cm long; stipes less than 1⁄6 the frond length, dark reddish brown, densely pubescent, hairs brownish, 2 mm long, to 3–4 mm long near the unrolling tip; blades linear, 1-pinnate, 0.4–1 cm wide; pinnae coriaceous, orbicular-cordate, 2.5–3 mm diam., stalks 0.5–0.8 mm long, adaxial surfaces with sparse hairs 1–1.5 mm long to glabrous with age, abaxial surfaces with abundant hairs 1–1.5 mm long; sori elongate along the veins, hidden by curled margins of the pinnae (modified indusia); indusia 0.8 mm wide with fimbriate margins; spores tan.

Distribution. Rocky soil by streams, flat meadows and steep slopes with pines, junipers, and Buddleia; 2750–3600 m. Mexico; Guat, CR; Col, Ec, Peru, Bol. Specimens Examined. Chis (Breedlove 24314, DS; Matuda 2864, S234, NY, US). Oax (Mickel 5513, NY; Sharp & Webster 720A NY, TENN). Unverified, Doubtful, or Mistaken Reports. Tab (reported by Magan˜a, 1992, but plant figured is Nephrolepis cordifolia, ‘Duffii’, escaped from cultivation).

This is the only species of Jamesonia in Mexico and grows at the most northern location in the genus (Sharp, Amer. Fern J. 61: 187. 1971). The Oaxaca locality is paramo-like in being high elevation, grassy, wet, and with cloud cover. Jamesonia alstonii grows with Huperzia crassa, another paramo plant reaching its northernmost locality there.

60 . L A S T R E O PS I S Lastreopsis Ching, Bull. Fan Mem. Inst. Biol. 8: 157. 1938. Type: Lastreopsis recedens (J. Sm. ex T. Moore) Ching [⬅ Las-

Terrestrial, rarely epiphytic; rhizomes short- to long-creeping or suberect, scaly; fronds monomorphic, clumped; stipes scaly at base, with two prominent ridges adaxially, the channel between ridges with abundant short, erect, ctenitoid hairs (short, articulate, unbranched, catenate, hyaline to reddish hairs), rachises with similar channels continuous with channels of costae, ridges continuous with thickened margin of ultimate segments; blades firm-herbaceous, (2–)3–5-pinnate, pentagonal, proximal pinnae more developed basiscopically in many species (ours), pinnae catadromous, apices pinnatifid; indument of adpressed glandular hairs on abaxial blade surfaces; buds present near apices of rachises or costae in many species (ours); veins free, readily visible on both sides, simple or forking, ending before the blade margins; sori usually round, indusiate or exindusiate, indusia roundreniform; sporangia often with 1–2 stalked glands on sporangial pedicels or within sori; spores bilateral with cristate perispore; x⫽41. Lastreopsis is a pantropical genus of about 36 species of southtemperate and rain forest regions, of which five to six are American; only two species, both wide-ranging, occur in Mexico. Species are most numerous in the Australasian area, especially Australia, with 15 species (Jones in McCarthy, 1998). The genus is often said to be closest to Ctenitis, which has similar (but not identical) hairs and very dissected fronds. However, the configuration of the ridges on the minor axes and rachises in Lastreopsis is distinctive; in Ctenitis such ridges are lacking or, if present, they are interrupted and not decurrent onto the axes above or below. Preliminary molecular data place Lastreopsis within the large dryopteroid clade, perhaps allied to Megalastrum and Rumohra more closely than to Ctenitis (Cranfill, unpubl. data). References Christensen, C. 1920. A monograph of the genus Dryopteris. Part II. The tropical American bipinnate-decompound species. Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 8, 6: 1–132; Tindale, M. 1965. A monograph of the genus Lastreopsis Ching. Contr. New South Wales Natl. Herb. 3: 249–339, t. 9–23, map.

Key to the Mexican Species of Lastreopsis 1. Sori exindusiate; fronds 1–2 m long. .................................. 1. L. effusa. 1. Sori indusiate; fronds usually 40–90 cm long. ................................................................ 2. L. exculta subsp. guatemalensis.

1. LASTREOPSIS EFFUSA Lastreopsis effusa (Sw.) Tindale, Victorian Naturalist 73: 184. 1957.

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Polypodium effusum Sw., Prodr. 134. 1788. Dryopteris effusa (Sw.) Urb., Symb. Ant. 4: 16. 1903. Parapolystichum effusum (Sw.) Ching, Sunyatsenia 5: 239. 1940. Ctenitis effusa (Sw.) Copel., Gen. Filic. 124. 1947. Lectotype (chosen by Tindale, 1965: 300). Jamaica. Swartz s.n. (B-Willd. 19724; isolectotypes BR, photo US; UPS-Thunb. 24483, 24484, 24485, 24486, 24487, microfiche UC!).

Rhizomes short-creeping, mostly 5–10 mm diam.; fronds commonly 1–2 m long; stipes stramineous to brown, 50–100 cm long, ca. 1⁄2 the frond length, scaly at bases, scales brown, lanceolate, to 5–10 ⫻ 1.5 mm, Ⳳ entire; blades 50–100 ⫻ 50–100 cm, 3–5pinnate, proximal pinnae more developed basiscopically, distal pinnae unequal-sided with acroscopic pinnules longer and more spreading than those of basiscopic side, which arise at ⬍45⬚ from costae, each blade usually with a scaly bud in axil of a distal pinna; costae narrowly winged all or nearly all their length, wings decurrent onto next axis; pinnules of proximal and medial pinnae short- to often long-stalked to ca. 1 cm; penultimate segments coarsely and acutely toothed; indument abaxially and adaxially of scattered to moderate hyaline, septate, blunt hairs 0.1–0.3 mm long on rachises, costae, costules, also with blade surfaces and veins abaxially with usually sparse adpressed orangish elongate glands 0.1–0.15 mm; sori exindusiate, with stalked reddish glandular paraphyses among sporangia; 2n⫽82 (Jam., for the sp.), ca. 164 (Trin., for subsp. divergens). Lastreopsis effusa is easily distinguished from L. exculta by the exindusiate sori and larger fronds. Tindale recognized four subspecies of L. effusa, two of which occur in Mexico. The type subspecies, from the Greater Antilles and Surinam, has more glandular hairs on the abaxial laminae and glabrous abaxial costae and costules. Subspecies confinis (C. Chr.) Tindale, also from the Greater Antilles, has thicker, bipinnate-pinnatifid to tripinnate fronds. Key to the Mexican Subspecies of Lastreopsis effusa 1. Blades dark green, 4-pinnate to 4-pinnate-pinnatifid; ultimate segments acute. ............................................ 1a. subsp. dilatata. 1. Blades dull gray-green or dark green, 3-pinnate to 3pinnate-pinnatifid; ultimate segments obtuse to acutish.

Distribution. Slopes and ravines in wet montane forests; 100– 600 m. Mexico; Guat, Hond, CR, Pan; Cuba, Hisp; Ec. Selected Specimens Examined. Chis (Breedlove 70864, CAS). Gro (Vela´zquez 41, FCME; Lozano V. 205, FCME). Hgo (Kenoyer & Crum 3981, GH). Oax (Herna´ndez G. 2543, CHAPA, NY; Mickel 5922, UC). SLP (Kenoyer s.n., ARIZ). Ver (Purpus 3008, NY, UC, US; Riba et al. 1224, UAMIZ, XAL). Unverified, Doubtful, or Mistaken Reports. Mich (cited by Mickel & Beitel, 1988, but not verified).

Occasional specimens have more than a single rachis bud per blade, the buds being multiple also in axils of costules (e.g., Riba et al. 1224, UAMIZ). 1b. Lastreopsis effusa subsp. divergens (Willd.) Tindale, Contr. New South Wales Natl. Herb. 3: 299, pl. 21. 1965. Fig. 170H. Polypodium divergens Willd. in Schkuhr, 24. Kl. Linn. Pfl.-Syst. [Kr. Gew.] 27, pl. 26b. 1804–1809. Type. Venezuela. Caracas, Bredemeyer s.n. (B-Willd. 19725–3 sheets).

Blades dull gray-green or dark green, 3-pinnate to 3-pinnatepinnatifid; ultimate segments obtuse to acutish; 2n⫽ca. 164 (Trin.).

..................................................................................... 1b. subsp. divergens.

1a. Lastreopsis effusa subsp. dilatata Tindale, Contr. New South Wales Natl. Herb. 3: 299. 1965. Figs. 170A–E. Polypodium dilatatum Liebm., Mexic. Bregn. 208 (reprint 56). 1849, hom. illeg., non Hoffm, 1795. Lectotype (chosen by Smith, 1981: 139). Mexico. Veracruz: Mirador, Liebmann s.n. [Pl. Mex. 2674, Fl. Mex. 723] (C!). Because Liebmann’s species is a later homonym, we regard the publication of subsp. dilatata as new, dating from 1965.

Blades dark green, 4-pinnate to 4-pinnate-pinnatifid; ultimate segments acute.

Distribution. Steep ravines in lower montane rain forests; 100– 1000 m. Mexico; Guat, Hond, Nic, CR, Pan; Hisp, L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol, Parag. Moran (in Davidse et al., 1995) also cited Belize in the range, but his concept included also subsp. dilatata.

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Specimens Examined. Chis (Breedlove 32834, 38683, DS). Oax (Herna´ndez G. 1426, NY, 1526, MEXU, NY; Mickel 6845, NY, UC).

All specimens from Veracruz cited as subsp. divergens by Palacios-Rios (1992) are subsp. dilatata. Also, most specimens (e.g., Brenckle 199; Hatch & Wilson 116, 128; Standley 68447; Heyde & Lux s.n. [Donn. Sm. 4682]; all US) identified by Stolze (1981) as subsp. divergens are subsp. dilatata. Subspecies divergens is less dissected and occurs at lower elevations than subsp. dilatata.

2. LASTREOPSIS EXCULTA SUBSP. GUATEMALENSIS Lastreopsis exculta (Mett.) Tindale subsp. guatemalensis (Baker) Tindale, Contr. New South Wales Natl. Herb. 3: 245. 1963. Figs. 170F, G. Nephrodium guatemalense Baker in Hooker & Baker, Syn. Fil., ed. 2, 498. 1874. Dryopteris guatemalensis (Baker) Kuntze, Revis. Gen. Pl. 2: 812. 1891. Dryopteris exculta (Mett.) C. Chr. var. guatemalensis (Baker) C. Chr., Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 8, 6: 96. 1920. Lastreopsis guatemalensis (Baker) L. D. Go´mez, Brenesia 8: 50. 1976. Lectotype (chosen by Christensen, 1920: 96). Guatemala. Choctum, Salvin & Godman s.n. (K!). Aspidium chontalense E. Fourn., Bull. Soc. Bot. France 19: 254. 1872. Lastreopsis chontalensis (E. Fourn.) Lellinger, Proc. Biol. Soc. Wash. 89: 716. 1977. Type. Nicaragua. Chontales, Levy 516 (P!–3 sheets; isotype frag. US!).

Rhizomes short-creeping, 3–6 mm diam.; fronds 40–90(–110) cm long; stipes stramineous to brownish, 12–40 cm ⫻ 1.5–3(–5) mm, dark brown and scaly only at bases, scales brown, lanceolate, to ca. 5(–8) ⫻ 1 mm, Ⳳ entire; blades 15–30(–40) ⫻ 9–15(– 20) cm, tripinnate-pinnatifid at the bases, bipinnate-pinnatifid distally, proximal pinnae more developed basiscopically, distal pinnae decidedly unequal-sided, with acroscopic pinnules or segments longer and more spreading than those of basiscopic side, which arise at ⬍ 45⬚ from costae; costae narrowly winged all or nearly all their length, wings decurrent onto next axis; pinnules slightly stalked to sessile; penultimate segments coarsely and acutely toothed; indument abaxially and adaxially of scattered to dense hyaline to yellowish, septate, blunt hairs 0.1–0.3 mm long on rachises, costae, costules, and veins, also with blade surfaces and veins abaxially with often numerous adpressed orangish elongate glands 0.1–0.2 mm; sori round, indusiate, indusia tan to brown, round-reniform, 0.5–1 mm diam., persistent, glabrous or sparsely glandular.

Distribution. Terrestrial in lowland wet forests, very rare, few recent collections; 200 m. Mexico; Guat, Bel (Davidse 36208, UC), Hond, Nic, CR, Pan. Specimens Examined. Chis (Ghiesbreght 232A, GH, 422, K, US; Purpus 6724, GH, NSW, NY, UC, US, 6869, NY). Oax (Herna´ndez G. 1548, MEXU, NY). Ver ? (see discussion below).

This differs from L. effusa by the indusiate sori and smaller fronds. Subsp. exculta is known from Colombia, Venezuela, and Ecuador, and was said by Tindale (1965) to differ by the broader pinnules (1–2 cm wide) and larger blades (to 40–50 ⫻ 25–30 cm). Tindale (1965) cited a specimen of L. exculta subsp. squamifera (C. Chr.) Tindale from Veracruz (Schaffner 1900, NSW, not seen), but it seems likely that this specimen is either misidentified, wrongly localized, or that the subspecies is not distinct from subsp. guatemalensis. Syntypes and all other specimens of subsp. squamifera cited by Tindale are from Costa Rica, and that subspecies was said by Tindale to differ in having the rachises and costae rather densely clothed abaxially with red-brown, lanceolate, thin scales.

6 1 . L E L L I N GE R I A Lellingeria A. R. Sm. & R. C. Moran, Amer. Fern J. 81: 76. 1991. Type: Lellingeria apiculata (Kunze ex Klotzsch) A. R. Sm. & R. C. Moran [⬅Polypodium apiculatum Kunze ex Klotzsch]. Epiphytic; rhizomes radially symmetrical, short-creeping, ascending, or erect; rhizome scales usually blackish, clathrate, glabrous or bearing hyaline marginal setulae, attached across the width of scale base; phyllopodia absent; stipes absent or much shorter than the bases, continuous with (not articulate to) the rhizomes; blades shallowly to deeply pinnatisect, some species (the L. myosuroides group) with the fertile distal portion entire or less divided than the sterile proximal portion of the blade, rarely the blades pinnate-pinnatifid (outside Mexico); indument of setae, setulae, or branched hairs, at least along the stipes and rachises, the setae hyaline to reddish, usually with a pale basal cell and forked unequally, the smaller cell oblique, glandular, the larger cell acicular; hydathodes present (viewed adaxially); veins simple, free; sori round or elliptic, often somewhat sunken, ex-

LELLINGERIA APICULATA

indusiate, lacking paraphyses; sporangial capsules glabrous, rarely setulose; x⫽32, 33. Lellingeria comprises about 60 species and is primarily neotropical, but a few species are known from Africa, Madagascar, Hawaii, and the southern Pacific (Smith et al., 1991). The species that occur in the Old World and Hawaii all belong to the L. myosuroides group. Although sampling is inadequate, preliminary molecular results indicate that the genus is monophyletic if several species that bear long tawny hairs are removed (Ranker et al., 2004). These possibly misplaced species may be diminuative members of the Terpsichore cultrata group, and include only L. mitchellae in Mexico, plus another species in southern Central America and two in southern Brazil (Smith et al., 1991). From Ceradenia and Enterosora, Lellingeria differs in having conspicuous hydathodes. Micropolypodium and Terpsichore differ

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by the presence of long, stout, reddish brown setae on the midribs and blades and the non-clathrate rhizome scales. From Melpomene, Lellingeria differs in the generally setose rhizome scales and the absence of long reddish brown setae. Several species of Lellingeria, in particular the narrow-bladed ones with a single sorus per segment, have often been treated in the genus Xiphopteris. This generic concept, based largely on blade dissection, has now been discredited by both morphological and molecular evidence (Smith et al., 1991; Ranker et al., 2004). References Copeland, E. B. 1952. The American species of Xiphopteris. Amer. Fern J. 42: 41–52, 93–110; Maxon, W. R. 1914. Studies of tropical American ferns– No. 5. Notes upon Polypodium duale and its allies. Contr. U.S. Natl. Herb. 17: 398– 406; Smith, A. R., R. C. Moran & L. E. Bishop. 1991. Lellingeria, a new genus of Grammitidaceae. Amer. Fern J. 81: 76–88.

Key to the Mexican Species of Lellingeria 1. Blades 6–55 mm wide; hydathodes or sori 3–30 per fertile segment; fertile veins not visible. ............................................................... 1. L. apiculata. 1. Blades 2–6 mm wide; hydathodes or sori 1–2 per fertile segment; fertile veins often darkened. 2. Sporangial capsules conspicuously setulose; rhizome scales with numerous, hyaline, marginal setulae. ...................................... 5. L. mitchellae. 2. Sporangial capsules glabrous; rhizome scales absent or present and glabrous, or each scale with a single apical, oblique hair. 3. Fertile portion of blades incised 2⁄3–7⁄8 of the way to the rachis. 4. Fertile portion of blades incised ca. 2⁄3 of the way to the rachis; hairs of the rachises pale, often twisted. ..................... 2. L. delitescens. 4. Fertile portion of blades incised 7⁄8 of the way to the rachis; hairs of the rachises darkened, acicular. ................................. 4. L. limula. 3. Fertile portion of blades entire or incised less than halfway to the rachis. 5. Fertile portion of blades undulate or shallowly lobed. ............................................................................................................... 3. L. hellwigii. 5. Fertile portion of blades incised 1⁄3–1⁄2 the way to the rachis. ................................................................................................. 6. L. prionodes.

1. LELLINGERIA APICULATA Lellingeria apiculata (Kunze ex Klotzsch) A. R. Sm. & R. C. Moran, Amer. Fern J. 81: 83. 1991. Figs. 171 N, O. Polypodium apiculatum Kunze ex Klotzsch, Linnaea 20: 378. 1847. Ctenopteris apiculata (Kunze ex Klotzsch) Copel., Philipp. J. Sci. 84: 17. 1955 [1956]. Grammitis apiculata (Kunze ex Klotzsch) F. Seym., Phytologia 31: 176. 1975. Lectotype (chosen by R. M. Tryon & Stolze, Fieldiana, Bot., n.s., 32: 94. 1993). Venezuela. [Aragua:] Colonia Tovar, Moritz 247 (B, photo F; isolectotypes NY!, US!).

Rhizomes ascending; rhizome scales castaneous, clathrate, conspicuous, linear, long-attenuate, 3–3.5 mm long with setulose margin, setulae 0.3 mm long; fronds clumped; stipes gray-brown to black, 1⁄6–1⁄4 the frond length, with dense, short hairs 0.1–0.3 mm long; blades pinnatisect, oblanceolate, 12–15 ⫻ 2–3 cm, gradually narrowed to base, apices caudate, abruptly narrowed; rachises black, with dense hairs 0.3–0.5 mm long; pinnae 25–30 pairs, linear, acute to obtuse, costae black abaxially, sometimes immersed, with sparse hairs 0.3–0.5 mm long, costae green adaxially, glabrous; laminar surfaces glabrous; veins obscure, unbranched, lime dots absent; sori oblong, glabrous; sporangia glabrous.

Distribution. Epiphytic in wet montane forests; 1150–1600 m. Mexico; Guat, Hond, CR, Pan; Hisp; Col, Ven, Guy, Ec, Peru, Bol, s Braz. Specimens Examined. Chis (Breedlove 42073, DS, 53419, 53568, 56220, CAS; 56221, ENCB; Te´llez 7993, MEXU). Oax (Mickel 7120, NY).

Lellingeria apiculata is distinct from congeners in Mexico by its setulose rhizome scales, densely short-hairy stipes, and much longer and broader blades. It more closely resembles Ceradenia margaritata, Melpomene firma, or M. xiphopteroides in blade form. From the first of these it differs by the readily visible vein tips adaxially, the lack of whitish soral glands, and the lack of

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dark reddish brown setae on the stipes and blades. From species of Melpomene it differs by the lack of dark reddish brown setae on the blades and the presences of setulose-margined rhizome scales. It was not recorded from Guatemala by Stolze (1981), but a specimen has been seen from there (Donnell Smith 1758, NY).

2. LELLINGERIA DELITESCENS Lellingeria delitescens (Maxon) A. R. Sm. & R. C. Moran, Amer. Fern J. 81: 84. 1991. Figs. 171H, J. Grammitis myosuroides Schkuhr, Krypt. Gew. 1: 9, t. 7. 1804, hom. illeg., non Swartz, 1801. Polypodium delitescens Maxon, Bull. Torrey Bot. Club 32: 74. 1905. Xiphopteris delitescens (Maxon) Copel., Amer. Fern J. 42: 51. 1952. Grammitis delitescens (Maxon) Proctor, Bull. Inst. Jamaica, Sci. Ser. 5: 32. 1953. Type. Schkuhr described and illustrated a plant probably received from Swartz and likely collected in Jamaica (HAL?). Smith (1981), Proctor (1985), and Mickel and Beitel (1988) also regarded this as type, but Copeland (1952) chose Maxon 1513 (US!), from Jamaica, Blue Mountain Peak, as lectotype, and in this he was followed by Lellinger (1989). Because Maxon’s Polypodium delitescens was an avowed “nom. nov.,” and because Maxon specifically selected “Schkuhr’s original plant, if existent,” as type, we reject Copeland’s lectotypification.

Rhizomes ascending; rhizome scales brown-black, clathrate, lacking setulae, ovate-lanceolate, 1.5–2 mm long; fronds clumped; stipes castaneous to black, 2–5 mm long, pilose with straight pale brownish hairs 0.3 mm long; blades linear, 4–8 cm long, gradually tapering at both ends, slightly hemidimorphic; rachises with sparse to scattered hairs 0.3 mm long, some of these unevenly bifurcating; proximal portion of blades pinnatisect; pinnae slightly ascending, (1–)1.5–2.5 ⫻ 1 mm, oblong to deltate, obtuse, costae obscure, not evident, terminal portion of blades pinnatifid, cut ca. 2⁄3 way to the rachises, 3.5–4 mm between opposing lobes, 1– 1.5 mm wide between opposing sinuses, sinuses 1.3–1.5 mm deep, costae often barely evident; sori oblong, one per lobe, basal, on terminal pinnatifid portion of blades and onto pinnatisect portion, extending beyond base of sinus on the veins, glabrous; sporangia glabrous; 2n⫽132–138 (Jam).

See discussions under Lellingeria prionodes and L. limula for comparison with those species which are closely related. All Chiapas specimens cited as Grammitis delitescens by Smith (1981) are here referred to either Lellingeria limula or L. prionodes. Also, specimens identified as L. delitescens from Guatemala (Stolze, 1981) are mostly L. prionodes; the sole specimen cited by Lellinger from Panama (Valdespino et al. 268, US) is L. myosuroides, and the Costa Rican one is likely also misidentified. Thus, the species appears to be absent from Central America.

3. LELLINGERIA HELLWIGII Lellingeria hellwigii (Mickel & Beitel) A. R. Sm. & R. C. Moran, Amer. Fern J. 81: 84. 1991. Figs. 171A, B. Grammitis hellwigii Mickel & Beitel, Mem. New York Bot. Gard. 46: 199. 1988. Type. Mexico. Oaxaca: Distrito Ixtla´n, road from Oaxaca to Tuxtepec, 30 mi beyond Ixtla´n, Hellwig 245 (NY!).

Rhizomes ascending, ca. 1 mm diam.; rhizome scales clathrate, lanceolate, 0.8–1 mm long, often with terminal setae; fronds clumped; stipes blackish, 3–10 mm long, glabrous to very sparsely hairy; blades strongly hemidimorphic, 3–10 cm long, sterile portion deeply pinnatifid, cut nearly to the rachis, lobes 7–12 pairs, nearly round to more elongate toward base of blades, 1.5–2 mm wide (widest near lobe tip), lobe bases narrowed, 1–1.5 mm wide, a few small (0.3 mm long), inconspicuous, unequally branched, bilobed hairs along the margin and underside of rachises; distal half of blades fertile, shallowly lobed to barely undulate, 3–3.5 mm between opposing lobes, 2.3–3 mm between opposing sinuses, sinuses 0.3–0.5 mm deep, veins in fertile portion darkened only beneath the sori; sori round to slightly oblong, in single rows along each side of rachises, glabrous; sporangia glabrous.

Distribution. Epiphytic, only in cloud forests of northeastern Oaxaca; 2450–2750 m. Mexico. Specimens Examined. Oax (Breedlove 65940, CAS; Mickel 5502, NY, UC, 4324, 4636, 4926, 6782, NY).

Distribution. Epiphytic in wet montane forests; 900–1350 m. Mexico; Cuba, Jam. Specimens Examined. Oax (Hallberg 1729, NY; Mexia 9124a, CAS, GH, MO, NY, UC, US). Ver (Nee et al. 25071, F, XAL).

This species is very closely allied to L. myosuroides (see Excluded Species), which is widespread in tropical America (see also discussion under L. prionodes for comparison with that species and L. delitescens). Lellingeria myosuroides has longer rhizome

LELLINGERIA PRIONODES

scales (1.5–2 mm long), scattered bifurcating hairs on the stipes and rachises, sinuses on the fertile portion 0.3 mm deep, sterile lobes narrow and broadest at the base (1.5–2 mm long, 0.5–0.8 mm wide at midpinna), and veins in the fertile portions darkened beyond the receptacle, almost to the blade margin.

4. LELLINGERIA LIMULA Lellingeria limula (Christ) A. R. Sm. & R. C. Moran, Amer. Fern J. 81: 84. 1991. Figs. 171C–E. Polypodium limulum Christ, Bull. Soc. Bot. Gene`ve, se´r. 2, 1: 218. 1909. Xiphopteris limula (Christ) Pic. Serm., Webbia 28: 472. 1973. Grammitis limula (Christ) L. D. Go´mez, Brenesia 8: 47. 1976. Syntypes. Costa Rica. San Jose´: La Palma, Pittier 708; Tonduz 12595; Brade 79; Werckle´ s.n. (all P; isosyntype of Tonduz 12595 at US!).

Rhizome scales dark brown to blackish, 0.5–1 mm long, ovate to lanceolate, glabrous; stipes 0–1 cm, sparsely to moderately setulose, the setulae castaneous, 0.3–0.5 mm, bifurcate with a pale glandular cell and short base; blades 4–15 ⫻ 0.3–0.5 cm, linear, pinnatisect all along the length of blades, glabrescent or sparsely setulose on the margins and on both surfaces; segments 1–1.5 mm wide at the base of the sinuses; rachises straight (not flexed), with hairs similar to those of stipes; fertile veins visible only proximal to the sori; hydathodes one per fertile segment; sori elongate, slightly imbedded and enclosed by the somewhat conduplicate segments; sporangia glabrous; 2n⫽64 (CR).

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5. LELLINGERIA MITCHELLAE Lellingeria mitchellae (Baker ex Hemsl.) A. R. Sm. & R. C. Moran, Amer. Fern J. 81: 85. 1991. Figs. 171K–M. Polypodium mitchellae Baker ex Hemsl., Biol. Cent.-Amer., Bot. 3: 664. 1885. Xiphopteris mitchellae (Baker ex Hemsl.) Copel., Amer. Fern J. 42: 97. 1952. Grammitis mitchellae (Baker ex Hemsl.) F. Seym., Phytologia 31: 174. 1975. Type. Belize. Orange Walk, Mitchell s.n. (K).

Rhizome scales 0.5–1 mm long, ovate to lanceolate, brownish to blackish, setulose; stipes absent; blades 1.5–6.5 ⫻ 0.2–0.6 cm, linear, incised to 1⁄2–2⁄3 of the way to the rachis along their length, setose, the hairs ca. 0.5 mm long, pale, spreading; segments 1–2 mm wide at the base of the sinuses; rachises straight, moderately to densely setose, the setae ca. 0.5 mm, pale, spreading; fertile veins not visible; hydathodes 2 per fertile segment; sori round, not immersed; sporangia conspicuously setose on capsules.

Distribution: Epiphytic in lower montane and montane rain forests; 200–1200 m. Mexico; Guat, Bel, Nic, CR, Pan. Specimens Examined. Chis (Breedlove 48307, CAS, ENCB, NY, 48271, 53561, 53562, 55404, 57682, CAS, 56226, CAS, ENCB; Martı´nez S. 18795, 18869, MEXU).

Distribution. Epiphytic in montane rain forests and cloud forests; 900–1300 m. Mexico; Guat, Hond, Nic, CR, Pan; Col, Ven, Ec.

This species is readily distinguished from congeners in Mexico by the conspicuously setose sporangial capsules. In blade form and venation (costae with a single acroscopic branch) it more closely resembles species of Micropolypodium, but species of Micropolypodium have lighter colored rhizome scales, dark setae on the blades, and glabrous sporangia. Unpublished molecular data suggest that this species may be better placed in the group of Terpsichore cultrata (Ranker et al., ined.).

Specimens Examined. Chis (Breedlove 30808, DS, MEXU, NY, 31532, 35225, DS).

Lellingeria limula is distinguished from L. delitescens by being pinnatisect to the apex of the blades, thus lacking the slightly modified terminal portion found in L. delitescens. Blade segments are often somewhat conduplicate, a feature not found in close allies. The Chiapas specimens were cited under Grammitis delitescens by Smith (1981).

6. LELLINGERIA PRIONODES Lellingeria prionodes (Mickel & Beitel) A. R. Sm. & R. C. Moran, Amer. Fern J. 81: 86. 1991. Figs. 171F, G. Grammitis prionodes Mickel & Beitel, Mem. New York Bot. Gard. 46: 203. 1988. Type. Mexico. Oaxaca: Dto. Ixtla´n, 58 km N of Ixtla´n, 11 km N of high point at Cerro Pelo´n, Mickel 4270 (NY! isotype UC!).

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Rhizomes ascending; rhizome scales brown-black, clathrate, ovate-lanceolate, 1.5–2 mm long; fronds clumped; stipes castaneous to black, 2–5 mm long, pilose with straight red-brown hairs 0.3 mm long; blades linear, 3–11 cm long, gradually tapered at both ends, strongly dimorphic, fertile part confined to distal half of blades, rachises with scattered, unevenly bifurcating hairs 0.3 mm; sterile portion of blades pinnatisect, pinnae slightly ascending, 2–3.5 ⫻ 0.8–1 mm, oblong to deltate, acute to obtuse, costae obscure; hydathodes present; fertile portion of blades deeply lobed, cut 1⁄3–1⁄2 way to the rachis, 2.5–4.5 mm wide between opposing lobes, 1.5–2 mm wide between opposing sinuses, sinuses 0.5–1 mm deep, costae often barely evident; sori oblong, restricted to the terminal deeply lobed portion of blades, one per lobe, basal, not extending on veins beyond bases of sinuses, glabrous; sporangia glabrous.

Distribution. Epiphytic in wet montane forests and cloud forests; 1050–2800 m. Mexico; Guat (Martı´nez S. 13152, MEXU; Maxon & Hay 3320, US), Hond, Salv, Nic. Selected Specimens Examined. Chis (Breedlove 22930, DS, NY, 53299, CAS, ENCB). Hgo (Gimate 799, CHAPA, MEXU, NY; Moore 5287, UC, US). Oax (Rzedowski 33365, ENCB; Smith 473, UC). Pue (Pringle 13853, CAS, LL, US, mounted with L. delitescens from Oaxaca; Riba et al. 285, MEXU). Ver (Lira 28, XAL; Palacios Rios 17, XAL).

The degree of blade dissection, especially that of the fertile distal portion, and soral placement relative to both the proximal portion of the blades and the base of the sinuses are critical features distinguishing taxa in the L. myosuroides complex. Species in which the distal portion of the blade is strictly entire occur outside Mexico. These include L. anamorphosa (Proctor) A. R. Sm. & R. C. Moran, from Jamaica, with blades 4–4.5 mm wide distally. Grammitis skutchii (Maxon) F. Seym., from Guatemala, and Xiphopteris jamesonii Hook., from South America, have a subentire to weakly crenulate-sinuate fertile blade tip, but are probably only extreme forms of Lellingeria myosuroides (see Excluded Species). In material of L. myosuroides from the Greater Antilles, the fertile portion of the blade is crenulate-sinuate (sinuses 0.5 mm or less deep). Other relatives include L. delitescens, which has the distal portion of the blades pinnatifid (cut 2⁄3 of the way to the rachises, sinuses 1–1.3 mm deep) and sori on the pinnatisect basal portion of the blades, as well as on the distal portion. Even on the distal

part of blades, the sori continue on the veins beyond the sinus bases. Lellingeria limula has the segments incised even further toward the rachises, ca. 7⁄8 of the distance. Lellingeria prionodes has blades pinnatisect proximally, but the fertile portion is only shallowly cut (less than halfway to rachis, sinuses 0.5–1 mm deep) into sharply acute lobes; sori are restricted to the modified terminal portions of the blades and do not extend beyond the sinus bases. Excluded Species Lellingeria myosuroides (Sw.) A. R. Sm. & R. C. Moran, Amer. Fern J. 81: 85. 1991. Polypodium myosuroides Sw., Prodr. 131. 1788. Grammitis myosuroides (Sw.) Sw., J. Bot. (Schrader) 1800(2): 18. 1802. Xiphopteris myosuroides (Sw.) Kaulf., Enum. Filic. 85, 275. 1824. Type. Jamaica. Swartz s.n. (S, frag. & photo US). Xiphopteris jamesonii Hook., Sec. Cent. Ferns, pl. 14. 1860. Polypodium jamesonii (Hook.) Jenman, Bull. Bot. Dept. Jamaica, n.s. 4: 112. 1897. Type. Ecuador. Jameson s.n. (K). Polypodium skutchii Maxon, Proc. Biol. Soc. Wash. 51: 34. 1938. Xiphopteris skutchii (Maxon) Copel., Amer. Fern. J. 42: 51. 1952. Grammitis skutchii (Maxon) F. Seym., Phytologia 31: 175. 1975. Type. Guatemala. Chimaltenango: Buena Vista, above Tecpam, Skutch 96 (US!; photo UC!). Also seen: Skutch 774, NY, from same locality, 10,000 ft. Distribution. Guat, Hond, CR, Pan; Cuba, Jam, PR; Col, Ven, Ec, Bol, Braz. This species has not yet been found in Mexico. Smith (1981) cited two collections, both of which are referable to L. prionodes.

6 2 . L I N D SA E A Lindsaea Dryand. in J. E. Smith, Me´m. Acad. Roy. Sci. (Turin) 5: 401. 1793. Type: Lindsaea trapeziformis Dryand. [⫽ Lindsaea lancea (L.) Bedd. var. lancea]. For additional synonymy, see Kramer (1957).

Terrestrial, rarely epipetric or (not in ours) epiphytic; rhizomes long- to short-creeping, scaly; fronds small to medium-sized, monomorphic; stipes glabrous, stramineous to castaneous, often quadrangular; blades (simple–)1–2(–3)-pinnate, anadromous; segments generally dimidiate, membranous to chartaceous; veins free (in ours), rarely netted without included veinlets; sori submarginal in a continuous or interrupted line, subtended by marginfacing indusia, paraphyses generally present; spores tetrahedral (in ours), rarely bilateral; x⫽34, 41, 47, 50. Lindsaea is a genus of 150 species, about 1⁄3 from the American tropics, mostly though of Southeast Asia and adjacent Pacific area. It is most closely allied to Odontosoria (Wolf, 1995). Lindsaea is probably an ancient genus, judging from its diverse chromosome numbers (x⫽34, 41, 47, 50), protostelic, scale-hair transitions, and broad distribution. At low to middle elevations, in wet forests, it often grows with Adiantum, with which it is superficially confused, and

LINDSAEA KLOTZSCHIANA

mixed collections or misidentifications frequently result. Lindsaea resembles Adiantum in blade dissection, segment shape and marginal sori, but is distinct in the paler axes and the sori with outward-opening (extrorse) indusia.

365 References

Kramer, K. U. 1957. A revision of the genus Lindsaea in the New World with notes of allied genera. Acta Bot. Neerl. 6: 97–290; Kramer, K. U. 1971. Lindsaea group. Flora Malesiana 1(3): 177–254.

Key to the Mexican Species of Lindsaea 1. Distal segments of individual pinnae little reduced, more than half as long as proximal ones; terminal segment of each pinna deltate-lanceolate, often pinnatifid at base; rachises at insertion of basal pinnae angular abaxially; indusial margins entire. .............. 3. L. lancea. 1. Distal segments of individual pinnae of individual pinnae gradually reduced, much less than half as long as proximal ones, confluent with small terminal segment; rachises at basal pinna insertion angular or terete abaxially; indusial margins entire, subentire or deeply erose. 2. Indusial margins erose; pinnae strongly ascending. .................................................................................................................................... 5. L. stricta. 2. Indusial margins entire; pinnae spreading. 3. Segment margins incised; indusia discontinuous. ....................................................................................................................... 2. L. klotzschiana. 3. Segment margins entire; indusia continuous along margins. 4. Rachises at basal pinnae insertions angular abaxially. ....................................................................................................... 4. L. quadrangularis. 4. Rachises at basal pinnae insertions terete or slightly flattened abaxially. .................................................................................. 1. L. arcuata.

1. LINDSAEA ARCUATA Lindsaea arcuata Kunze, Linnaea 9: 86. 1834. Figs. 172N–P. Lindsaea trapeziformis Dryand. var. arcuata (Kunze) Baker in Martius, Fl. Bras. 1(2): 355. 1870. Lindsaea lancea (L.) Bedd. var. arcuata (Kunze) Rosenst., Hedwigia 46: 80. 1906. Type. Peru. Pampayaco, Poeppig 1133 (LZ, destroyed; isotype B!). Lindsaea curvans Fe´e, Me´m. Foug. 5: 106. 1852. Type. Mexico. Oaxaca: Galeotti 6489 (P!). Lindsaea galeottii Fe´e, Me´m. Foug. 5: 107. 1852. Type. Mexico. Oaxaca: Galeotti 6469 (P!). For additional synonymy, see Kramer (1957).

Rhizomes short- to long-creeping, 2–3 mm diam.; rhizome scales lustrous dark brown, lanceolate, 1–2 ⫻ 0.3 mm; fronds 40– 70 cm long; stipes ca. 1⁄2 the frond length, yellow-green to castaneous to atropurpureous, quadrangular in cross-section, glabrous; rachises at insertion of basal pinnae terete or slightly flattened abaxially; blades bipinnate; pinnae (1–)2–3(–5) pairs, segments arcuate, dimidiate, lacking midveins; sori 1–2 per segment; indusia continuous, 0.5 mm wide, margins entire; spores tetrahedral; 2n⫽ca. 168 (Oax), ca. 176 (Chis).

Distribution. Wet montane forests; 250–1500 m. Mexico; Guat, Hond, Nic, CR, Pan; Cuba, Hisp; Col, Ven, Ec, Peru, se Braz, Bol. Selected Specimens Examined. Chis (Breedlove 31517, DS, NY; Castillo C. et al. 3810, XAL; Mu¨nch 7, DS, US; Rovirosa 963, PH). Oax (Hallberg 1554, NY; Herna´ndez G. 2492, CHAPA, NY; Maya J. 2138, NY; Mickel 5698, 6695, NY; Rzedowski 30665, NY).

Lindsaea arcuata is distinct by its entire indusia and terete rachises at insertion of basal pinnae. Most Mexican collections have segments twice as long as wide whereas Central American specimens have segments 3–3.5 times as long as broad.

2. LINDSAEA KLOTZSCHIANA Lindsaea klotzschiana Moritz in Ettingshausen, Farnkr. 212, t. 145, f. 12. 1865 [1864]. Type. Venezuela. [Aragua:] Tovar, Moritz 238 (B; isotypes HBG, P). Figs. 173D–F. Rhizomes short-creeping, 2–3 mm diam.; rhizome scales reddish brown, lanceolate-deltate, 1–2 mm; fronds 40–46 cm, 3–5 mm distant; stipes nearly 1⁄2 the frond length, stramineous to brown, 1–1.5 mm diam., glabrous; rachises sharply quadrangular at attachment of basal pinnae; blades bipinnate; pinnae spread-

366

LINDSAEA

ing, 2–3 pairs; segments arcuate, dimidiate, lacking midveins, acroscopic margins incised; sori 3–6 per segment, generally 1 on each segment lobe; indusia discontinuous, 0.5 mm wide, margins entire; spores tetrahedral.

Distribution. Terrestrial in wet forests; 40–1150 m. Mexico; Bel, Guat, Hond, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Sur, Ec, Peru, Braz, Bol, Parag. Distribution. Terrestrial on steep slopes in wet forests, 750– 900 m. Mexico; Guat, Bel, Hond, CR, Pan; Col, Ven. Specimen Examined. Ver (Nee & Calzada 22710, NY, XAL).

This species is the only Mexican species to have incised pinna margins with discontinuous sori, although one specimen of L. arcuata (Fryxell 2562, NY, from Oaxaca) has one frond with continuous sori and two apparently juvenile fronds with irregularly discontinuous sori.

3. LINDSAEA LANCEA Lindsaea lancea (L.) Bedd., Suppl. Ferns Brit. Ind. 6. 1876. Figs. 172J–M. Adiantum lanceum L., Sp. Pl., ed 2, 2: 1557. 1763. Type. Seba, Thes. 2: t. 64, f. 7, 8, 1735, based on a specimen from Surinam. Lindsaea falcata Dryand., Trans. Linn. Soc. London 3: 41, t. 7, f. 2. 1797. Lindsaea lancea (L.) Bedd. var. falcata (Dryand.) Rosenst., Hedwigia 46: 79. 1906. Type. French Guiana, Aublet s.n. (BM, photo US). Lindsaea imbricata Liebm., Mexic. Bregn. 269 (reprint 117). 1849, hom. illeg., non Desv., 1811. Lectotype (chosen by Smith, 1981: 140). Mexico. Oaxaca: San Pedro Tepinapa, Liebmann s.n. [Pl. Mex. 2447, Fl. Mex. 450 (annotated by Kramer as holotype)] (C!, photos US!). Other syntype is Fl. Mex. 450a. For additional synonymy, see Kramer (1957).

Rhizomes short- to long-creeping, 1.5–3 mm diam.; rhizome scales lustrous, dark brown, lanceolate, 1–2 ⫻ 0.3 mm; fronds clumped, 10–50 cm long; stipes 1⁄3–1⁄2 the frond length, yellowish to dark brown especially at bases, angular; rachises at insertion of basal pinnae sharply angular abaxially; blades 1–2-pinnate; pinnae 10–20(–35) cm long, pinnae or pinnules subtrapezoidal, lunulate to subfalcate, glabrous; veins free, 2-forked; sori continuous on segment margins; indusia continuous, 0.2–0.5 mm wide, margins entire, 2n⫽82–88 (Trin).

Selected Specimens Examined. Chis (Breedlove 34289, DS, NY). Oax (Hallberg 1581, 1603, NY; Herna´ndez G., 1447, 2109, NY; Mickel 1048, ENCB, ISC, NY, 6516, ENCB, NY; Rzedowski 32965, ENCB). Tab (Ventura A. 20131, ENCB). Ver (Ventura A. 17958, XAL). Unverified, Doubtful, or Mistaken Reports. Gro (Lorea 2884, FCME, cited by Lorea-Herna´ndez & Vela´zquez M., 1998, but not verified).

Lindsaea lancea is the most common species of the genus in Mexico and distinct by its somewhat falcate pinnae/pinnules and the distal pinnules of a pinna being only slightly reduced. Frond dissection is too variable to distinguish var. falcata (1-pinnate) from other material in Mexico. Nearly half the specimens are 1pinnate, with some specimens having both 1-pinnate and 2pinnate fronds.

4. LINDSAEA QUADRANGULARIS Lindsaea quadrangularis Raddi, Opusc. Sci. 3: 294. 1819. Lindsaea lancea (L.) Bedd. var. quadrangularis (Raddi) Rosenst., Hedwigia 46: 80. 1906. Type. Brazil. Vic. of Rio de Janeiro, Raddi s.n. (FI).

Rhizomes short-creeping, 1.5–3 mm diam.; rhizome scales castaneous, 1–2 ⫻ 0.3 mm; fronds 15–40 cm long, clumped; stipes 1 ⁄2–2⁄3 the frond length, yellow or brown at bases, grooved, glabrous; rachises at insertion of basal pinnae strongly angular abaxially, rest of rachis grooved abaxially; blades bipinnate, each with 1–4 pairs of alternate to subopposite pinnae; pinnae with 15–40 pairs of segments, mostly 12–18 ⫻ 5–8 mm, glabrous; veins 1- or 2-forked; indusia continuous, 0.5 mm wide, margins subentire. Distribution. Terrestrial and moist forest; 0–1500 m. Mexico; Guat, Bel, Hond, Nic, CR, Pan; Cuba, PR, L Ant; Col, Ven, Ec, Braz, Bol, Parag. Key to the Mexican Subspecies of Lindsaea quadrangularis 1. Blade segments generally 5–6 mm long; Jal. ........... 4a. subsp. mexiae. 1. Blade segments mostly 8–17 mm long; Ver, Oax, Gro, Chis. ........................................................................... 4b. subsp. subalata.

LINDSAEA STRICTA

4a. Lindsaea quadrangularis subsp. mexiae Mickel, subsp. nov. Type. Mexico. Jalisco: West of San Sebastian, Hacienda del Ototal. Arroyo de Los Hornos, Mexia 1837 (holotype NY!; isotypes F!, G, GH!, MICH!, MO!, UC!, US!). Figs. 173A–C. Ab varietatibus aliis L. quadrangularis segmentis arcuatis 5–6 mm abiens. (Honoring Ynes Mexia, who made many important collections of Mexican ferns.)

367

Mickel and Beitel (1988) mistakenly reported subsp. antillensis Kramer from Mexico (Ver, Oax, Chis) and Central America, but these specimens belong to subsp. subalata.

5. LINDSAEA STRICTA VAR. STRICTA Lindsaea stricta (Sw.) Dryand, Trans. Linn. Soc. London 3: 42. 1797. Figs. 172A–C, Q–S. Adiantum strictum Sw., Prodr. 135. 1788. Type. Jamaica. Swartz s.n. (S, photo & frag. US!; isotypes B-Willd. 20055, BM, photo US). Lindsaea portoricensis Desv., Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck. Gesammten Naturk. 5: 326. 1811. Type. “Antilles,” apparently from Puerto Rico. s. coll. s.n. (P!; the smaller part of the material belongs to L. lancea). Lindsaea microphylla C. Presl, Reliq. Haenk. 1(1): 59, t. 10, f. 2. 1825, hom. illeg., non Swartz, 1801. Type. Mexico. Haenke s.n. (PRC). Lindsaea multifrondosa Fe´e, Me´m. Foug. 5: 105, t. 28, f. 1. 1852. Type. Mexico. Oaxaca: Talea, Galeotti 6470 p.p. (P!; isotypes B, BR!–3 sheets, photo US!, G, LE!). For additional synonymy, see Kramer (1957).

Distribution. Shady streambanks; 1500 m. Known only from the type collection. Kramer (1957) considered this to be L. stricta f. moritziana (Klotzsch) Kramer, but its characters match those of L. quadrangularis much more closely. 4b. Lindsaea quadrangularis subsp. subalata Kramer, Acta Bot. Neerl. 6: 190. 1957. Type. Costa Rica. El General, Skutch 2241 (US!; isotypes GH, MO, NY!). Figs. 172D–H.

Distribution. Terrestrial in wet forests; 0–1000(–1400) m. Mexico; Guat, Bel, Hond, Nic, CR, Pan; Cuba; Col, Ven.

Rhizomes short-creeping, ca. 2–3 mm diam.; rhizome scales castaneous, linear-lanceolate, 1–2 mm long; fronds 25–85 cm long, clumped; stipes ca. 2 mm diam., stramineous to dark castaneous toward the bases, glabrous; blades 1-pinnate or with 1– 4 pairs of pinnate pinnae, each ascending and nearly as long as the conform terminal pinna; rachises at insertion of basal pinnae terete to slightly flattened abaxially, flattened to shallowly grooved adaxially; pinna rachises terete at base, distally becoming quadrangular in cross section; segments many, mostly 10 ⫻ 5 (to 6 ⫻ 4) mm, arcuate; sori generally continuous, 1 per segment; indusia 0.2–0.5 mm wide, erose; 2n⫽ca. 196 (Trin).

Distribution. Dry clay banks on Atlantic lowlands; 300–500 m. Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Gr Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, s Braz, Bol.

Specimens Examined. Chis (Breedlove 21891, DS). Oax (Herna´ndez G. 1676, 2035, 2536, 2675, NY; Martı´nez C. 415, UC, US; Mickel 6517, NY). Ver (Nee et al. 25042, XAL; Smith 2078, MO p.p.).

Selected Specimens Examined. Chis (Martı´nez S. 10084, 12044, 13889, MEXU). Oax (Figueroa L. 91, ENCB; Galeotti 6470, BR; Herna´ndez G. 248, NY; Mexia 9138, CAS, NY, US; Mickel 7372, NY, US; Torres C. 12055, MEXU). Ver (Orcutt 3197, DS, US; Smith 57, US).

Unverified, Doubtful, or Mistaken Reports. Gro (Campos 1692, FCME, cited by Lorea-Herna´ndez & Vela´zquez M., 1998, but not verified).

Unverified, Doubtful, or Mistaken Reports. Camp (Martı´nez S. et al. 27824, MEXU, XAL, cited by Palacios-Rios, 2002a, but not verified).

368

LLAVEA

According to Kramer (1957), Lindsaea stricta is closely related to L. portoricensis and often difficult to separate from it, the latter having larger segments, thinner texture, and evident veins. These characters, however, do not seem to hold and may reflect a more shaded habitat. Lindsaea portoricensis is here included within L. stricta. Mexican material belongs to var. stricta. Two other varieties of L. stricta occur in Central and South America, Cuba, and Puerto Rico.

segments linear, 2.5–8 cm ⫻ 1.2–2 mm, margins differentiated, 0.5 mm wide, erose, thin, continuous, covering the sori, with short, glandular hairs on inside of indusial flap; 2n⫽58 (Hgo, NL).

63. LLAVEA Llavea Lag., Gen. Sp., pl. 33. 1816. Type: Llavea cordifolia Lag. Ceratodactylis Hook. in Hooker & Bauer, Gen. Fil., t. 36. 1840. Type: Ceratodactylis osmundoides Hook. ⫽ Llavea cordifolia Lag. Botryogramma Fe´e, Me´m. Foug. 5: 166. 1852. Type: Botryogramma karwinskii (Kunze) Fe´e [⬅ Allosorus karwinskii Kunze] ⫽ Llavea cordifolia Lag.

Terrestrial or epipetric; rhizomes stout, horizontal; rhizome scales mostly black, to bright yellow at stipe bases; fronds hemidimorphic, 2–3-pinnate, glabrous; veins free; sterile segments ovate with truncate to narrowly cuneate bases, 8–12 mm wide; blades fertile in distal third, fertile segments constricted, linear, 1.5–2 mm wide, margins inrolled, covering the sori, those slightly elongate along the veins; indusia modified, glandular hairs on inside surface; spores tetrahedral-globose; x⫽29. Llavea is distinct by its hemidimorphic fronds, recurved margin on the fertile segments, elliptic sterile segments, and the large, black rhizome scales, bright yellow at stipe bases. Llavea is a monotypic genus. Copeland (1947) regarded it as a derivative of Pellaea, but Tryon & Tryon (1982) cited recent placement of Llavea near Cryptogramma and Onychium. Molecular evidence (Gastony & Rollo, 1995) has shown Llavea to be not at all close to the cheilanthoid ferns, but instead related to Coniogramme of Asia.

1. LLAVEA CORDIFOLIA Llavea cordifolia Lag., Gen. Sp., pl. 33. 1816. Type. “Nova Hispania.” [Mexico.] s. coll. s.n. (MA or FI). Figs. 174A–D. Allosorus karwinskii Kunze, Linnaea 13: 138. 1839. Botryogramma karwinskii (Kunze) Fe´e, Me´m. Foug. 5: 166. 1852. Type. Mexico. Karwinski s.n.. (M?). Ceratodactylis osmundoides J. Sm. in Hooker & Bauer, Gen. Fil., t. 36. 1839. Type. Mexico. Lambert s.n. (K).

Rhizomes horizontal, compact, to 2.5 cm diam.; rhizome scales concolorous at bases, lustrous black, linear-lanceolate, 0.6–1.4 cm ⫻ 1–2 mm; fronds 60–120 cm long, clumped; stipes ca. 1⁄2 the frond length, stramineous, grooved, glabrous except for large, stramineous, linear-lanceolate scales 1–3 cm ⫻ 1–2 mm at stipe bases; blades broadly ovate, mostly 3-pinnate at bases; segments ovate-deltate, 2–3 ⫻ 0.8–2.2 cm, truncate to narrowly cuneate at bases, obtuse to acute at apices, glabrous, margins serrulate and clear; veins free, 1–2-forked, vein endings prominent; fertile

Distribution. Terrestrial on rocky slopes of moist woods and cliffs; 1450–2500 m. Mexico; Guat. Selected Specimens Examined. Chis (Matuda 4642, F, GH, MEXU, MICH). Coah (Lyonnet 421000003, MEXU). Dgo (Houck 4, MICH). Gro (Vela´zquez 476, FCME). Gto (Rzedowski 41546, XAL). Hgo (Copeland herb. s.n., 27 Dec 1937, MICH). NL (Pringle 1990, DS, F, GH, MICH, NY, UC). Oax (Camp 2750, GH, MICH, NY). Pue (Arse`ne 2180, MEXU, UC). Qro (Zamudio 6148, IEB, MEXU). SLP (Sa´nchez 460, MEXU). Tam (Stanford et al. 2012, DS, GH, NY). Ver (Va´zquez T. 00219, XAL).

6 4 . L O M A R I O P SI S Lomariopsis Fe´e, Me´m. Foug. 2: 10, 66. 1845. Lectotype (chosen by Holttum, Gard. Bull. Straits Settlem., ser. 3, 5: 264. 1932): Lomariopsis cochinchinensis Fe´e. Hemiepiphytic; rhizomes long-creeping, dorsiventral, climbing, scaly; fronds holodimorphic; stipes scaly; sterile blades 1pinnate, terminal pinna conform (ours) or abortive with articulate lateral pinna acting as false terminal pinna; pinnae articulate; rachises alate or not; juvenile fronds with more deeply toothed pinnae (rachises more broadly alate); veins free, simple to forking; fertile blades much constricted, laminar tissue greatly reduced, pinnae linear, articulate, stalked or not; sori acrostichoid, exindusiate; spores bilateral, with broadly and irregularly winged, long-spinulose perispore; x⫽41 (lower nos. known in some African spp.). Lomariopsis is a pantropical genus of nearly 40 species, with about 15 in the American tropics (Moran, 2000). The genus is uncommon in Mexico, where it is found primarily in lowland Atlantic slopes. Three Mexican species are known, and a fourth, L. japurensis (Mart.) J. Sm., is to be expected, as it occurs from Guatemala and Belize to Bolivia, the Guianas, and Brazil. Lomariopsis japurensis differs from congeners in Mexico by the broader sterile pinnae, 3.5–6 cm wide, the generally fewer pinna

LOMARIOPSIS RECURVATA

pairs (5–14), and the ferrugineous rhizome apex and stipe base scales. Affinities of Lomariopsis are generally with the dryopteroid ferns, but some systematists (e.g., Kramer in Kubitzki, 1990; Moran in Davidse et al., 1995; Moran, 2000) have treated Lomariopsis in a family Lomariopsidaceae, along with Elaphoglossum and Bolbitis. Unpublished molecular data indicate that such a family is polyphyletic, and that the affinities of Lomariopsis may instead be with Cyclopeltis; the relationships of Elaphoglossum, Bolbitis, and other genera often treated in Lomariopsidaceae appear to lie elsewhere in the dryopteroid assemblage (Cranfill, pers. comm.).

369

Lomariopsis is distinct by its hemiepiphytic habit, dimorphic fronds, free veins, and acrostichoid sporangia. Plants begin life on the ground and climb trees, where greater light intensity towards the canopy seems to trigger the production of fertile fronds. References Holttum, R. E. 1940. New species of Lomariopsis. Kew Bull. 1939: 613–628; Moran, R. C. 2000. Monograph of the neotropical species of Lomariopsis (Lomariopsidaceae). Brittonia 52: 55–111; Underwood, L. M. 1906. American ferns. VII. A. The American species of Stenochlaena. Bull. Torrey Bot. Club 33: 591– 603.

Key to the Mexican Species of Lomariopsis 1. Pinnae 20–35 pairs; sterile pinnae truncate at their bases, 4–10 ⫻ 1–1.8 cm; fertile pinnae 4–6 cm long; rachises conspicuously alate; proximal several pairs of pinnae reduced to 1⁄3 or less the length of longest pinnae; stipes 2–7 cm long, 1⁄10–1⁄5 of frond length. ...................................................................................................................................................................................................................... 3. L. vestita. 1. Pinnae 12–20 pairs; sterile pinnae rounded to cuneate at their bases, 6–15 ⫻ 1.5–2.7 cm; fertile pinnae 10–15 cm long; rachises nonalate or only slightly so; proximal pinnae not reduced or reduced to only 2⁄3 of length of longest pinnae; stipes (5–)10–30 cm long, 1⁄4–1⁄2 of frond length. 2. Sterile pinnae rounded to broadly cuneate at their bases, 6–10 ⫻ 1.5–2.5 cm, proximal 1–3 pinnae somewhat reduced. ...........1. L. mexicana. 2. Sterile pinnae cuneate at their bases, 8–15 ⫻ 2.2–2.7 cm, proximal pinna longest or nearly so. ................................................... 2. L. recurvata.

1. LOMARIOPSIS MEXICANA Lomariopsis mexicana Holttum, Kew Bull. 1939: 617. 1940. Type. Mexico. [Hidalgo:] Patrero, Hahn 56 (P!, frag. NY!). Figs. 175A–C. Rhizome scales stramineous to pale, lustrous, margins ciliate; stipes 5–15 cm long, 1⁄4–1⁄3 the frond length, stramineous to brown, scaly; sterile blades lanceolate, 30–60 ⫻ 12–20 cm, proximal 1–3 pinna pairs somewhat reduced (to ca. 2⁄3 of longest pinnae); rachises slightly alate; pinnae 12–15 pairs, 6–10 ⫻ 1.5–2.5 cm, rounded to broadly cuneate at bases, apices abruptly acuminate, margins entire to crenulate; fertile fronds with 12–15 pinna pairs, pinnae 12–13 ⫻ 0.3–0.4 cm.

UC; Mickel 6395, ENCB, NY, UC). Tab (Rovirosa 469, NY). Ver (Sohmer 9460, MEXU; Va´zquez T. 2561, CHAPA, NY).

This taxon, first mentioned by Underwood (1906) and named by Holttum (1940), is somewhat intermediate between L. vestita and L. recurvata, and Smith (1981) postulated that this poorly known species might be a hybrid. In characters of both the sterile and fertile blades it is intermediate in most characteristics (e.g., pinna base, sterile blade base, dimensions of fertile pinnae). However, the spores of the type fragment and other specimens examined (Cruden 1114, Va´zquez T. 2561) appear well formed and are highly ornamented and cristate (cf. R. M. Tryon & A. F. Tryon, 1982: 611). Moran (in Davidse et al., 1995: 284; 2000) synonymized L. mexicana under L. recurvata, but to us they seem adequately distinct. Because of its morphological intermediacy between L. recurvata and L. vestita, the possibility that it was formed through hybridization should be investigated.

2. LOMARIOPSIS RECURVATA Lomariopsis recurvata Fe´e, Me´m. Foug. 2: 68, pl. 28. 1845. Fig. 175F. Stenochlaena recurvata (Fe´e) Underw., Bull. Torrey Bot. Club 33: 600. 1906. Type. Mexico. Tabasco: Teapa, Linden s.n. (P!, photo NY!; isotypes P! photo US!, frag. B!, NY!).

Distribution. Hemiepiphytic in wet forest; 100–750 m. Mexico. Selected Specimens Examined. Chis (Breedlove 22473, 34075, DS; Palacios-Rios 2819, CHAPA, IEB). Hgo (Hahn 56, P). Oax (Cruden 1114,

Rhizome scales stramineous to pale, lustrous, margins ciliate; stipes (9–)11–30 cm long, 1⁄3–1⁄2 the frond length, stramineous to brown, scaly; sterile blades lanceolate, 30–100 ⫻ 15–24 cm, proximal pinnae longest or nearly so; rachises not alate or obscurely so; pinnae 12–20 pairs, (8–)9–15 ⫻ 2.2–2.7 cm, sessile, broadest in middle, narrowly to broadly cuneate at base, apices acumi-

370

LONCHITIS

nate, margins subentire to denticulate; fertile fronds with 14–20 pinna pairs, pinnae 10–15 ⫻ 0.2–0.3 cm.

Distribution. Hemiepiphytic in wet lowland forests; 50–1250 m. Mexico; Guat, Bel, Hond, Nic, CR, Pan; Col, Ec. Distribution. Hemiepiphytic in wet forests; 100–1000 m. Mexico; Guat, Bel, Hond. Selected Specimens Examined. Chis (Breedlove 21675, DS, NY, 35427, DS, MO, aberrant in having more widely placed veins; Martı´nez S. 10486, IEB, 24930, 24960, 26131, MEXU). Oax (Herna´ndez G. 1740, CHAPA, 1752, CHAPA, NY; Mickel 7171, ENCB, NY, UC, US). Tab (Cowan 3934, CHAPA, ENCB, UC). Ver (Riba 793, 824, 849, XAL; Rzedowski 20373, ENCB, NY).

The fewer pinna pairs, which are longer, broader, cuneate at their bases, and not reduced in size toward the frond bases, distinguish this species from L. vestita. Lomariopsis sorbifolium (L.) Fe´e of the Lesser Antilles, Puerto Rico, and Hispaniola, which also has 10–15 pinna pairs with cuneate bases, differs in having pinnae 4–8 cm long, rhizome scales pale brown to reddish brown, and entire, stalked fertile pinnae and winged rachises.

Selected Specimens Examined. Chis (Breedlove 21610, 38078, DS; Mell 2035, NY, US; Purpus 6770, UC, US, 6873, NY, PH, UC, US). Oax (Herna´ndez G. 1223, CHAPA; Wendt 3822, CHAPA, ENCB, NY, UC). Tab (Cowan 2073, CAS; Rovirosa 468, 469, PH).

The greatly reduced proximal pinna pairs, great number of pinnae, short stipes, and narrow, short pinnae with truncate to subcordate bases characterize this species. Lomariopsis fendleri D. C. Eaton (Mem. Amer. Acad. Arts, ser. 2, 8: 195. 1860), based on Fendler 335 (syntypes YU!, 3 sheets, collected on two dates; isosyntype K, frag. NY!) from Edo. Vargas, Venezuela, was considered by Tryon and Tryon (1982: 610) and R. Tryon and Stolze (1991) to be an earlier name for this species, but Moran (in Davidse et al., 1995: 284) indicated that the name L. fendleri should be restricted to plants occurring from Costa Rica, Trinidad, and Venezuela to Bolivia. It differs from L. vestita in having broader blades and proximal pinnae little or not reduced.

3. LOMARIOPSIS VESTITA Lomariopsis vestita E. Fourn., Bull. Soc. Bot. France 19: 250. 1872. Figs. 175D, E. Stenochlaena vestita (E. Fourn.) Underw., Bull. Torrey Bot. Club 33: 600. 1906. Type. Nicaragua. Chontales: Le´vy 476 (P!–4 sheets; isotype K! frag. NY!). Acrostichum pittieri Christ, Bull. Soc. Roy. Bot. Belgique 35(1): 243. 1896. Stenochlaena pittieri (Christ) Diels in Engler & Prantl, Nat. Pflanzenfam. 1(4): 251. 1899. Type. Costa Rica. Pittier 6931 (isotype NY!).

Rhizome scales stramineous to pale, dull, margins ciliate; stipes 2–7 cm long, 1⁄10–1⁄5 the frond length, stramineous to brown, scaly; sterile blades lanceolate, 30–60 ⫻ 8–20 cm, proximal several pairs of pinnae reduced to 1⁄3 or less length of longest pinnae; rachises conspicuously alate; pinnae 20–30 pairs, 4–10 ⫻ 1–1.8 cm, sessile or short-stalked (1 mm long), truncate to subcordate at bases, apices acuminate, margins subentire to denticulate; fertile fronds with 20–35 pinna pairs, pinnae 4–6 ⫻ 0.2–0.3 cm.

Excluded Species Lomaria ? juglandifolia C. Presl, Rel. Haenk. 52. 1825. Type. Said to be from Mexico, but later (Epimel. Bot. 164. 1849 [1851]) emended to Luzon (Philippines). According to Holttum (Novit. Bot. Delect. Seminum Horti Bot. Univ. Carol. Prag. 1969: 14), this name is a synonym of Stenochlaena palustris (Burm.) Bedd. Lomariopsis latiuscula (Maxon) Holttum was attributed to Mexico by A. F. Tryon and R. M. Tryon (1982: 611) on the basis of Dressler & Jones 142 (GH), from Veracruz. This specimen is L. recurvata. Lomariopsis latiuscula is a species of southern Central America (Nic, CR, Pan) and probably does not occur in Mexico. It was treated as a synonym of L. maxonii (Underw.) Holttum by Moran (in Davidse et al., 1995: 284; Moran, 2000).

65. LONCHITIS Lonchitis L., Sp. Pl. 2: 1078. 1753. Lectotype (first chosen by Brongniant in Bory, Dict. Class. Hist. Nat. 9: 490. 1825): Lonchitis hirsuta L. See Lellinger, Taxon 26: 578–580. 1977, for discussion of typification.

LOPHOSORIA

371

Anisosorus Trevis. ex Maxon, Pteridophyt. Porto Rico 429. 1926. Type: Pteris laciniata Willd. [⫽ Anisosorus hirsutus (L.) Underw. & Maxon] ⬅ Lonchitis hirsuta L.

Terrestrial; rhizomes creeping, fleshy, stout, with pale hairs; fronds medium to large, monomorphic; stipes stout, fleshy, hairy, green, terete abaxially, ridged adaxially; blades ovate, bipinnatepinnatifid, ridges continuous with those of costae; laminae membranaceous, herbaceous, with numerous septate hairs (in ours) or glabrous; veins free (in ours) to casually anastomosing; sori marginal on sides of segments, 2 per segment, lacking paraphyses; indusia submarginal from inner edge of marginal flange, broad, reflexed over sorus; spores tetrahedral-globose; x⫽50. Lonchitis has two species, one in Africa and one in America. The Amercian one is widespread in wet forests at middle elevations. The species is sometimes placed in Anisosorus due to confusion with typification of Lonchitis and Blotiella (Lellinger, 1977). Lonchitis is distinct in its hairy rhizomes, stipes, and blades, and the membranous blades with free veins. References Lellinger, D. B. 1977. The identity of Lonchitis aurita and the generic names Anisosorus and Lonchitis. Taxon 26: 578–580; Tryon, R. M. 1962. Taxonomic fern notes. III. The genera Lonchitis and Blotiella. Contr. Gray Herb. 191: 91–107.

Distribution. Wet montane forests; 350–1700 m. Mexico; Guat, Hond, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Sur, Ec, Peru, se Braz, Bol. Selected Specimens Examined. Chis (Breedlove 42100, CAS, 56489, CAS, NY; Mu¨nch 150, DS, US). Oax (Hallberg 1474, NY; Martı´nez S. 2801 MEXU; Mickel 4002, 5233, MEXU, NY). Pue (Riba et al. 316, MEXU). Ver (Purpus 2928, NY; Ventura A. 14881, MEXU).

6 6 . L O P H O SO R I A 1. LONCHITIS HIRSUTA Lonchitis hirsuta L., Sp. Pl. 2: 1078. 1753.

Figs. 176A–C.

Anisosorus hirsutus (L.) Maxon, Pteridophyt. Porto Rico 429. 1926. Lectotype (chosen by Brongniant in Bory, Dict. Class. Hist. Nat. 9: 490. 1825). Plumier, Traite´ Foug. Ame´r., pl. 20, 1705, illustrating a plant from Morne Rouge, Martinique. Pteris laciniata Willd., Sp. Pl., ed. 4, 5(1): 397. 1810. Type. “India orientale.” Flu¨gge´ s.n. (B-Willd. 20013!).

Rhizomes creeping, 10 mm diam.; rhizome hairs light brown, septate, 1–3 mm long; fronds 1–3 m long, subdistant; stipes ca. 1⁄3 the frond length, light green to stramineous, densely hairy, soft; blades deltate or ovate-deltate, hairy throughout; pinnae 12–16 pairs, alternate to subopposite, stalked, unequally deltate, segments oblong or roundish, 5–9 mm long and wide, lobed to entire; veins free, ending short of margins on sterile segments; sori submarginal, scarious, subentire, 1–10 ⫻ 1 mm; 2n⫽200 (Jam).

Lophosoria C. Presl, Gefa¨ssbu¨ndel Farrn 36. 1847 (preprint from Abh. Ko¨nigl. Bo¨hm. Ges. Wiss., ser. 5, 5: 344. 1848.). Lectotype (first chosen by J. Smith?, Hist. Fil. 251. 1875): Lophosoria pruinata (Sw.) C. Presl [⬅ Polypodium pruinatum Sw., nom superfl. for Polypodium quadripinnatum J. F. Gmel., 1792, both based on Polypodium glaucum Sw., 1788, non Thunb., 1784] ⬅ Lophosoria quadripinnata (J. F. Gmel.) C. Chr.]. Trichosorus Liebm., Mexic. Bregn. 281 (reprint 129). 1849. Type: Alsophila pruinata (Sw.) Kunze (combination Trichosorus pruinatus not made) [⬅ Polypodium pruinatum Sw.] ⫽ Lophosoria quadripinnata (J. F. Gmel.) C. Chr.].

Terrestrial; rhizomes creeping to ascending or erect, trunks short, with dense, long, golden hairs; fronds large, monomorphic; stipes stout, hairy, brown to stramineous, lacking spines; blades deltate-ovate, tripinnate-pinnatifid, anadromous, coriaceous or subcoriaceous, glaucous abaxially, axes and veins with golden hairs; veins free or casually areolate; sori round, medial, abaxial, exindusiate, with abundant paraphyses; spores with equatorial flange; x⫽65. Lophosoria is a genus of three species, one of which is widespread and common in high elevation wet forests. The genus resembles the dicksonioid tree ferns in stomata, hairs, stipe anatomy, and chromosome number (x⫽65). It is distinct by its stout, hairy rhizomes, large fronds, blades glaucous abaxially, and exindusiate abaxial sori. Molecular data (Wolf et al., 1999) have shown Lophosoria to be sister to Dicksonia. Reference Wolf, P. G., S. D. Sipes, M. R. White, M. L. Martines, K. M. Pryer, A. R. Smith & K. Veda. 1999. Phylogenetic relationships of the enigmatic fern families

372

LOXOGRAMME

Hymenophyllopsidaceae and Lophosoriaceae: evidence from rbcL nucleotide sequences. Pl. Syst. Evol. 219: 263–270.

1. LOPHOSORIA QUADRIPINNATA Lophosoria quadripinnata (J. F. Gmel.) C. Chr. in Skottsberg, Nat. Hist. Juan Fernandez 1: 16. 1920. Figs. 177A–C. Polypodium glaucum Sw., Prodr. 134. 1788, hom. illeg, non Thunb., 1784. Polypodium quadripinnatum J. F. Gmel., Syst. Nat. 2(2): 1314. 1791 [1792]. Polypodium pruinatum Sw., J. Bot. (Schrader) 1800(2): 29 1801, nom. superfl. Lophosoria pruinata (Sw.) C. Presl, Gefa¨ssbu¨ndel Farrn 36. 1847. Alsophila quadripinnata (J. F. Gmel.) C. Chr., Index Filic. 47. 1905. Type. Jamaica, Swartz s.n. (S?). Trichosorus glaucescens Liebm., Mexic. Bregn. 283 (reprint 131). 1849. Lectotype (chosen by Smith, 1981: 144). Mexico. Oaxaca: “Inter Talea and Hac. de Sta. Gertrudes,” Liebmann s.n. [Fl. Me´x. 877] (C!). Trichosorus densus Liebm., Mexic. Bregn. 284 (reprint 132). 1849. Lectotype (chosen by Smith, 1981: 144) Mexico. Puebla: “Pr. Chinautla,” Liebmann s.n. [Fl. Mex. 881] (C!, photo BR!, photo US, frag. NY!). Trichosorus frigidus Liebm., Mexic. Bregn. 284 (reprint 132). 1849. Lectotype (chosen by Smith, 1981: 144). Mexico. Oaxaca: “Cerro de Sempoaltepec,” Liebmann s.n. [Pl. Me´x 884] (C!, photo US; isotype US!); isotype Liebmann s.n. [Pl. Mex. 507] (C!, photo US).

Stems creeping, densely clothed with long, golden hairs; fronds 2–4 m long; stipes mostly stramineous, nearly black and very hairy at bases with golden hairs to 2 cm; blades tripinnatepinnatifid to quadripinnate, subcoriaceous; pinnae subsessile to petiolulate, petiolules to 13 cm long, pinnules also stalked to 2 mm; adaxial surfaces nearly glabrous, lustrous, hairs limited to axes and major veins; abaxial surfaces with orange-tan hairs 1–2 mm long, especially on midveins, to nearly glabrous; 2n⫽130 (Jam).

6 7 . L O X O GR A M M E Loxogramme (Blume) C. Presl, Suppl. Tent. Pterid. 214, t. 9, f. 8. 1836. Antrophyum Kaulf. sect. Loxogramme Blume, Enum. Pl. Javae 1: 73. 1828. Lectotype (first chosen by J. Sm.?, Hist. Fil. 156. 1875): Loxogramme lanceolata (Sw.) C. Presl [⬅ Grammitis lanceolata Sw.].

Epiphytic and epipetric; rhizomes long-creeping, slender, scaly, bearing densely matted roots; fronds monomorphic (or some dimorphic in Old World), small, distant to clumped, pendent; stipes short to nearly absent, not articulate; blades simple, entire, glabrous, fleshy; veins immersed, obscure, netted with included veinlets; sori elongate, oblique to midrib, superficial or in slight grooves; indusia absent; paraphyses absent (in ours) or present; spores green, bilateral or tetrahedral, perispore papillate to shallowly tuberculate; x⫽35, 36. There are about 30 species of Loxogramme, most of eastern and southeastern Asia and adjacent Pacific islands, two in Africa and on offshore islands, and only one in Mesoamerica. Copeland (1947) suggested a grammitid alliance for Loxogramme, but Price (pers. comm.) believed it to be a distinct group that is coordinate with polypods and grammitids, perhaps more closely allied to Polypodiaceae s.str. Recent molecular analyses show that Loxogramme and the Australasian genera Anarthropteris (monotypic, New Zealand) and Dictymia (two or three species, eastern Australia and Melanesia) form the basalmost clade within Polypodiaceae, and this clade has no close alliance with Grammitidaceae (Schneider et al., ms. in prep.). The sole Mexican species is rare, growing with, and sometimes confused with, Scoliosorus ensiforme. Loxogramme is characterized by anastomosing veins, clathrate rhizome scales, entire, glabrescent leaves and green spores.

1. LOXOGRAMME MEXICANA Loxogramme mexicana (Fe´e) C. Chr., Index Filic., Suppl. 3: 25. 1934. Figs. 178A–E.

Distribution. Wet upper montane forests and cloud forests on slopes, oak woods, with pine, beech, alder; 1050–3050 m. Mexico; Guat, Hond, Salv, Nic, CR, Pan; Gr Ant; Col, Ven, Trin, Ec, Peru, Bol, s Chile, J Fdez, Arg. Selected Specimens Examined. Chis (Breedlove 15314, DS, US; Mickel 1253, ISC, NY, US). DF (Riba et al. s.n., in 1964, FCME). Gro (Rzedowski 16424, ENCB, NY). Hgo (Vera C. 7, CAS, ENCB, MEXU). Oax (Rzedowski 21035, MICH, NY, US). Pue (Orcutt 3997, US). Qro (Rzedowski 42871, CHAPA, ENCB, IEB, MICH, UAMIZ, XAL). Tab (Zenteno 19, UAMIZ). Ver (Taylor & Nee 258, F, NY, XAL).

Selliguea mexicana Fe´e, Me´m. Foug. 7: 43, t. 10, f. 4. 1857 (nom. nud. in Me´m. Foug. 5: 177. 1852). Gymnogramma mexicana (Fe´e) Baker in Hooker & Baker, Syn. Fil. 387. 1868. Type. Mexico. Oaxaca: Sierra San Pedro Nolasco, Talea, etc., Jungersen [Ju¨rgensen] 945 (P!, frag. NY!; isotypes BM!, K!). Grammitis salvinii Hook., Sec. Cent. Ferns, t. 71. 1860 [1861]. Gymnogramma salvinii (Hook.) Hook., Sp. Fil. 5: 157. 1864. Loxogramme salvinii (Hook.) Maxon, Contr. U.S. Natl. Herb. 13: 17. 1909. Type. Guatemala. Verapaz, Salvin s.n. (K!).

Rhizomes long-creeping, 0.5–1.4 mm diam., with heavy, straight brown hairs and black clathrate scales, the scales linearlanceolate, long-acuminate, 2–5(–8) ⫻ 1 mm; fronds mostly 10– 25 ⫻ 1–3.5 cm, distant, often 1.5–3 cm apart; stipes absent; blades fleshy, simple, narrowly oblanceolate, gradually tapered to frond base, apex long-acuminate, margins entire; veins obscure, midvein visible below as a lighter streak, above by a slight ridge; sori sunken in grooves, mostly 7–12 mm long, as 4–8 streaks in a

LOXOSCAPHE

row (one row on each side of midrib), positioned submedially between midrib and margins, more or less parallel to midrib but slightly oblique to it; spores bilateral.

373

chromosome base number (x⫽35), unknown in Asplenium. However, Gastony and Johnson (2001) have recently shown that Loxoscaphe thecifera is “robustly and surprisingly nested” within a clade containing a number of Old World (African and Asian), simple-bladed species allied to the Bird’s nest fern, Asplenium nidus L. This group has sometimes been treated as the segregate genus Neottopteris. Reference Gastony, G. J. & W. P. Johnson. 2001. Phylogenetic placements of Loxoscaphe thecifera (Aspleniaceae) and Actiniopteris radiata (Pteridaceae) based on analysis of rbcL nucleotide sequences. Amer. Fern J. 91: 197–213.

1. LOXOSCAPHE THECIFERA Loxoscaphe thecifera (Kunth) T. Moore, Index Fil. 302. 1861. Figs. 179A–C. Distribution. Epiphytic or on logs or at base of trees, in wet montane forest of Atlantic slope, cloud forests, pine-oak forests, montane cloud forests; 800–2100 m. Mexico; Guat, Salv, Nic, CR, Pan. Selected Specimens Examined. Chis (Breedlove 22844, DS, MEXU, MICH, MO, NY, 31970, DS, MEXU, 35201, DS, MEXU, MO, 56609, CAS, ENCB; Martı´nez S. 20667, MEXU, XAL). Jal (McVaugh 21441, CAS, IEB, MEXU, MO, MICH). Oax (Garcı´a Mendoza 1532, MEXU; Maya J. 4042, NY; Mickel 949b, NY, US). Ver (Copeland herb. 156, B, BM, MEXU, MICH, UC, US).

In Oaxaca, this species was found mixed with Scoliosorus ensiformis (Mickel 949a, NY), which it closely resembles in the blade shape and texture, clathrate scales, and oblique sori, but it is distinct in the long-creeping rhizomes and straighter, fewer, non-anastomosing sori. Under dry conditions, L. mexicana becomes flaccid and limp, but revives remarkably with moisture.

Davallia thecifera Kunth, Nov. Gen. Sp. 1: 23. 1816 [“Theciferum”]. Darea thecifera (Kunth) E. Fourn., Mexic. Pl. 1: 132. 1872. Type. Venezuela. “In monte Saraquen,” Humboldt s.n. (P-Humb.!). Davallia lindenii Hook., Sp. Fil. 1: 193, t. 56B. 1846. Loxoscaphe lindenii (Hook.) T. Moore, Hooker’s J. Bot. Kew. Gard. Misc. 5: 227. 1853. Syntypes. Venezuela. Caracas, Linden 70 (K; isosyntype BM, photo NY!, FI), Hartweg 1507 (K; isosyntype BM, photo NY!, FI), Funck 648 (K); Brazil. Organ Mts., Gardner 200 (K).

Rhizomes suberect; rhizome scales clathrate, 0.8–1 ⫻ 0.5–0.8 mm, ovate-lanceolate, denticulate; fronds clumped, 4–15 ⫻ 2–5 cm, 2–3-pinnate, linear-lanceolate, apices pinnatifid; stipes 2–10 cm long, 1⁄2–2⁄3 the frond length, stramineous, with scattered scales similar to rhizome scales; pinnae 4–6 pairs, pinnules fleshy; sori in pouches near margin, indusia 1 mm wide; 2n⫽140 (CR).

68 . L O X O S C A PH E Loxoscaphe T. Moore, Hooker’s J. Bot. Kew Gard. Misc. 5: 227. 1853. Type. Loxoscaphe concinna (Schrad.) T. Moore [⫽ Loxoscaphe thecifera (Kunth) T. Moore]. Epiphytic; rhizomes short-creeping to suberect, with clathrate scales; fronds relatively small, clumped, monomorphic; stipes scaly at bases; blades finely dissected, 2–4 times pinnate; segments slender, linear, fleshy; veins free; sori near margin protected by pouch-like indusia, facing outward, similar to those of Davallia; spores bilateral, perispore winged; x⫽35. Loxoscaphe comprises four or five species, one in the neotropics, two in Africa (one usually considered only varietally different from the neotropical species), and two in the South Pacific and Indonesia. Although it is most closely allied to and sometimes placed in synonymy under Asplenium, it is maintained as separate here because of the peculiar pocket-like sori (superficially similar to those of Odontosoria) and the apparent different

Distribution. Epiphytic in montane rain forests; 1350–1900. Mexico; Nic, CR, Pan; Hisp; Col, Ven, Ec, Peru, Bol; Africa, Madagascar. Also cited from Honduras and Brazil by Adams (in Davidse et al., 1995: 322). Specimens Examined. Chis (Breedlove 52078, CAS). Ver (Bourgeau 3611, cited in Fournier, 1872: 132).

The African variant, L. concinna, appears conspecific with material from the neotropics; it is often recognized at varietal rank [as var. concinna (Schrad.) Schelpe], but even that level of recognition is questionable.

374

LYCOPODIELLA

69. L YC O PO DI E L L A Lycopodiella Holub, Preslia 36: 22. 1964. Type: Lycopodiella inundata (L.) Holub [⬅ Lycopodium inundatum L.]. Palhinhaea Vasc. & Franco, Bol. Soc. Brot., ser. 2, 41: 24. 1967. Type: Palhinhaea cernua (L.) Vasc. & Franco [⬅ Lycopodium cernuum L.] ⬅ Lycopodiella cernua (L.) Pic. Serm. Pseudolycopodiella Holub, Folia Geobot. Phytotax. 18: 441. 1983. Type: Pseudolycopodiella caroliniana (L.) Holub [⬅ Lycopodium carolinianum L.] ⬅ Lycopodiella caroliniana (L.) Pic. Serm.

Terrestrial in boggy areas; stems short-creeping on ground or stolon-like, rooting where touching the ground, with erect shoots, these unbranched, to 30 cm tall (rare species) or highly branched, 30–150 cm tall (common species); leaves not in distinct ranks, ascending or spreading, monomorphic, linear to linear-

lanceolate, entire; gemmiferous branchlets absent; strobili terminal on erect stems and indistinctly differentiated, or nodding on branches and distinct; sporangia globose to reniform, axillary or on the base of sporophylls; spores rugulate; gametophytes photosynthetic, on soil surface, pin-cushion-shaped or tuber-shaped; x⫽35, 39, 78, 104 (USA). Lycopodiella contains about 40 species, of which three occur in Mexico. The genus as construed here is divided by some authors into three genera: Lycopodiella, Palhinhaea, and Pseudolycopodiella. The three Mexican Lycopodiellas are diverse, but details of sporangia, spores, and gametophytes suggest that they are closely allied. For discussion of relationships, see references under Huperzia and Lycopodium.

Key to the Mexican Species of Lycopodiella 1. Plants erect, dendroid, 30–150 cm tall; strobili sessile, pendent; leaves 2–4 mm long, each without a bristle-tip. ............................... 3. L. cernua. 1. Plants creeping; strobili erect, borne on simple erect branches (peduncles). 2. Peduncles with abundant imbricate leaves similar to those of horizontal stems; horizontal stems with leaves all alike and not lying on the substrate. .......................................................................................................................................................................... 1. L. alopecuroides. 2. Peduncles nearly naked, with scattered small leaves; horizontal stems appearing flattened, with lateral leaves longer than the median leaves and more or less appressed to the substrate. .............................................................................................................. 2. L. caroliniana.

1. LYCOPODIELLA ALOPECUROIDES

Specimens Examined. Chis (Breedlove 15165, CAS, US).

Lycopodiella alopecuroides (L.) Cranfill, Amer. Fern J. 71: 97. 1981. Figs. 180E–H.

2. LYCOPODIELLA CAROLINIANA

Lycopodium alopecuroides L., Sp. Pl. 2: 1102. 1753. Syntype. U.S.A. Virginia: Kalm s.n., LINN-1257.7.

Lycopodiella caroliniana (L.) Pic. Serm., Webbia 23: 165. 1968. Figs. 180A–D.

Horizontal branches strongly arching, rooting at tips; stems (excluding leaves) 2–4 mm diam.; leaves monomorphic, spreading to ascending, 5–7 mm long, marginal teeth 1–7 per side; peduncles 6–30 cm long with abundant imbricate leaves similar to those of horizontal stems, leaves spreading to ascending, 6–7 mm long, marginal teeth 1–10 per side; strobili erect, 20–60 ⫻ 12–20 mm, 1⁄7–1⁄3 the branch length; sporophylls wide-spreading, 6–7 mm long, marginal teeth 1–5 per side in proximal 1⁄2; 2n⫽156 (USA).

Lycopodium carolinianum L., Sp. Pl. 2: 1104. 1753. Syntype. Dillenius s.n., Hist. Musc., t. 62, f. 6. Lycopodium meridionale Underw. & F. E. Lloyd, Bull. Torrey Bot. Club 33: 121. 1906. Lycopodium carolinianum L. var. meridionale (Underw. & F. E. Lloyd) Nessel, Ba¨rlappgewa¨chse 271. 1939. Pseudolycopodiella meridionalis (Underw. & F. E. Lloyd) Holub, Folia Geobot. Phytotax. 18: 442. 1983. Lycopodiella caroliniana (L.) Pic. Serm. var. meridionalis (Underw. & F. E. Lloyd) B. Øllg. & P. G. Windisch, Bradea 5: 27. 1987. Type. Puerto Rico. Luquillo Mts, Wilson 94 (NY!).

Distribution. Bogs, marshes, ditches; 2700 m. E USA; Mexico; Guat, Salv; Cuba; Col, Ven, Ec, Peru, Braz, Bol, Parag, Uru.

Horizontal branches appressed to the soil, rooting along their length; stems (excl. leaves) ca. 1 mm diam.; leaves somewhat dimorphic, the upperside ones linear, more or less appressed, entire, 3–4 mm long, the lateral ones somewhat appressed to the substrate, the underside ones lanceolate, spreading, 4–5 mm long, entire; peduncles 20–30 cm long, with appressed leaves, these distant in pseudowhorls of 3, 3–4 mm long, entire; strobili erect, 2–6 cm ⫻ 5–8 mm (incl. spreading sporophylls), sporophylls 1⁄6–1⁄8 the branch length, lanceolate to linear-lanceolate, entire, ca. 3 mm long; 2n⫽70, 140 (USA), 78 (Europe).

LYCOPODIUM

Distribution. Wet meadows, ditches; 0–50 m. E USA; Mexico; Guat, Bel, Hond, Salv, Nic, Pan; Gr & L Ant; Col, Ven, Guy, Sur, Fr Gui, Ec, Peru. Specimens Examined. Tab (Cowan et al. 2233, MEXU, NY; Orozco, S. et al. 2233, MO).

This species is distinct by the flattened terrestrial stems and sparse leaves of the peduncles. Material of this species from tropical America is usually placed in var. meridionalis.

3. LYCOPODIELLA CERNUA Lycopodiella cernua (L.) Pic. Serm., Webbia 23: 166. 1968. Figs. 180J–L. Lycopodium cernuum L., Sp. Pl. 2: 1103. 1753. Lepidotis cernua (L.) P.Beauv., Prodr. Aethe´ogam. 108. 1805. Palhinhaea cernua (L.) Vasc. & Franco, Bol. Soc. Brot., ser. 2, 41: 25. 1967. Type. Uncertain, “habitat in Indis.” LINN 1257.13–1257.17 (frag. NY!) plus several cited illustrations.

Plants 30–50 cm tall; main stems usually erect (sometimes with horizontal arching stolon-like branches from the bases), 2–3(–5) mm diam., stramineous, with leaves indistinctly spirally arranged, spreading to ascending, linear, 3–5 ⫻ 0.25 mm; lateral branches spreading, pinnately to dichotomously branched, with leaves abundant, spreading-falcate, linear, 2–4 ⫻ 0.2 mm, indistinctly spirally arranged, with sparse hairs 0.2–0.5 mm along decurrent leaf bases; strobili sessile, pendent at the ends of branchlets, 0.5–1 cm ⫻ 2–3 mm; sporophylls appressed, tightly overlapping, deltate, acuminate, margins erose-fimbriate; sporangia globose, valves unequal; 2n⫽208, 312 (India), ca. 320 (Jam).

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Distribution. Terrestrial, on roadbanks in cultivated areas and other disturbed places; common at lower to middle elevation wet forests, tropical montane forests, savannas, coffee plantations, oak and pine woods, montane rain forests; 100–2000 m. SE USA; Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol, Parag; Old World tropics and subtropics. Selected Specimens Examined. Chis (Matuda 3781, MEXU, NY, US). Gro (Tenorio L. et al. 1381, ENCB, MEXU). Hgo (Sharp 441787, MEXU, US). Jal (Quintero 6420, ENCB). Me´x (Hinton 9017, MO, NY). Mich (Dı´az & Zamudio 2700, ENCB, IEB, XAL). Nay (Tenorio L. et al. 17017, MEXU). Oax (Mexia 9143, MO, NY). Pue (Ventura 376, ENCB, NY). Tab (Cowan 3172, MO, NY). Ver (Purpus 2176, NY).

The Mexican material of L. cernua appears quite uniform though pantropically it forms a complex. It is abundant along roadsides and other open disturbed soil and is distinct by its upright stems with spreading branches and pendent strobili.

70. LYCOPODIUM Lycopodium L., Sp. Pl. 2: 1100. 1753. Lepidotis Mirbel in Lamarck & Mirbel. Hist. Nat. Ve´g. 3: 477; 4: 311. 1802, nom. superfl. for Lycopodium by lectotypification (chosen by Pichi Sermolli, Webbia 26: 145–149. 1971). Type: Lycopodium clavatum L. Diphasiastrum Holub, Preslia 47: 104. 1975. Type: Diphasiastrum complanatum (L.) Holub [⬅ Lycopodium complanatum L.]

Plants trailing on the ground; horizontal stems long-creeping with scattered upright shoots; leaves linear to linear-lanceolate, appressed, ascending, or spreading, in 6–8 ranks or scale-like reduced leaves in four ranks, monomorphic on round branches or dimorphic on flattened branches; leaf margins entire; gemmiferous branchlets absent; strobili single and sessile or multiple and pedunculate, peduncles leafy; sporangia reniform in axils of sporophylls aggregated in strobili; spores reticulate; gametophytes nonphotosynthetic, subterranean, flat and irregularly buttonshaped or carrot-shaped; x⫽23, 34 (USA). Lycopodium s.l., as used by most previous authors (e.g., Smith, 1981, and Mickel & Beitel, 1988) is here treated as comprising

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three genera: Lycopodium, Lycopodiella, and Huperzia, but has been broken into seven to 14 genera by some other authors (e.g., Wagner & Beitel, 1992, 1993). For further discussion of relationships, see references under Huperzia. References Øllgaard, B. 1983. Lycopodiaceae. In: Stolze, R., Ferns and fern allies of Guatemala, Part III. Fieldiana, Bot., n.s., 12: 20–44; Wagner, F. S. 1980. [Abstract] Chromosome behavior in three interspecific hybrids of Diphasiastrum (Lycopodiaceae). Bot. Soc. Amer. Misc. Ser. Publ. 158: 121–122; Wagner, W. H., Jr. & J. M. Beitel. 1992. Generic classification of modern North American Lycopodiaceae. Ann. Missouri Bot. Gard. 79: 676–686; Wagner, W. H., Jr. & J. M. Beitel. 1993. Lycopodiaceae. In: FNA Ed. Com, pp. 18–37; Wikstro¨m, N. & P. Kenrick. 2000. Relationships of Lycopodium and Lycopodiella based on combined plastid rbcL gene and trnL intron sequence data. Syst. Bot. 25: 495–510; Wilce, J. 1965. Section Complanata of the genus Lycopodium. Beih. Nova Hedwigia 19: i-x, 1-233; Wilce, J. 1972. Lycopod spores. I. General spore patterns and the generic segregates of Lycopodium. Amer. Fern J. 62: 65–79.

Key to the Mexican Species of Lycopodium 1. Branches terete; leaves spirally arranged throughout in many ranks; strobili 1–3 per long peduncle or peduncles absent. ................................................................................ 1. L. clavatum. 1. Branches flattened; branchlet leaves in 4 ranks; strobili 4– 12 per long peduncle. ......................................................... 2. L. thyoides.

1. LYCOPODIUM CLAVATUM Lycopodium clavatum L., Sp. Pl. 2: 1101. 1753. Type. Uncertain; LINN 1257.2 is this species, but according to Jackson (1912) was incorporated in LINN after 1755. Linnaeus cited several references. Figs. 181D, E. For additional synonymy, see Øllgaard (1983).

Stems prostrate, long-creeping, 1–3 mm diam., sparsely rooting; lateral branches terete, ascending to erect, 4–25 cm long, subdichotomously forked to pinnately branched; leaves appressed to patent, ascending or appressed, 4–8 ⫻ 0.3–0.9 mm, spirally arranged, each ending in a hair-tip 1–3 mm long; strobili cylindrical, 1–3 per peduncle, 2–9 cm ⫻ 3–5 mm, on erect branches at the end of sparsely leaved peduncles, peduncles simple to 3-times sympodially forked; sporophylls deltate-ovate, margins erose, hair-tips 1–2 mm; 2n⫽68 (Europe, USA).

Distribution. Terrestrial on banks and exposed slopes, cloud forests, montane rain forests or pine-oak forests, pine-fir forests; 650–3200 m. North temperate regions of North America; Mexico; Guat, Hond, Nic, CR, Pan; Gr & L Ant; Col, Ven, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol, Parag; n Europe, temperate and tropical regions of Asia and Pacific Basin. Selected Specimens Examined. Chis (Breedlove 52225, ENCB, NY; Ventura & Lo´pez 3962, ENCB, NY). Gro (Rzedowski & McVaugh 130, IEB, MEXU, NY). Hgo (Pringle 11885, US). Me´x (Boege 1811, MEXU). Mich (Carranza 1191, IEB). Oax (Hallberg 840, DS; Mickel 4935, NY). Pue (Arse`ne 1944, MEXU, NY, US). Qro (Carranza 1191, ENCB, IEB, MEXU). Ver (Nee 22468, NY).

A highly variable species or species complex, L. clavatum s.l. is in need of worldwide revision. Specimens from Mexico seem to exhibit two extremes: (1) Strobili on peduncles 9–16 cm long, peduncle leaves entire to slightly erose, each with scarious tip, 2–4 strobili per peduncle, each strobilus on a separate modified branch of main peduncle, branch leaves spreading to ascending, annual constrictions none or indistinct, 600–2100 m; and (2) strobili sessile or on peduncles to 4 cm long, peduncle leaves erose, each with a scarious tip and distal leaf margins, 1 strobilus per peduncle (rarely 2, then paired at base of strobili, no peduncle branch), branch leaves ascending to appressed, distinct annual constrictions, 2700–3000 m. Intermediate specimens occur with 1–4 strobili on the same specimens, usually with a short peduncle branch for each strobilus; peduncles 4–13 cm; branch leaves ascending to spreading or appressed. These occur at intermediate elevations of ca. 2500 m.

2. LYCOPODIUM THYOIDES Lycopodium thyoides Humb. & Bonpl. ex Willd., Sp. Pl., ed. 4, 5(1): 18. 1810. Figs. 181A–C. Diphasiastrum thyoides (Humb. & Bonpl. ex Willd.) Holub, Preslia 47: 108. 1975. Type. Venezuela. Silla de Caracas, Humboldt s.n. (B-Willd. 19352!, frag. NY; isotype P-Humb.!). Lycopodium complanatum L. var. validum Weath., Proc. Amer. Acad. Arts 45: 414. 1910. Syntypes. Mexico. Chiapas: Between San Cristo´bal Las Casas and Huista´n, Seler & Seler 2273 (GH!; isosyntype NY!), Ghiesbreght 600 (GH!– 2 sheets; isosyntype NY!–2 sheets); Oaxaca: Cerro San Felipe, Gonza´lez &

LYGODIUM Conzatti 889 (GH!); Veracruz: Regio´n d’Orizaba, Bourgeau 3159 (GH!; isosyntype US!); Hidalgo: Trinidad, Pringle 11856 (GH!). Wilce (1965: 157) called this an “incorrect” name; we are uncertain what this means. Under the rules of nomenclature, var. validum seems effectively and validly published.

Subterranean stems long-creeping, just under the ground level, ca. 1.2–2.5 mm diam., with erect aerial branches mostly 15–45 cm long, leaves scattered, linear-lanceolate, spirally arranged, ca. 2–4 mm long; erect aerial stems mostly monopodial with several flattened, flabellate, lateral branches 1.5–3 mm wide and 1–2 terminal fertile peduncles; seasonal growth constrictions evident on the flattened branches; branchlet leaves in 4 ranks, the lateral two pseudodecussate with slightly to widely spreading Ⳳ deltate tips, the upper and lower ranks appressed, needle-like, 1–2 mm long, the lower leaves less conspicuous than the upper ones, occasionally 3-ranked with lower rank missing, lateral rank losing pseudodecussate arrangement; fertile peduncles dichotomously branched 2–4 times resulting in 4–12 strobili; pedicels subtending strobili usually twice dichotomously branched; strobili cylindrical, 2–5 cm ⫻ 2.5–3 mm, sometimes with a caudate sterile tip to 8 mm long.

Distribution. Terrestrial in humid forests at upper elevations, matorral, cloud forests, pine-oak forests, pine forests; 1300– 3050 m. Mexico; Guat, Hond, Salv, CR, Pan; Col, Ven, Ec, Peru, Braz, Bol, n Arg. Selected Specimens Examined. Chis (Breedlove 19992, DS, US). Gro (Vela´zquez & Zlotnik s.n., 21 Jul 1990, FCME). Hgo (Gimate 798, CAS, ENCB; Pringle 11856, US). Oax (Anderson & Anderson 4716, MICH,

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NY; Camp 2727, GH, MICH, NY). Pue (Purpus 2691, US; Ventura A. 375, ENCB, NY). Ver (Ma´rquez R. 916, F, NY, XAL, UAMIZ).

This name is applied to all members of the L. complanatum complex found in continental Latin America (Antillean material is L. fawcettii Underw. & F. E. Lloyd). Wilce (1965) treated the complex in the rest of the world, but believed the Latin American material required further study. Lycopodium thyoides appears more irregular in growth characters than taxa north of Mexico.

7 1 . L Y GO D I U M Lygodium Sw., J. Bot. (Schrader) 1800(2): 106. 1801, nom. cons. Type: Lygodium scandens (L.) Sw. [⬅ Ophioglossum scandens L.] ⫽ Lygodium flexuosum (L.) Sw. For further synonymy, see Lellinger, Mem. New York Bot. Gard. 18: 5. 1969.

Plants with climbing, twining fronds; rhizomes shortcreeping, hairy; stipes and rachises wiry, stramineous to brown, rachises twining, of indefinite growth; pinna axes short with dormant apices and two lateral pinnules, these sometimes Ⳳ dichotomously forked or pinnately divided; pinnules usually palmately lobed, with free or netted veins; sporangia on narrow finger-like projections from the pinnules, in two rows on each projection; sporangia oblong with an apical annulus, each covered by a basally attached indusium; spores tetrahedral, yellow, verrucose or reticulate; x⫽29, 30. Lygodium comprises about 25 species, all but one of tropical or subtropical regions, three-fourths of them of the Old World. It is distinct by its twining fronds and monangial, indusiate sori. It has traditionally been placed with Anemia, Mohria, Schizaea, and Actinostachys in the Schizaeaceae, and this placement has been supported by molecular data (Hasebe et al., 1995; Pryer et al., 1995). References Duek, J. J. 1978. A taxonomic revision of Lygodium (Filicinae) in America. Repert. Spec. Nov. Regni Veg. 89: 411-423; Hanks, J. G. 1998. A monographic study of Lygodium Swartz (Pteridophyta: Lygodiaceae). Unpublished doctoral dissertation, City Univ. of New York; Lellinger, D. B. 1969. The botany of the Guayana Highland. Part VIII. Schizaeaceae (Filicales). Mem. New York Bot. Gard. 18: 2–11. 1969; Maxon, W. R. 1909. Schizaeaceae. N. Amer. Fl. 16: 31-52.

Key to the Mexican Species of Lygodium 1. Veins netted, each vein connecting with another after only 1 or 2 forkings from the pinnule midvein. ........................................ 1. L. heterodoxum. 1. Veins free. 2. Pinnules lobed at bases, distal ones mostly shorter than the proximal. ............................................................................................. 2. L. venustum. 2. Pinnules mostly not lobed at bases, all nearly equal. ............................................................................................................................... 3. L. volubile.

1. LYGODIUM HETERODOXUM Lygodium heterodoxum Kunze, Farrnkra¨uter 2: 32, t. 113. 1849. Fig. 182E.

Hydroglossum heterodoxum (Kunze) T. Moore, Index Fil. cxiv. 1857. Lygodictyon heterodoxum (Kunze) J. Sm., Ferns Brit. For. 259. 1866. Syntypes. Mexico. Oaxaca, Galeotti 6419 bis (LZ, destroyed; isosyntypes B!, BR!); cultivated specimens grown at Leipzig Botanical Garden, from plants originally collected in Guatemala. Hydroglossum spectabile Liebm., Mexic. Bregn. 299 (reprint 147). 1849. Lec-

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totype (chosen by Smith, 1981: 145). Mexico. Oaxaca: S. Pedro Tepinapa, Liebmann s.n. [Pl. Mex. 2634, Fl. Mex. 936] (C!; isolectotype US!).

Rhizomes short-creeping, hairs brown; stipes and rachises stramineous, glabrous; pinna apices dormant with two pinnules laterally divergent, each in turn Ⳳ dichotomously forked or nearly pinnately divided, pinnule axes flexuous; pinnae 2–3-forked, forming 4 (to several) trifid (or narrowly oblong) segments, axes and fertile lobes minutely strigose; segments strongly cordate to auriculate at base, each lobe linear-oblong, 8–20 ⫻ 1.5–2 cm, glabrous, margins entire; veins netted, reticulate after only 1–2 branchings from pinnule midvein; fertile lobes finger-like, ca. 1 mm wide with two rows of sporangia, each sporangium covered with a yellow, pocket-like, glabrous indusium. Distribution. Scrubby open areas of low elevations; 0–900 (–1300) m. Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Gr Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol, Parag. Selected Specimens Examined. Ags (Siqueiros D. 2367, IEB). Camp (Lundell 1415, NY). Chis (Breedlove 26554, NY). Col (McVaugh 15669, NY). Gro (Hinton 10794, NY). Hgo (Clark 6850, NY). Jal (McVaugh 25536, NY). Me´x (Hinton 7392, NY). Mich (King & Soderstrom 4952, NY). Nay (Rose 14309, NY). Oax (Mexia 9184, NY). Pue (Boege 430, MEXU). QR (Boom 282, NY). Qro (Carranza 1220, MEXU). Sin (Gonza´lez Ortega 4812, NY). SLP (Pringle 3318, MEXU). Tab (Cowan 1732, NY). Tam (Clark 7424, NY). Ver (Ventura A. 8184, NY). Yuc (Lundell 4843, MO).

Distribution. Frequent in open to lightly shaded scrubby habitats at lower elevations on Atlantic slope; 50–900(–1200) m. Mexico; Guat, Bel, Hond, Nic, CR, Pan. Selected Specimens Examined. Chis (Breedlove 57741 NY; Matuda 3720, NY). Oax (Mexia 9212, NY; Mickel 6842, NY; Torres C. 11128, MEXU). Pue (Vera Santos 2567, US). Tab (Cowan 2002, NY; Haynes 5624, NY). Ver (Nee 29972, NY; Williams 9027a, NY).

See discussion of the hybrid with L. venustum under that species.

2. LYGODIUM VENUSTUM Lygodium venustum Sw., J. Bot. (Schrader) 1801(2): 303. 1803. Lectotype (chosen by Proctor, Fl. L. Antill. 2: 51. 1977). Brazil. Breynius s.n. (S?). Figs. 182F–H. Lygodium mexicanum C. Presl, Reliq. Haenk. 1(1): 72. 1825. Type. Mexico. Haenke s.n. (PRC!).

Rhizomes short-creeping, 1–2 mm diam., hairs brown; stipes and rachises stramineous, wiry; axes glabrescent to densely hairy with short stiff hairs; pinna axes very short; pinnule axes pinnate, the segments alternating, each segment trifid to quinquifid, the central lobe much the largest, 3–10 ⫻ 0.5–1.4 cm, margins finely serrulate; veins free, sparsely strigose to densely hairy, hairs 0.5 mm long, whitish; fertile lobes ca. 1 mm wide, indusia with many 0.5 mm long white hairs; 2n⫽116 (Trin).

Lygodium venustum is easily distinguished from L. heterodoxum by its free veins, pubescent laminae, and palmately 3–5parted pinnules. Mexican specimens have often been named L. mexicanum and L. polymorphum (Cav.) Kunth, based on degree of pubescence, but these variants are merely part of the wide variation in this species. The name L. polymorphum is probably misapplied to this species; the type is from the Philippines, fide Christensen (Dansk. Bot. Ark. 9(3): 3–32. 1937), and probably referable to L. flexuosum (L.) Sw. In any case, L. polymorphum is illegitimate, since Cavanilles cited Ophioglossum scandens L. as a synonym (see Alston & Holttum, Reinwardtia 5: 15. 1959). A putative hybrid of L. heterodoxum and L. venustum, with malformed spores, was collected in Oaxaca (Mickel 4179, NY). It has some free and some anastomosing veins, fertile lobes with associated whitish hairs, pinnules of intermediate width (11–14 mm wide), pinnule axes flexuous, and inequilaterally 2–4 palmately lobed pinnules.

3. LYGODIUM VOLUBILE Lygodium volubile Sw., J. Bot. (Schrader) 1801(2): 304. 1803. Type. Jamaica. Swartz s.n. (S). Figs. 182A–D. Lygodium volubile differs from L. venustum in the ultimate pinnules all about the same size, unlobed at bases and rather long-stalked; 2n⫽58, 174–180 (Jam.).

MACROTHELYPTERIS

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spores. Also, the stipe vasculature of Macrothelypteris is of the typical thelypteroid type (two vascular bundles), whereas in Ctenitis and Lastreopsis there are numerous bundles arranged in a partial circle. Recent molecular evidence shows that Macrothelypteris, together with the north temperate Phegopteris (beech ferns) and the paleotropical genus Pseudophegopteris, form the basalmost clade and sister group to all other genera and species within the Thelypteridaceae, a family comprising nearly 1000 species worldwide (Smith & Cranfill, 2002). References Holttum, R. E. 1969. Studies in the family Thelypteridaceae. The genera Phegopteris, Pseudophegopteris, and Macrothelypteris. Blumea 17: 5–32; Smith, A. R. & R. Cranfill. 2002. Intrafamilial relationships of the thelypteroid ferns (Thelypteridaceae). Amer. Fern J. 92: 131–149.

Distribution. Lowland wet forests; 0–200 m. Mexico; Bel, Guat, Hond, Nic, CR, Pan; Jam, Cuba; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol, Arg. Specimens Examined. Chis (Mu¨nch 98, DS, MICH). Ver (Lot 1244, MEXU).

This is very close to, and possibly not distinct from, L. micans J. W. Sturm in Martius, type from Guyana.

7 2 . M A C R O T H E L YPT E R I S Macrothelypteris (H. Itoˆ) Ching, Acta Phytotax. Sin. 8: 308. 1963. Thelypteris sect. Macrothelypteris H. Itoˆ in Nakai & Honda, Nov. Fl. Jap. 4: 141. 1939. Thelypteris subg. Macrothelypteris (H. Itoˆ) A. R. Sm., Phytologia 34: 233. 1976. Type: Thelypteris torresiana (Gaudich.) Alston [⬅ Polystichum torresianum Gaudich.]⬅ Macrothelypteris torresiana (Gaudich.) Ching.

Terrestrial; caudices short, creeping or suberect; blades bipinnate to bipinnate-pinnatifid with Ⳳ adnate pinnules; proximal pinnae not reduced or only slightly reduced; aerophores absent; scales on rachises often with thickened bases and an acicular hairtip, sometimes with marginal hairs; veins free, usually branched, ending before the margin in a clavate tip (as seen adaxially); hairs on blades acicular and capitate, often with some long-septate hairs; sori usually with small indusia; sporangia usually with small capitate glands near annulus; spores with a Ⳳ winged perispore; x⫽31. Macrothelypteris comprises nine species in the Old World tropics and subtropics, from Africa to Queensland and Hawaii. A single species is widely naturalized in various parts of the New World. This genus is easily distinguished from Thelypteris in Mexico by the more dissected blades. In frond form it is more likely to be confused with dissected species of Ctenitis, such as C. equestris, or Lastreopsis spp. From Ctenitis it differs in lacking short, reddish, multicellular hairs on the costae adaxially, lack of scales on the rachis and costae below, presence of hyaline, acicular hairs on the lamina below, and in base chromosome number and

1. MACROTHELYPTERIS TORRESIANA Macrothelypteris torresiana (Gaudich.) Ching, Acta Phytotax. Sin. 8: 310. 1963. Figs. 183A–C. Polystichum torresianum Gaudich. in Freycinet, Voy. Uranie 333. 1828. Thelypteris torresiana (Gaudich.) Alston, Lilloa 30: 111. 1960. Type. Mariana Islands. Gaudichaud s.n. (P). Aspidium uliginosum Kunze, Linnaea 20: 6. 1847. Dryopteris uliginosa (Kunze) C. Chr., Index Filic., Suppl. 3: 100. 1934. Thelypteris uliginosa (Kunze) Ching, Bull. Fan Mem. Inst. Biol. 6: 342. 1936. Lectotype (chosen by Morton, Contr. U.S. Natl. Herb. 38: 219. 1973). Cultivated in Hort. Bot. Leipzig, origin Java (BR, photo US). Additional synonymy cited in Holttum (Blumea 17: 5–32. 1969).

Rhizomes short-creeping, stout; fronds 60–150 cm; stipes green-glaucous when living, stramineous when dry, to 75 cm ⫻ 3–12 mm, at base with narrow castaneous hairy scales, the rest of stipe glabrous or glabrescent; blades to ca. 70 ⫻ 50 cm, deeply bipinnate-pinnatifid, with 10–18 pairs of free pinnae, proximal or sub-proximal ones the longest; largest pinnae to 35 ⫻ 17 cm, deltate, pinnate with all pinnules but the larger ones Ⳳ adnate to a narrowly green-winged costa; pinnules oblique to costae, basal ones of proximal (but not distal) pinnae reduced, largest 2–8 ⫻ 0.8–2.5 cm, cut almost to costule into oblique, dentate to deeply lobed segments 2.5–5 mm wide; veins in largest segments 7–12 pairs, forked or pinnate, distal part of each vein thickened; costae and costules abaxially clothed with scattered pale, stiff, slender hairs, some multicellular and over 1 mm long; laminar tissue abaxially with short, erect, unicellular capitate glands; sori round, indusia very small or seemingly absent; sporangia each with 2–3 short capitate glands near annulus; n⫽62, 93 (OW).

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MARATTIA References Lavalle, M. D. C. 2003. Taxonomia de las especies neotropicales de Marattia (Marattiaceae). Darwiniana 4: 6–86; Underwood, L. M. 1909. Marattiaceae. N. Amer. Fl. 16: 15–23.

Key to the Mexican Species of Marattia l. Blades bipinnate, segments 10–30 ⫻ 1–3.2 cm. ..................2. M. laxa. l. Blades tripinnate, segments 0.5(–4) cm long. 2. Ultimate segments mostly 2–3 ⫻ 1–1.3 cm. ............ 1. M. excavata. 2. Ultimate segments mostly 0.7–2 ⫻ 0.6–0.8 cm. ....................................................................3. M. weinmanniifolia.

1. MARATTIA EXCAVATA Distribution. Along roadsides and trails, wet woods, seasonal evergreen forests, pine-oak forests; 50–1900 m. Se USA; Mexico; Guat, Bel (Davidse 36036, UC), Hond, Salv (Seiler, 1982: 387), Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Ec, Peru, Braz, Bol, Parag, n Arg; Africa, Asia, Pacific islands. Selected Specimens Examined. Chis (Breedlove 56800, CAS). Gro (Lorea 4071, FCME; Gallego H. 253, FCME). Me´x (Tejero-Dı´ez 4103, IZTA). Oax (Mickel 5921, NY, UC., 6400, NY). Mor (Sandoval 98, UAMIZ). Qro (Ferna´ndez N. 3895, IEB; Rzedowski 46385, IEB, XAL). Tab (Cowan 1723, CAS, CHAPA, CSAT). Ver (Breckon & Breckon 2050, UC; Nee & Calzada 22727, LL).

Marattia excavata Underw., N. Amer. Fl. 16: 22. 1909. Type. Costa Rica. Coliblanco, Maxon 272 (NY!). Figs. 184A, B. Similar to M. weinmanniifolia, but differing in the ultimate segments mostly 2–3 ⫻ 1–1.3 cm; axes less scaly below, glabrescent or with a few hair-like scales.

The distinguishing characters of this species are those mentioned under the genus. Macrothelypteris torresiana is widely naturalized in the neotropics and is rapidly expanding its range. The first Mexican collections were made in Oaxaca in 1964, in Veracruz in 1970, and in Chiapas in 1971. Now it is a commonly collected species in these three states, with over 50 collections seen.

7 3. M A R A T T I A Marattia Sw., Prodr. 8, 128. 1788. Type: Marattia alata Sw. Terrestrial; rhizomes compact, stout, fleshy, horizontal to erect, with scales amorphous, thin, light brown, and with large, fleshy stipules; roots large, fleshy; fronds large, coarse, fleshy; blades deltate, 2–4 pinnate, swollen at branching points; veins free; laminae glabrous to slightly hairy; ultimate axes often winged; sori exindusiate; sporangia in elliptic, exindusiate, compound structures (synangia) near the ends of the veins, each sporangium opening by a pore into a 2-valved, clam-shaped structure; spores bilateral; x⫽40(39). Marattia is a pantropical genus of about 60 species, mostly of the South Pacific. It grows in wet forests of higher elevations. The large stipules are sometimes used for food, hence the local name “maı´z del monte.” The genus is distinct by the large, fleshy fronds, huge stipules, and clam-like synangia. Among the Mexican ferns, Marattia is most closely related to Danaea.

Distribution. Terrestrial in montane rain forests, evergreen rain forests, cloud forests; 1900–2400 m. Mexico; Guat, Hond, Salv, Nic, CR, Pan. Specimens Examined. Chis (Breedlove 22649, F, MEXU, MICH, MO, NY, 31624, DS; Croat & Hannon 65190, MEXU, US; MacDougall 7921, MEXU; Martı´nez S. 19640, MEXU; Purpus 6750, NY, UC, US; Riba 1403, ENCB, MEXU).

MARSILEA

The size of the ultimate segments within and among species of Marattia varies greatly, depending on whether the segments are from distal or proximal pinnae. In areas where both M. excavata and M. weinmanniifolia occur, it may be difficult to place all collections with confidence.

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mostly 7–20 ⫻ 6–8 mm, elliptic, bases rounded, apices obtuse to rarely acuminate; ultimate rachises winged between segments; margins serrate; veins mostly simple; laminae glabrous adaxially, glabrous abaxially or with sparse brown hairs or narrow scales; 2n⫽160 (Chis).

2. MARATTIA LAXA Marattia laxa Kunze, Linnaea 18: 306. 1844. Figs. 184G, H, J. Gymnotheca laxa (Kunze) C. Presl, Gefa¨ssbu¨ndel Farrn 275. 1845. Type. “In humidis region. temper.” (Herb. Roemer) Leibold 62 (Leibold s.n. at K! with Kunze writing “Marattia laxa mihi” and frag. NY! from K–3 sheets!).

Fronds bipinnate, 1.5–4 m long; pinnules 10–30 ⫻ 1–3.2 cm, narrowly oblong, bases rounded, truncate or cordate to auriculate, apices acuminate, margins serrulate; veins free, 1–2 times forked. Distribution. Dense montane forests or cloud forests on Atlantic slope, oak forests with Liquidambar, Drimys, or pines; 800–2700 m. Mexico; Guat, Hond, Salv, CR, Pan. Selected Specimens Examined. Chis (Breedlove 32343, DS, MEXU). Gro (Rzedowski & McVaugh 141, ENCB, MEXU, MICH). Hgo (Sa´nchez-Mejorada 227, MEXU, US). Jal (Cochrane & Judziewicz 10669, CAS, IBUG, UC, WIS; McVaugh 21417, MEXU, MICH, NY). Oax (Hallberg 1797 NY, US; Mickel 5573, MEXU, NY, UC). Pue (Pringle 10299, C, CAS, CHAPA, GH, K, MEXU, MICH, MO, NY, UC, US). Qro (Rzedowski 46391, ENCB, IEB, MEXU, NY). Ver (Ramı´rez 693, XAL).

Distribution. Steep, rocky slopes in wet oak forests of upper elevations on the Atlantic slope, pine-oak forests, limestone; 750–2500 m. Mexico; Hond, CR, Pan. Selected Specimens Examined. Oax (Conzatti & Gonza´lez 723, GH, MEXU, US; Mickel 4117, NY, UC). Pue (Lorea 1087, FCME; Ma´rquez et al. 782, ENCB, NY, XAL; Tenorio L. 15441, GH, MEXU; Ventura A. 21996, ENCB). Ver (Cha´zaro & Cha´zaro 4037, US; Nee & Diggs 24881, XAL; Purpus 11140, CAS, F, K, MO, NY, US; Ventura 194, ENCB, UC).

This species is readily distinct from M. weinmanniifolia and M. excavata by having bipinnate fronds and large blade segments.

3. MARATTIA WEINMANNIIFOLIA Marattia weinmanniifolia Liebm., Mexic. Bregn. 308 (reprint 156). 1849. Lectotype (chosen by Smith, 1981: 147). Mexico. Oaxaca: Dist.Villa Alta, between Tonaguia and Roayaga, Liebmann s.n. [Fl. Mex. 960] (C!–in folia herbarium). Figs. 184C–F. Rhizomes massive, covered with old stipules, ca. 15 cm diam.; fronds 2–4 m long, tripinnate, ovate or deltate-ovate; axes sparsely to abundantly clothed with light brown scales; segments

Marattia weinmanniifolia is similar to M. alata Sw. of the West Indies, but that species has abundant, broader scales on the midveins and veins of the ultimate segments, teeth more curved, and the ultimate segments 2 or more times as long as wide. Marattia excavata has larger segments 20–30 ⫻ 10–13 mm.

7 4 . M A R SI L E A Marsilea L., Sp. Pl. 2: 1099. 1753. Lectotype (chosen by Maxon, Pteridophyt. Porto Rico 509. 1926): Marsilea quadrifolia L. Zaluzianskia Neck., Hist. & Commentat. Acad. Elect. Sci. Theod.-Palat. 3: 303. 1775, non F. W. Schmidt, 1793. Type: Zaluzianskia marsiloides Neck. [⫽ Marsilea quadrifolia L.].

Aquatic, subaquatic, or amphibious, forming diffuse or dense colonies, usually rooted, or entire plant floating; rhizomes shortto long-creeping, slender, with lateral shoots arising at nodes, bearing hairs; roots adventitious, thread-like, present only at nodes or also on internodes; leaves monomorphous, but emergent ones often somewhat different from floating or nonemergent ones, distichous, generally less than 30 cm long; stipes slender, upright or decumbent, glabrous or hairy, bearing at the apex two pairs of cuneate-flabellate leaflets that are pulvinate at

382

MARSILEA

the base; leaflets (pinnae) dichotomously veined, the veins anastomosing regularly, especially near the margin, forming a reticulum with elongate areoles; lateral margins of leaflets concave, straight, or convex, terminal margin entire or denticulate; indument of rhizomes, leaves, and sporocarps often with 2–9-celled uniseriate trichomes; fertile leaves usually terrestrial, with branched or unbranched peduncles bearing hardened, globose or discoid, nut-like sporocarps at the tips and diverging from the stipe at or above its base; sporocarps attached laterally or terminally to peduncles, which may be adnate for a short distance to the sporocarp (the adnate portion called a raphe), peduncles often ending in a blunt tooth, an additional superior tooth sometimes present; sori enclosed in sporocarps, disposed in two rows, sausage-shaped, with microsporangia borne along the margins of the sori and megasporangia borne medially, sori attached by a thin medial membrane to an elongate gelatinous mass (sorophore); sporangia hyaline, lacking annuli; microspores 16–64 per sporangium, white or pale brown, tetrahedral, smooth or variously ornamented; megaspores one per sporangium, ovoid, white, surface smooth or indistinctly reticulate; x⫽20. Marsilea comprises 45 to 70 species, and is subcosmopolitan, but primarily in temperate areas and subtropical areas; the greatest number of species is found in Africa and Australia. The species grow in ephemeral pools in drier areas (seasonally wet habitats), savannas, wet ditches, along creek and river margins, or in permanent pools and marshes. The New World species, nine native and two naturalized, were monographed by Johnson

(1986), and our treatment is based primarily on this work. Seven species occur sporadically in Mexico. Non-reproductive specimens can be difficult or impossible to identify, and some populations apparently remain vegetative for long periods. Sporocarps of Marsilea often have great longevity. Johnson (Amer. Fern J. 75: 30. 1985) was able to grow sporophytes of M. mollis from sporocarps taken from a herbarium specimen that had been dry for 89 years. Sporocarps of Marsilea are known to be transported by waterfowl, on muddy feet, or may be ingested and passed intact through digestive tracts (Johnson, 1986: 24). The sporocarps of an Australian species, M. drummondii A. Braun, commonly called nardoo, are ground into flour by Aboriginals, then mixed with water to form an edible paste. The Marsileaceae were once thought to be related to the Schizaeaceae, but molecular and fossil evidence now suggests that the family is most closely allied and sister to the two other heterosporous fern families, Salviniaceae and Azollaceae (Pryer et al., 1995; Rothwell & Stockey, 1994). References Johnson, D. M. 1986. Systematics of the New World species of Marsilea (Marsileaceae). Syst. Bot. Monogr. 11: 1–87; Johnson, D. M. 1993. Marsilea. pp. 332– 335. In: FNA Ed. Comm.; Palacios-Rios, M. & D. M. Johnson. 1992. Marsileaceae. In: V. Sosa (ed.), Flora de Veracruz 70: 1–11; Pe´rez-Garcı´a, B., R. Riba & D. M. Johnson. 1999. Pteridofitas, Familia Marsileaceae. Flora de Mexico 5: 1–17; Rothwell, G. W. & R. A. Stockey. 1994. The role of Hyropteris pinnata gen. et sp. nov. in reconstructing the cladistics of heterosporous ferns. Amer. J. Bot. 81: 479–492; Schneider, H. & K. M. Pryer. 2002. Structure and function of spores in the aquatic heterosporous fern family Marsileaceae. Int. J. Pl. Sci. 163: 485–505.

Key to the Mexican Species of Marsilea 1. Rhizomes bearing roots only at nodes. 2. Distal tooth of sporocarps 0.4–1.2 mm, acute. ......................................................................................................................................... 7. M. vestita. 2. Distal tooth of sporocarps absent or to 0.4 mm and blunt. 3. Sporocarps 6–9 mm, strongly ascending; peduncles usually branched. .................................................................................... 4. M. macropoda. 3. Sporocarps 2.4–6 mm, perpendicular to strongly nodding, i.e., peduncles curved or bent, unbranched. 4. Peduncles recurved or prostrate, sporocarps underground or below rhizome level, often hooked at base of raphe; sporocarps usually lacking a superior tooth, often covered with shaggy, reddish hairs. ................................................... 1. M. ancylopoda. 4. Peduncles erect or slightly decumbent, sporocarps never underground or below rhizome level, not hooked at base of raphe; sporocarps with superior tooth or not, covered with thin, uniform, white to pale brown hairs. .............................. 5. M. mollis. 1. Rhizomes bearing roots both at nodes and on internodes. 5. Sporocarps with a raphe and often with a superior tooth, usually one sporocarp per leaf. ................................................................ 5. M. mollis. 5. Sporocarps each lacking a raphe and superior tooth, 1–25 sporocarps per leaf. 6. Sporocarps each 1–4, on proximal 1⁄4 of stipes; sporocarps 3.5–6 mm long, angled in cross section, with conspicuous lateral ribs. ............................................................................................................................................................................................................ 3. M. deflexa. 6. Sporocarps 3–25, on proximal 2⁄3 of stipes; sporocarps 4 mm long or less, round, ovate, or slightly angled in cross section. 7. Sporocarps 3.1–3.9 mm long, slightly angled or ovate in cross section; rhizomes 1.1–2.3 mm diam.; sporocarps 5–20 per stipe, usually crowded and overlapping. .................................................................................................................................. 2. M. crotophora. 7. Sporocarps less than 3 mm long, round in cross section; rhizomes 0.5–0.7 mm diam.; sporocarps 7–25 per stipe, distant, not overlapping. .............................................................................................................................................................. 6. M. polycarpa.

1. MARSILEA ANCYLOPODA Marsilea ancylopoda A. Braun, Monatsber. Ko¨nigl. Preuss. Akad. Wiss. Berlin 1863: 434. 1864. Figs. 186E, F.

Zaluzianskia ancylopoda (A. Braun) Kuntze, Revis. Gen. Pl. 2: 823. 1891. Type. Ecuador. Environs of Guayaquil, Jameson 394 (G; isotypes BM!, G, K!). Marsilea mexicana A. Braun, Monatsber. Ko¨nigl. Preuss. Akad. Wiss. Berlin 1870: 747. 1871. Zaluzianskia mexicana (A. Braun) Kuntze, Revis. Gen. Pl. 2: 823. 1891. Type. Mexico. Julisca [Jalisco:] Beechey s.n. (K!).

Differs from Marsilea mollis by: rhizomes with short nodal shoots, dark red to black, apices with rusty hairs, glabrate be-

MARSILEA DEFLEXA

383

neath; land fronds with glabrous stipes, leaflets covered with nonoverlapping hairs or glabrous; abaxial intercostal striae irregularly distributed or absent; sporocarps solitary, on spreading or retrorse peduncles often hooked near the apices, rectangular to round in lateral view, lateral ribs none, covered with shaggy pelt of white to fulvous hairs, sometimes glabrate, (2.5–)4–6 ⫻ (2–)2.5–5 mm, 2.1–3.2 mm thick, superior tooth a broad bump 0.1 mm high.

Distribution. Ditches and lake margins, stagnant pools, and flooded areas; 0–100 m. Mexico; Nic; Ven, Braz, Bol, Parag. Specimens Examined. Tab (Saunders 231, US). Ver (Calı´x de D. & Castillo 95, XAL; Lot 1295, CAS, GH, MO, MEXU, XAL-photo ex MO; Palacios-Rios et al. 3236, 3238, XAL). Unverified, Doubtful, or Mistaken Reports. Gro (Lorea 5129, FCME, cited by Pe´rez-Garcı´a et al., 1999, but not verified).

Distribution. Dense clay substrates on the borders of temporary ponds and ditches in otherwise arid regions; 0–2400 m. Mexico; Guat, Hond, Nic, CR, Pan; Jam, Hisp, PR; Col, Ven, Ec, Peru, Braz, Bol, Parag, Arg, Uru. Specimens Examined. Jal (Lott et al. 3321, CAS, MO; Lott 3987, CAS, MO, UC, UCR; McVaugh 26057, CAS, IEB, MO). Mich (Dı´az Barriga 6422, IEB). Nay (Jones 23510, MO, UC; Murray & Johnson 1458, MICH, NY, UC). Unverified, Doubtful, or Mistaken Reports. Chih (Spellenberg 9557, MEXU, NMC, NY, cited by Spellenberg et al., 1996; the specimen is vegetative, and possibly M. vestita, as annotated by Yatskievych in 1996). Qro (Rzedowski 48832, 48589, both IEB, cited by Pe´rezGarcı´a et al., 1999, but not verified). Tab (reported by Magan˜a, 1992, but not verified and possibly based on a misidentification).

Marsilea ancylopoda produces sporocarps throughout the year, perennating solely by sporocarps. The distinctive recurved peduncle with the sporocarps developing and maturing underground is unique among New World marsileas. The broad virtually glabrous leaflets and pelt of appressed reddish hairs on the sporocarps are also characteristic.

2. MARSILEA CROTOPHORA Marsilea crotophora D. M. Johnson, Syst. Bot. Monogr. 11: 46. 1986. Type. Brazil. Mato Grosso do Sul: Corumba, Hatschbach & Scherer 30470 (US!; isotypes C, LP, M, MICH!, NY!, UC!). Figs. 185H, J. Differs from M. polycarpa by: rhizomes thick (1.1–2.3 mm diam.), with 5–20 crowded and overlapping sporocarps in a single row on stipes; sporocarps hairy, dark gray to black, 3.1–3.9 ⫻ 2.9–3.7 mm, 1.2–2.2 mm thick, raphe and teeth absent.

Marsilea crotophora is distinguishable from M. polycarpa (q.v.) by the thicker rhizomes and the larger and overlapping sporocarps, which extend to 2⁄3 of way up the stipes.

3. MARSILEA DEFLEXA Marsilea deflexa A. Braun, Monatsber. Ko¨nigl. Preuss. Akad. Wiss. Berlin 1863: 421. 1864. Figs. 185D–G. Zaluzianskia deflexa (A. Braun) Kuntze, Revis. Gen. Pl. 2: 823. 1891. Type. Brazil. Piauı´: Gardner 2760 (G; isotypes BM, G, K, P).

Main rhizomes with nodal and internodal roots, internodes 1.5–6.5 cm long, apices with dense, tan hairs, glabrate beneath, rhizomes in aquatic habitats green; land fronds 2–9.3 cm long, stipes terete and glabrate, leaflets 1.1–2 ⫻ 0.9–1.7 cm, glabrous to sparsely hairy, lateral margin concave to convex, terminal margin entire; floating leaves 4–18 cm long, stipes terete and glabrous, leaflets 1.4–3.1 ⫻ 1.6–2.9 cm, lateral margin straight to convex; abaxial intercostal striae almost always present and uniformly distributed, except for small patch just distal to the pulvinus; fertile leaves in water and on land, bearing 1–4 sporocarps on lower 1⁄4 of stipes; sporocarps rectangular, quadrate in lateral view, with conspicuous lateral ribs, covered with flattened matted hairs, eventually glabrate, brown to purplish black, 3.5–6 ⫻ 2–4.5 mm, 1.7–2.8 mm thick, raphe and teeth absent.

384

MARSILEA

Distribution. Aquatic on clay substrates or floating in seasonal ponds, marshes and ditches; 0–1100 m (in total range). Mexico; Guat, Hond, CR; Col, Ven, Peru, Braz, Parag.

Distribution. Aquatic along seasonally wet roadsides, black clays (pH 7–8), and less frequently on sandy soils; 0–1400 m. USA (Ala, La, Tex); Mexico.

Specimens Examined. Jal (Rzedowski & McVaugh 1331, ENCB). Nay (McVaugh 19287, CAS, IEB, MICH, MO). Oax (Lachica & Sa´nchez FaI-2118, CAS, ENCB, MEXU). Pue (Bravo-Hollis 523, MEXU). Ver (Ortega O. 583, NY, UC, XAL).

Specimens Examined. Coah (Gould 6405, ANSM, CAS, MICH, UC). Hgo (Edwards 789, CAS, F, MO, TEX; Fisher 3798, F, GH, MICH). NL (Barkley 14344, ENCB, GH, MO, NY, TEX, UC). Tam (Harriman 11408, NY; Haynes 5440, MICH; Pringle 1975, BM, F, K, LL, MO, NY, UC, US; Wussow & Landry 164, LAF, US).

Unverified, Doubtful, or Mistaken Reports. Yuc (Tun G. 544, UADY, cited by Palacios-Rios, 2002b, but not verified).

Marsilea deflexa produces sporocarps throughout the year. It does not persist as sporophytes with viable rhizomes through the dry seasons, but new plants are produced each wet season by dehiscence of sporocarps. Hybrids are known between this species and M. polycarpa in Venezuela (Johnson, 1986). See M. polycarpa for comparison.

4. MARSILEA MACROPODA Marsilea macropoda Engelm. ex A. Braun, Amer. J. Sci., ser. 2, 6: 88. 1848. Figs. 186A–D. Zaluzianskia macropus (A. Braun) Kuntze, Revis. Gen. Pl. 2: 823. 1891. Type. U.S.A. Texas: swamps of Guadalupe bottoms near Victoria, Lindheimer III 573 (MO; isotypes B, BM, G, K, LL, MO, PH, UC!, US).

Plants forming dense clones; roots only at nodes; stipes 5–39 cm long, hairy; pinnae 9–35 ⫻ 8–39 mm, conspicuously whitehairy abaxially, hairy adaxially, hairs often exceeding margin to make pinnae appear white-margined from above; sporocarp peduncles erect, frequently branched, attached 2–4.5 mm above bases of stipes, stalks 8–17 mm long, common trunk of branched stalks 1–5 mm (rarely 2 or 3 unbranched stalks attached separately to same stipe); sporocarps strongly ascending, 6–9 ⫻ 4.5–5.5 mm, 2 mm thick, elliptic or quadrate in lateral view, covered with matted and often twisted hairs, proximal tooth of each sporocarp 0.4–0.5 mm long, blunt, distal tooth absent or present as slightly raised area; raphes 1.6–2.4 mm; sori 19–23 per sporocarp.

Hybrids with M. vestita have been reported from Texas (Johnson, 1986: 73).

5. MARSILEA MOLLIS Marsilea mollis B. L. Rob. & Fernald, Proc. Amer. Acad. Arts 30: 123. 1895. Type. Mexico. Chihuahua: St. Diego, Hartman 604 (GH; isotypes F, GH, K!, MEXU!, MSC, NY!, UC!, US!). Figs. 187C, D. Main rhizomes brown to black with nodal roots, occasionally with an internodal root, internodes 0.8–2.3 cm long, apices densely pilose, glabrate beneath, rhizomes in water green; land fronds 1.1–14 cm long, stipes terete and pilose, leaflets .25–1.7 ⫻ 0.13–1.6 mm, densely pilose beneath, sparsely pilose or glabrous above, lateral margin convex to straight; floating leaves 12.5–100 cm long, leaflets 0.5–2.6 ⫻ 0.5–2.9 cm; abaxial intercostal striae irregularly distributed; fertile leaves on land, bearing one sporocarp on erect to slightly decumbent peduncle; sporocarps ovate in lateral view, often with conspicuous lateral ribs, gray-brown to brownish black, covered with acicular 4–5-celled verrucose hairs, soon glabrate, 2.4–4 ⫻ 2–3 mm, 1.3–1.7 mm thick; raphe 0.6–1.4 mm long, inferior tooth blunt or absent, superior tooth blunt or absent.

MARSILEA VESTITA

Distribution. Clay substrates on sandy soils, in temporary ponds and lakes; 1200–2500 m. USA (Ariz, Tex); Mexico; Peru, Chile, Arg. Probably also Colombia and Ecuador, where represented only by sterile collections (Johnson, 1986: 58).

385

Main rhizomes green to black, with nodal and internodal roots, 0.5–3 mm diam., internodes 0.8–15⫹ cm long, apices with sparse to dense, tan hairs, glabrate proximally; land fronds 3–15 cm long, stipes usually channeled, glabrous or glabrate, leaflets flabellate, 0.6–3 ⫻ 0.5–3 cm, glabrous to moderately hairy, lateral margins slightly concave to straight, distal margin rounded; floating leaves 6–28 cm long, stipes terete, glabrous, leaflets 2.4–4 ⫻ 1.9–3.5 cm, lateral margins straight to slightly convex; abaxial intercostal striae often present and uniformly distributed; fertile leaves on land, rarely on floating plants, bearing 4–26 sporocarps in a single row on unbranched, recurved peduncles, on proximal 1 ⁄2 of stipes and beginning 0.5–4 cm above stipe bases; sporocarps round to ovate in lateral view, without conspicuous lateral ribs, villous, eventually glabrate, brown, 2–2.6 ⫻ 1.6–2 mm, 1.7–1.9 mm thick, raphe and teeth absent.

Selected Specimens Examined. Ags (Rzedowski & McVaugh 882, MICH). BCS (Brandegee s.n., Sep 1893, DS). Chih (Pringle 2007, MEXU, MO, NY, TEX, UC, US). Chis (Murray & Johnson 1404, NY, UC). DF (Schaffner 17, 18, NY). Dgo (Palmer 13, F, LL, K, MEXU, MO, NY, US). Gto (Dziekanowski et al. 1797, MO, NY). Hgo (Moore 3452, UC, US). Jal (Pringle 2434, CAS, F, K, MEXU, MO, NY, UC, US). Me´x (Garcı´a P. 987, CHAPA, F, LL, MO, NY; Koch 74196, CAS, CHAPA, ENCB, MEXU). Mich (Dı´az Barriga 2194, IEB). Oax (Pringle s.n., 7 Jun 1894, F). SLP (Rollins & Tryon 58319, GH, US). Son (Hartman 893, GH, MO, UC, US). Ver (Brigada Vegetacio´n Acua´tica 339, MEXU, XAL). Unverified, Doubtful, or Mistaken Reports. Mor (Jackson 19, MICH, cited by Pe´rez-Garcı´a et al., 1999, but not verified). Nay (Alava & Cook 1529, UC, sterile, identification not certain). Pue (Arse`ne 520, 2283, US, cited by Pe´rez-Garcı´a et al., 1999, but not verified). Qro (Rzedowski 48832, IEB, not seen, cited by Arreguı´n et al., 2001, but this is M. ancylopoda, according to Pe´rez-Garcı´a et al., 1999).

The small sporocarps with spreading hairs, lacking a superior tooth and attached to wiry peduncles characterize this species. Marsilea mollis bears sporocarps March through December (and perhaps all year) in Mexico. See M. ancylopoda for comparison with that species. The following sterile collections are probably this species, but in the absence of sporocarps, we cannot be certain: Mor (Lyons 63, US; Bonilla 316, HUMO, MEXU, cited by Riba et al., 1996, but not verified). Pue (Arse`ne 520, 2283, US). Qro (Rzedowski 42619, 43101, IEB).

6. MARSILEA POLYCARPA Marsilea polycarpa Hook. & Grev., Icon. Filic. 2: t. 160. 1831. Figs. 185A–C. Zaluzianskia polycarpa (Hook. & Grev.) Kuntze, Revis. Gen. Pl. 2: 823. 1891. Lectotype (chosen by Johnson, 1986: 44). Guyana. Demerara, Parker s.n. (K!; isolectotype K). Marsilea polycarpa Hook. & Grev. var. mexicana A. Braun, Monatsber. Ko¨nigl. Preuss. Akad. Wiss. Berlin 1870: 713. 1871. Type. Mexico. [Veracruz?:] Mesachica, Schiede 836 (B!; isotype G).

Distribution. Forming diffuse colonies on mud or on floating vegetation, edges of marshes, ponds and seasonal lakes on clay substrates of subacid to neutral pH. 0–100 m. Mexico; Guat, Hond, Nic, CR, Pan; Gr Ant, Dominica; Col, Ven, Trin, Guy, Sur, Ec, Braz; Society Isl. Specimens Examined. Jal (Feddema 2532, IEB, MICH, MO). Unverified, Doubtful, or Mistaken Reports. Gro (Fonseca & Vela´zquez 1282, FCME, ENCB, cited by Pe´rez-Garcı´a et al., 1999, but not verified; also cited by Lozada, 1994, but not verified). Tab (Cowan 1593, CAS, PH; Ca´lix de Dios & Castillo 488, CSAT, and Orozco & Cowan 2953, CAS, cited by Pe´rez-Garcı´a et al., 1999, but not verified). Ver (Liebmann 2189 BM, C; Schiede 836, B, G).

Marsilea polycarpa is distinctive in its numerous, small sporocarps borne in a single row on the stipes and the lack of raphe and teeth on the sporocarps. Marsilea crotophora (q.v.) has a similar number and attachment of sporocarps, but the sporocarps are larger and the distalmost sporocarp is above the midpoint of the stipe. In sterile condition, M. polycarpa can be distinguished from M. deflexa by the former having abaxial intercostal striae all the way to the pulvinus and the tendency to have floating rather than submerged rooted rhizomes.

7. MARSILEA VESTITA Marsilea vestita Hook. & Grev., Icon. Filic. 2: t. 159. 1830. Figs. 187A, B.

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MEGALASTRUM

Zaluzianskia vestita (Hook. & Grev.) Kuntze, Revis. Gen. Pl. 2: 823. 1891. Lectotype (designated by Johnson, 1986: 62). U.S.A. Columbia River, Scouler 338 (K; isolectotypes GH, NY!). Marsilea mucronata A. Braun, Amer. J. Sci., ser. 2, 3: 55. 1847. Type. U.S.A. [North Dakota:] near Devil’s Lake, Geyer 71 (MO!; isotypes K, MO, NY!, US!). Marsilea minuta E. Fourn., Bull. Soc. Bot. Fr. 27: 329. 1880, hom. illeg., non L., 1771. Type. Mexico. San Luis Potosı´: in 1876 & 1882, Schaffner s.n. (P? isotype NY!).

Plants forming diffuse or dense clones; roots arising at nodes; petioles 2–20 cm long, sparsely hairy; pinnae 4–20 ⫻ 4–20 mm, hairy to glabrescent; sporocarp stalks erect, unbranched, attached at bases of petioles (occasionally up to 3 mm above bases), 0.5–25 mm, not hooked at tips; sporocarps perpendicular to stalk tips or slighly nodding, 3.5–7.5 ⫻ 3–6.5 mm, 1.5–2 mm thick, elliptic to nearly round in lateral view, hairy but soon glabrate, scars left by abraded hairs often appearing as purple or brown specks; proximal tooth of each sporocarp 0.3–0.6 mm long, blunt, distal tooth 0.4–1.2 mm, acute, often hooked at tip; raphes 1.1–1.7 mm; sori 14–22 per sporocarp.

Distribution. Edges of marshes, ponds, and seasonal lakes; 0– 500(–1900) m. Sw Canada, w USA; Mexico; Peru. Selected Specimens Examined. BCS (Carter & Moran 5934, BM, MEXU, UC; Wiggins 15162, CAS, ENCB, GH, MEXU, TEX, UC). Coah (Palmer 467, K, MO, NY, UC, US). DF (Schaffner 20, in 1875, NY). Dgo (Johnston et al. 11448, LL). NL (Seigler et al. 1292, TEX). SLP (Parry & Palmer 1010, F, K, MO, NY, US). Sin (Jones 23509, MO, UC). Son (Felger 85–991, ARIZ, TEX, UC). Zac (Jones 157, GH, DS, MO, US). Unverified, Doubtful, or Mistaken Reports. Chih (Spellenberg 9557, MEXU, NMC, NY cited by Spellenberg et al., 1996, as M. ancyclopoda, but not verified; the specimen is vegetative, and possibly M. vestita, as annotated by Yatskievych, 1996). Gro (cited by Lozada, 1994, but not verified).

7 5 . M E GA L A ST R U M Megalastrum Holttum, Gard. Bull. Straits Settlem. 39: 161. 1986 [1987]. Type: Megalastrum villosum (L.) Holttum [⬅ Polypodium villosum L.]. Dryopteris Adans. subg. Ctenitis C. Chr. sect. Subincisae C. Chr., Index Filic., Suppl. 3: 7. 1934. Ctenitis C. Chr. sect. Subincisae (C. Chr.) Tindale, Contr. New South Wales Natl. Herb. 3: 252. 1965. Type: Dryopteris subincisa (Willd.) Urb. [⬅ Megalastrum subincisum (Willd.) A. R. Sm. & R. C. Moran].

Terrestrial; stipes scaly, especially near their bases; blades 2–4pinnate, catadromous at the bases of proximal pinnae, the surfaces without cylindrical glands; proximal pinnae more developed basiscopically; veins simple or furcate, ending on adaxial surface of laminae in clavate tips well back from the blade margins, basal basiscopic veins of the distal pinnules arising from the costae, not from the costules; indument adaxially usually of stout, hyaline, acicular, falcate, septate hairs on rachises, costae, costules, and lesser axes, abaxially the midribs with similar or shorter hairs, also with linear to ovate, often marginally toothed scales, blades lacking short-segmented hairs of the type found in Ctenitis; indusia absent (present in a few spp. outside Mexico); spores echinate or with low, subparallel wings; x⫽41. In the neotropics, Megalastrum comprises about 40 species, and the species of South America are especially poorly known and virtually unstudied since the monograph by Christensen (1920). In addition, there is one species in Africa and five species in Madagascar and the Mascarene Islands. Megalastrum was relatively recently segregated from Ctenitis on the basis of the venation and trichome type (Holttum, 1986; Smith & Moran, 1987). The venation is characterized by the basal basiscopic veins of the ultimate segments arising from the penultimate costae, not from the midribs of the segments themselves; individual veins terminate well before the blade margins in clavate tips, as viewed adaxially. Distal segments of the pinnae tend to be strongly adnate, even decurrent basiscopically. Species lack ctenitoid hairs (short, twisted, often reddish, septate hairs) but have, on the midribs adaxially, stiff, antrorse, acicular hairs like those found on cyatheoid and thelypteroid ferns. The relationships of Megalastrum are certainly with the dryopteroid genera, as indicated by the base chromosome number of x⫽41, but beyond that they are uncertain. Unpublished molecular studies by Cranfill (pers. comm.) indicate that the relationship with Ctenitis is not particularly close, rather Megalastrum may be related to Lastreopsis (the species of which also often bear acicular hyaline hairs) and Rumohra. Further studies are needed. References

The most widespread subspecies, subsp. vestita, occurs throughout the range of the species and is the only one known from Mexico. A second subspecies, subsp. tenuifolia (Engelm. ex A. Braun) D. M. Johnson, is confined to central Texas (Johnson, 1986), but this apparently intergrades with subsp. vestita and is no longer considered separable by Johnson (1993).

Christensen, C. 1920. A monograph of the genus Dryopteris, part II. The tropical American bipinnate-decompound species. Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 8, 6: 3–132; Holttum, R. E. 1986. Studies in the fern-genera allied to Tectaria Cav. VI. A conspectus of genera in the Old World regarded as related to Tectaria, with descriptions of two genera. Gard. Bull. Straits Settlem. 39: 153–167; Smith, A. R. & R. C. Moran. 1987 [1988]. New combinations in Megalastrum (Dryopteridaceae). Amer. Fern J. 77: 124–130.

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387

Key to the Mexican Species of Megalastrum 1. Costae and costules abaxially lacking hairs, or if hairy, the hairs less than 0.5 mm, 1–3-celled. 2. Fronds 2–5 m long; costal scales with teeth 0.1 mm long. .......................................................................................................... 3. M. pulverulentum. 2. Fronds 1–2 m long; costal scales with teeth 0.05 mm long. ............................................................................................................ 4. M. subincisum. 1. Costae and costules abaxially hairy, the hairs 0.5–2 mm, 4–8-celled. 3. Scales of the rachises and costae dark brown, broadly ovate-lanceolate, entire or faintly erose. ...................................................... 2. M. lunense. 3. Scales of the rachises and costae brown or blackish, linear-lanceolate, denticulate. 4. Fronds mostly 1–2 m long; blades 2–3-pinnate-pinnatifid at bases; scales on midribs abaxially with teeth less than 0.05 mm long; blades densely hairy. ............................................................................................................................................................... 1. M. atrogriseum. 4. Fronds mostly 2–5 m long; blades 4-pinnate at bases; scales on midribs abaxially with teeth to 0.1 mm long; blades sparsely hairy. .............................................................................................................................................................................................. 3. M. pulverulentum.

1. MEGALASTRUM ATROGRISEUM

2. MEGALASTRUM LUNENSE

Megalastrum atrogriseum (C. Chr.) A. R. Sm. & R. C. Moran, Amer. Fern J. 77: 127. 1987 [1988]. Figs. 188C–E.

Megalastrum lunense (Christ) A. R. Sm. & R. C. Moran, Amer. Fern J. 77: 128. 1987 [1988]. Figs. 188A, B.

Dryopteris atrogrisea C. Chr., Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 8, 6: 70, f. 15. 1920. Ctenitis atrogrisea (C. Chr.) Ching, Sunyatsenia 5: 250. 1940. Type. Costa Rica. Tablazo, Biolley 70 (BM; isotype US!).

Differs from M. subincisum by the much longer hairs on the midribs and veins abaxially, the presence of long hairs on the veins adaxially, and the more weakly denticulate scales; from M. pulverulentum differing by the smaller fronds, and less decidedly toothed scales.

Distribution. Terrestrial in wet lowland forests, 250–400 m. Mexico; CR, Pan. Specimens Examined. Oax (Herna´ndez G. 1414, 1616, 1704, CHAPA, NY, 1962, CAS, CHAPA, MEXU). Ver (Ramı´rez 1095, XAL; Sinaca C. 1154, MEXU, MO).

This species differs from M. subincisum primarily in the much longer hairs on the costae, costules, and veins abaxially, and by having similar hairs on the veins adaxially; it also differs by the scales being less strongly denticulate along the midribs abaxially. From M. pulverulentum, with which it is also often confused, it differs in the much smaller fronds, less strongly denticulate scales, and more abundant hairs.

Aspidium lunense Christ, Bull. Herb. Boissier, se´r. 2, 6: 55. 1906. Dryopteris lunensis (Christ) C. Chr., Index Filic., Suppl. 1: 35. 1913. Ctenitis lunensis (Christ) Lellinger, Proc. Biol. Soc. Wash. 89: 709. 1977. Syntypes. Costa Rica. Werckle´ s.n. (P; isosyntypes NY!, UC!, US!). Dryopteris pansamalensis C. Chr., Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 8, 6: 72. 1920. Ctenitis pansamalensis (C. Chr.) Ching, Sunyatsenia 5: 250. 1940. Megalastrum pansamalense (C. Chr.) A. R. Sm. & R. C. Moran, Amer. Fern J. 77: 126. 1987 [1988]. Lectotype (here designated). Guatemala. Alta Verapaz: Pansamala´, Tu¨rckheim s.n. [Donn. Sm. 1055] (US 831791!; isotypes UC!, US 831790!).

Rhizomes short-creeping to suberect, plants appearing acaulescent; stipes stramineous to light brown, 30–65 cm ⫻ 4–6 mm, 1 ⁄3–1⁄2 the frond length, persistently scaly in distal part; stipe base scales dense, brown, spreading to appressed, broadly ovatelanceolate, mostly 5–15 ⫻ 1.5–4 mm, Ⳳ entire and lacking definite teeth, the cells isodiametric; blades deltate, 50–100⫹ cm long, 3-pinnate-pinnatifid or nearly 4-pinnate at bases, proximal pinna pair (14–)23–34 ⫻ 7–20 cm, much more strongly developed basiscopically (basal basiscopic pinnule to 15 ⫻ 6 cm) than acroscopically; middle pinnae 2-pinnate-pinnatifid, sessile to stalked to 1 cm; indument abaxially on rachises, costae, and costules of numerous brown, lustrous, ovate-lanceolate, entire scales, also with rather dense, hyaline, acicular septate hairs mostly 0.5–1.5(–2) mm long, veins with similar hairs, also often with shorter, erect, gland-tipped hairs to 0.4 mm on tissue between veins; indument adaxially of stout, curved acicular septate hairs ca. 1 mm long on costae, costules, and veins, sometimes with a few shorter hairs between veins; sori medial, exindusiate.

388

MEGALASTRUM

Distribution. Terrestrial in wet montane forests; 800–1000. Mexico; Guat, Bel (Holland 14A, UC), Nic, CR, Pan. Specimens Examined. Chis (Breedlove 42014, DS, MEXU, 56669, CAS, ENCB, NY).

This species can easily be distinguished from congeners in Mexico by the broadly ovate-lanceolate (vs. linear-lanceolate), entire or faintly toothed (vs. distinctly toothed) scales of the stipe bases, rachises, and costae.

3. MEGALASTRUM PULVERULENTUM Megalastrum pulverulentum (Poir.) A. R. Sm. & R. C. Moran, Amer. Fern J. 77: 129. 1987 [1988]. Figs. 188K–M. Ctenitis pulverulenta (Poir.) Copel., Gen. Fil. 124. 1947. Polypodium pulverulentum Poir. in Lamarck, Encycl. 5: 555. 1804. Dryopteris pulverulenta (Poir.) C. Chr. in Urban, Symb. Antill. 9: 305. 1925. Type. Plumier, Traite´ Foug. Ame´r. 27, pl. 34, 1705, illustrating a plant from Hispaniola. Polypodium karstenianum Klotzsch, Linnaea 20: 390. 1847. Dryopteris karsteniana (Klotzsch) Hieron., Hedwigia 46: 348. 1907. Ctenitis karsteniana (Klotzsch) Vareschi, Fl. Venez. 1(1): 405. 1969. Type. Venezuela. Karsten II. 3 (B). ?Phegopteris hirsuta Fe´e, Me´m. Foug. 5: 248. 1852. Type. Mexico. Funck & Schlim 975 (P?). Dryopteris karsteniana (Klotzsch) Hieron.var. heydei C. Chr., Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 8, 6: 77. 1920. Ctenitis pulverulenta (Poir.) Copel. var. heydei (C. Chr.) Stolze, Amer. Fern J. 67: 43. 1977. Megalastrum pulverulentum (Poir.) A. R. Sm. & R. C. Moran var. heydei (C. Chr.) A. R. Sm. & R. C. Moran, Amer. Fern. J. 77: 129. 1987. Type. Guatemala. Santa Rosa: Rı´o de los Esclavos, Heyde & Lux s.n. [Donn. Sm. 3249 p.p.] (US!; isotypes GH, MO).

Similar to Megalastrum subincisum except: plants with erect trunks to 0.7 m tall; fronds 2–5 m long; blades 4-pinnatepinnatifid at bases; indument abaxially on costae, costules, and ultimate segment midveins of soft, lax, hyaline hairs 0.8–1 mm long; costal scales with longer teeth ca. 0.1 mm; 2n⫽82 (Jam.).

Distribution. Terrestrial in wet lowland and montane forests; (250–)1200–2300 m. Mexico; Guat, Hond, CR, Pan; Jam, Hisp; Col, Ven, Ec, Peru, Bol, nw Arg. Selected Specimens Examined. Chis (Breedlove 22622, 31677, 32364, 33687, DS, MEXU, NY). Pue (Ventura 399, ARIZ, ENCB, LL, NY; Ventura A. 13640, IEB, MEXU, XAL). Qro (Rzedowski 42866, IEB). Ver (Castillo C. et al. 2543, XAL; Palacios-Rios 3314, XAL; Sa´nchez S. 345, US).

This is the largest species in the genus, with the fronds as long as or longer than many tree ferns. Specimens of Ventura 399 have fronds to 4.6 m long. Smith et al. 4139 (US), from Edo. Puebla, was noted as having a trunk two feet tall, an unusual condition in the genus as a whole, but possibly normal for this species.

4. MEGALASTRUM SUBINCISUM Megalastrum subincisum (Willd.) A. R. Sm. & R. C. Moran, Amer. Fern J. 77: 129. 1987 [1988]. Figs. 188F, G, H, J. Polypodium subincisum Willd., Sp. Pl., ed. 4, 5(1): 202. 1810. Dryopteris subincisa (Willd.) Urb., Symb. Antill. 4: 19. 1903. Phegopteris subincisa (Willd.) Fe´e, Me´m. Foug. 5: 243. 1852. Ctenitis subincisa (Willd.) Ching, Sunyatsenia 5: 250. 1940. Type. Venezuela. Caracas, Bredemeyer s.n. (B-Willd. 19701!). Polypodium galeottii M. Martens, Me´m. Foug. Mexique 43, pl. 7, f. 3. 1842. Type. Mexico. [Veracruz:] “Mirador” [on specimen], Galeotti 6321 (BR!; isotype BR! frag. US!). Phegopteris stenolepis Fe´e, Me´m. Foug. 8: 89. 1857 [“stenolepsis”]. Type. Mexico. Veracruz: Near Huatusco, Schaffner 239 (RB; isotype K!).

Rhizomes suberect, woody, stout; stipes stramineous to brownish, to 50⫹ cm ⫻ 7 mm, ca. 1⁄3 the frond length, glabrescent or with scattered, linear, fibrillose scales distally; stipe base scales dense, castaneous, lustrous, linear, 10–20(–30) ⫻ 0.1–0.3 mm, with prominent marginal falcate (hooked) teeth, the cells elongate; blades deltate, to 1(–1.5) m long, 3-pinnate-pinnatifid at bases, proximal pinna pair to 1 m long, much more developed basiscopically than acroscopically; middle pinnae mostly 2pinnate-pinnatifid, sessile to stalked to ca. 1 cm; indument abaxially on rachises, costae, and midveins, of scattered hair-like toothed scales (0.5–)1–4 mm long, sometimes also with sparse hyaline acicular hairs 0.1–0.3 mm long on costae and costules, and sometimes even a few short hairs on laminar tissue between

MELPOMENE

389

veins; indument adaxially of stout, curved, acicular, septate hairs 0.3–0.8 mm long on costae and costules, veins and tissue between veins usually glabrous; sori supramedial, exindusiate; 2n⫽ca. 164 (Jam).

entire, apices acute to attenuate, margins and surfaces lacking hairs or teeth, often with one to several papillose cells (glandular?) ca. 0.1 mm long at the scale tips, these cells often dislodged and not evident on older scales; fronds monomorphic or nearly so, determinate; stipes not articulate, 0.5–1.5 mm diam.; blades pinnatifid, pinnatisect, or 1-pinnate proximally, linear to elliptic, pinnae (segments) broadly adnate; rachises dark red-brown to blackish; veins free, often not readily visible, pinnate in the segments; hydathodes present and usually readily visible adaxially, not producing lime-dots; indument of stipes and blades of castaneous or deep red setae 0.3–3 mm long, setae multicellular but with cross walls difficult to see, also of lax pale hairs with reddish color concentrated at the cross walls, also of branched hairs 0.1– 0.2 mm long, branches 2–8-celled, some species nearly devoid of setae and hairs; sori Ⳳ round or slightly oblong, discrete, superficial or slightly sunken, ca. 1 mm diam., exindusiate; soral paraphyses absent, but some species with castaneous receptacular setae or with circumsoral setae; sporangia glabrous; x⫽37.

Distribution. Terrestrial in wet lowland and montane forests; 100–2000 m. Mexico; Guat, Hond, Nic, CR, Pan; Gr & L Ant; Col, Ven, Ec, Peru, Bol.

Melpomene comprises about 20 species, most of them in the neotropics, but at least one, M. flabelliformis, in Africa and on islands in the Indian Ocean. The genus differs from other genera of Grammitidaceae by the combination of strongly clathrate, non-setose, cordate-based rhizome scales that are attached at a point and often bear one to several gland-like papillae at the tip, pronounced hydathodes adaxially, pinnate, free veins, the presence (usually) of dark reddish brown to castaneous setae on the midribs and sometimes laminae, and branched pale hairs on the stipes and midribs. It is in need of modern taxonomic revision, especially in South America, where the variation and number of species are greater, especially at high elevations in the Andes. One suspects that much of the radiation in the genus is quite recent, corresponding to the uplift of the Andes within the last 15 million years. Unpublished molecular data show that the genus is monophyletic, possibly the sister group to Lellingeria (Ranker et al., 2004). When dried, several species of Melpomene emit a sweet, spicy fragrance that is not found or is uncommonly noted in other genera of grammitid ferns. This aroma can be almost overpowering to some individuals, while scarcely noticeable by others (Smith & Moran, 1992). With the recent transfer of Melpomene anfractuosa to Terpsichore, based in part on molecular evidence (Ranker et al., unpubl. data), the distinctive black clavate fungus Acrospermum is known to occur only on the blades of some species of Terpsichore.

Selected Specimens Examined. Chis (Calzada et al. 8770, UC). Gro (Hinton 14060, ARIZ, LL, NY, TEX, US; Lorea 2999B, 3294, FCME). Oax (Mickel 6191, NY, UC, US). Ver (Purpus 1980, NY, UC, US; Ventura A. 8217, ASU, 16270, ENCB, IEB, XAL).

Megalastrum subincisum and its close relatives are distinct from Ctenitis species in their toothed rhizome scales and long, often curved, acicular hairs adaxially on the costae and segments; the toothed, narrow scales abaxially on the costae will separate M. subincisum from M. lunense. From M. atrogriseum and M. pulverulentum it differs by the shorter hairs along the costae and costules abaxially. Breedlove 22633 (DS), from Chiapas, is a much scalier variant of M. subincisum; the ultimate segments are also broader and shorter than typical.

7 6 . M E L PO M E N E Melpomene A. R. Sm. & R. C. Moran, Novon 2: 426. 1992. Type: Melpomene moniliformis (Lag. ex Sw.) A. R. Sm. & R. C. Moran [⬅ Polypodium moniliforme Lag. ex Sw.]. Epiphytic, epipetric, or terrestrial; rhizomes erect or short- to long-creeping; rhizome scales usually blackish or dark reddish brown en masse, strongly clathrate, lanceolate to ovatelanceolate, cordate basally and attached at a central point, Ⳳ

Reference Smith, A. R. & R. C. Moran. 1992. Melpomene, a new genus of Grammitidaceae (Pteridophyta). Novon 2: 426–432.

Key to the Mexican Species of Melpomene 1. Rachises glabrous adaxially. 2. Segments ascending with respect to the rachises. 3. Sori lacking setae among the sporangia. .............................................................................................................................................. 1. M. deltata. 3. Sori with setae among the sporangia. .................................................................................................................................. 9. M. zempoaltepetlensis. 2. Segments perpendicular to the rachises. 4. Rhizomes suberect, the scales 2–3 mm long, blackish; stipes 0–1 cm long. ........................................................................... 5. M. melanosticta.

390

MELPOMENE

4. Rhizomes short- to long-creeping, the scales 3–6 mm long, reddish brown; stipes (1–)2–11 cm long. 5. Sori and abaxial surface of the fertile segments setose, the setae 2–3 times the length of the sporangia; segments ca. 2 times longer than wide. ........................................................................................................................................................... 3. M. flabelliformis. 5. Sori and abaxial surface of the fertile segments glabrous or, rarely sparsely setose and the setae only slightly longer than the sporangia; segments about as wide as long, or sometimes wider than long. ............................................................ 6. M. moniliformis. 1. Rachises setose adaxially. 6. Pinna (segment) length less than 3 times the width; rhizome scales 0.3–1 mm; rachises often with black clavate fungi abaxially; new plants formed from filamentous stolon-like roots, forming colonies. ............................................................................. Terpsichore anfractuosa. 6. Pinnae (segments) length (2–)3–10 times the width; rhizome scales 2–5 mm; rachises lacking black clavate fungi abaxially; new plants not formed on stolon-like roots. 7. Costae evident, black along the length of the pinnae. 8. Sori with a few setae from the receptacles; segments perpendicular to rachis, or almost so, the margins glabrous; stipes not setose abaxially (but with branched hairs). ............................................................................................................................... 2. M. firma. 8. Sori without setae from the receptacles; segments strongly ascending, the margins ciliolate; stipes setose abaxially. .. 4. M. leptostoma. 7. Costae not visible, inconspicuous, or only partially visible in some of the pinnae. 9. Segments lacking marginal setae. ................................................................................................................................... 9. M. zempoaltepetlensis. 9. Segments bearing at least a few marginal setae. 10. Segments usually acute at tips, ascending; rhizomes 2–6 cm long, the scales often spreading towards the tips; setae present on the surfaces of the sterile segments; 2700–3600 m. ..................................................................................... 7. M. pilosissima. 10. Segments obtuse, perpendicular to rachis or only slightly ascending; rhizomes 1–2 cm long, the scales appressed; setae absent on the abaxial surface of the sterile segments; 1200–2700 m. ............................................................. 8. M. xiphopteroides.

1. MELPOMENE DELTATA Melpomene deltata (Mickel & Beitel) A. R. Sm. & R. C. Moran, Novon 2: 429. 1992. Figs. 190C, D. Grammitis deltata Mickel & Beitel, Mem. New York Bot. Gard. 46: 198. 1988. Type. Mexico. Oaxaca: Distrito Mixe, NW slope of Cerro Zempoalte´petl, trail from Yacoche to Totontepec, Mickel 4637a (NY!).

Rhizomes creeping, 1–1.5 mm diam.; rhizome scales blackish to dark red-brown, 2–3 mm long, appressed, acute at tips; fronds clumped; stipes brown to black, 1.5–4 cm long, ca. 1⁄6 the frond length, glabrous except for scattered branched hairs 0.3 mm long; blades linear-elliptic, 13–16 ⫻ 0.9–1.2 cm, gradually tapering at both ends; rachises glabrous on both sides, or with rare transparent acicular hairs 0.1 mm long; pinnae mostly 20–30 pairs, obliquely deltate, 6–7 mm long, 1–2.5 times longer than wide, ascending, basiscopic side ca. 40–50⬚ to the rachis, decurrent, acroscopic side 70–90⬚, rounded at tips; costae visible and blackish abaxially; indument absent abaxially and along pinna margins, or nearly so; sori 4–6 pairs per pinna, lacking setae among sporangia.

Distribution. Epiphytic or occasionally epipetric in cloud forests; 2400–3000 m. Mexico; Hond, Salv. Specimens Examined. Chis (Breedlove 40337A, DS; Heath & Long 299, CAS; Mickel 1247d, NY). Oax (Breedlove 65943A, CAS; Mickel 4894, NY, UC).

Melpomene deltata is close to M. moniliformis and M. flabelliformis and grows with them, but differs in its glabrous nature (no setae in sori or on rachis) and segment shape (obliquely deltate, not narrowly oblong). Melpomene moniliformis is also nearly glabrous, but it is more often terrestrial, has short, rounded (not deltate) segments 2–4 mm long, perpendicular to the rachis, and sometimes has sparse soral setae. Melpomene deltata is similar to M. melanosticta in its lack of hairs, but its segments are more ascending and deltate. In Chiapas, M. deltata (Mickel 1247d; Breedlove 40337A) has been collected with M. moniliformis (Mickel 1247a, NY; Breedlove 40337B), M. zempoaltepetlensis (Mickel 1247b), and M. flabelliformis (Mickel 1247c). In Oaxaca, M. deltata (Breedlove 65943A) grows with M. moniliformis (Breedlove 65943B).

MELPOMENE FLABELLIFORMIS

2. MELPOMENE FIRMA Melpomene firma (J. Sm.) A. R. Sm. & R. C. Moran, Novon 2: 430. 1992. Figs. 189E–G. Polypodium firmum Klotzsch, Linnaea 20: 378. 1847, hom. illeg., non Kaulf., 1827. Ctenopteris firma J. Sm., Hist. Fil. 184. 1875. Grammitis firma (J. Sm.) C. V. Morton, Contr. U.S. Natl. Herb. 38: 110. 1967. Lectotype (inferentially chosen by Looser, Revista Univ. (Santiago) 36(1): 75. 1951). Guyana. Schomburgk 1170 (B, not seen; isolectotype K; frag. US!). Polypodium aromaticum Maxon, Proc. U.S. Natl. Mus. 27: 743. 1904. Ctenopteris aromatica (Maxon) Copel., Philipp. J. Sci. 84: 418. 1955 [1956]. Grammitis aromatica (Maxon) Proctor, Brit. Fern Gaz. 9: 218. 1965. Type. Jamaica. Blue Mt. Peak, Underwood 1449 (NY!; isotype US!).

Rhizomes suberect, 2–4 mm diam.; rhizome scales reddish brown, 3–5 mm long, appressed, acuminate at tips; fronds clumped; stipes dark brown to blackish, mostly (2–)4–8 cm long, 1 ⁄5–1⁄3 the frond length, glabrescent abaxially, setose adaxially, the setae red-brown, 0.1–0.4 mm long, also with a few branched lax pale reddish hairs on both sides; blades mostly 10–20(–27) ⫻ 2–6 cm, elliptic to oblanceolate, often completely 1-pinnate proximally and even in the middle, proximal several pairs of pinnae gradually reduced, gradually to subabruptly tapering distally; rachises setose on both sides, the setae 0.3–1 mm long, hairs absent; pinnae ca. 30–35 pairs, linear, mostly 15–30 mm long, 6–12 times longer than broad, perpendicular to the rachis and often upcurved, slightly excised basiscopically and sursumcurrent acroscopically, obtuse to subacute at the tips; costae readily visible and black abaxially; indument abaxially of sparse to numerous setae 1 mm long on costae, with veins, laminae between veins, and pinna margins glabrous; sori to ca. 15 pairs per pinna, sporangia intermixed with setae in and around the sori.

391

ially and the broader, 1-pinnate blades with more widely separated proximal pinnae. Melpomene firma is one of the most strongly aromatic species in the genus. Proctor (1985) reported the aroma persisting in dried specimens nearly a hundred years old.

3. MELPOMENE FLABELLIFORMIS Melpomene flabelliformis (Poir.) A. R. Sm. & R. C. Moran, Novon 2: 430. 1992. Figs. 190G, H. Polypodium flabelliforme Poir. in Lamarck, Encycl. 5: 519. 1804. Grammitis flabelliformis (Poir.) C. V. Morton, Contr. U.S. Natl. Herb. 38: 57. 1967. Xiphopteris flabelliformis (Poir.) Schelpe, Bol. Soc. Brot., se´r. 2, 41: 217. 1967. Type. Re´union. Commerson s.n. (P-JU1098C) Polypodium rigescens Bory ex Willd., Sp. Pl., ed. 4, 5(1): 183. 1810. Ctenopteris rigescens (Bory ex Willd.) J. Sm., Hist. Fil. 184. 1875. Xiphopteris rigescens (Bory ex Willd.) Alston, Bol. Soc. Brot., se´r. 2, 30: 27. 1956. Grammitis rigescens (Bory ex Willd.) Lellinger, Proc. Biol. Soc. Wash. 89: 383. 1985. Type. Re´union. Bory s.n. (B-W 19668; isotype, P-JU 1098C). Grammitis katasophistes Mickel & Beitel, Mem. New York Bot. Gard. 46: 200. 1988. Type. Mexico. Oaxaca: Distrito Mixe, NW slope of Cerro Zempoalte´petl, trail from Yacoche to Totontepec, Mickel 7637b (NY!). Grammitis andicola Stolze, Fieldiana, Bot., n.s., 32: 103. 1993. Type. Peru. Amazonas: Prov. Chachapoyas, Cerros Calla Calla, Hutchison & Wright 5826 (F; isotypes GH!, UC!).

Rhizomes long-creeping, 1.5–2 mm diam.; rhizome scales reddish brown, 3–4 mm long, appressed, acute at tips; fronds distant; stipes castaneous, gray-brown or black, 1–6(–11) cm long, 1⁄8–1⁄3 the frond length, lacking setae but with very short 0.3 mm pale branched hairs; blades linear, 10–20 ⫻ 1–1.6 cm, slightly reduced at bases, gradually tapering distally; rachises abaxially bearing scattered reddish setae 1–1.5 mm long and some smaller branched hairs, glabrous adaxially; pinnae ca. 25–45 pairs, oblong, 4.5–7 mm long, ca. 2 times longer than wide, perpendicular to rachises, obtuse at tips; costae not visible abaxially; indument abaxially of sparse setae 0.8–1 mm long on costae, segment margins with rare twisted hairs 0.3 mm long; sori 1–3 pairs per pinna, sporangia intermixed with reddish setae 0.5–1 mm long.

Distribution. Epiphytic in cloud forests; 1700–2300 m. Mexico; Guat, Hond, Salv, CR, Pan; Jam; Col, Ven, Guy, Ec, Peru, Bol. Selected Specimens Examined. Chis (Breedlove 22657, 25420 p.p., 28212 p.p., 29970 p.p., 30093, DS, 30158, DS, MEXU, 32642, 33665, DS). Oax (Mickel 1008, US; Mickel 4287, NY p.p.).

This species is most similar to M. xiphopteroides, and small forms of M. firma can be confused with that species. Melpomene firma can usually be distinguished by the non-setose stipes abax-

Distribution. Epiphytic in wet montane forests and cloud forests; 2150–2800 m. Mexico; CR, Pan; Hisp; Col, Ven, Ec, Peru, Bol, s Braz; Africa, Madagascar, Re´union.

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Selected Specimens Examined. Chis (Breedlove 22070B, DS; Mickel 1247c, NY). Oax (Hallberg 847a, NY, US; Mickel 3823b, 4325, 5251, 6784b, 6786, NY; Smith 470, UC p.p.).

Melpomene flabelliformis is very close to M. moniliformis and M. deltata (see discussion under M. moniliformis). A specimen from Chiapas (Mickel 1247c) was growing with M. deltata, M. moniliformis, and M. zempoaltepetlensis, at 2850 meters. Smith 470, from Oaxaca, was growing mixed with M. moniliformis. Grammitis andicola, recently described from Peru and attributed also to Colombia, seems to be exactly Melpomene flabelliformis from Africa and offshore islands. Furthermore, it is conspecific with specimens of M. flabelliformis from Mexico and Central America. What Stolze called Grammitis flabelliformis from Peru (and matched by some specimens from Costa Rica to Bolivia), is perhaps a different species, distinguished by narrower, more acuminate, and more spreading scales and smaller fronds. However, this group is sufficiently problematic taxonomically that confident conclusions are impossible at present.

4. MELPOMENE LEPTOSTOMA Melpomene leptostoma (Fe´e) A. R. Sm. & R. C. Moran, Novon 2: 430. 1992. Figs. 190A, B. Polypodium leptostomum Fe´e, Me´m. Foug. 7: 58, t. 21, f. 2. 1857. Ctenopteris leptostoma (Fe´e) Copel., Philipp. J. Sci. 84: 421. 1955 [1956]. Grammitis leptostoma (Fe´e) F. Seym., Phytologia 31: 179. 1975. Type. Mexico. [Veracruz:] Near Orizaba, Schaffner 210 (RB; not found at P). Polypodium productum Maxon, Contr. U.S. Natl. Herb. 13: 11. 1909. Type. Guatemala. Alta Verapaz: Coba´n, Tu¨rckheim II.1347 (US!).

Rhizomes short-creeping to suberect, compact, 1–2 mm diam.; rhizome scales blackish, 2–4 mm long, appressed, acute at tips; fronds tightly clumped; stipes dark brown to black, 1–4 cm long, ca. 1⁄6–1⁄20 the frond length, with stiff reddish setae 0.3–0.8 mm long and scattered, branched hairs 0.3 mm long; blades 11–20(– 35) ⫻ 1.3–1.7 cm, narrowly elliptic, very gradually tapering at both ends; rachises with stiff setae 0.3–1 mm long on both sides; pinnae to ca. 120 pairs, linear, 6–11 mm long, to 6 times longer than wide, decidedly ascending, slightly decurrent at bases, acute at tips; costae brown to black abaxially, but color not visible at pinna bases; indument absent except for occasional hairs 0.1–0.3 mm long on costae and pinna margins; sori mostly 4–6 pairs per pinna, lacking setae among the sporangia.

Distribution. Epiphytic in cloud forests; 1000–2050(–2700) m. Mexico; Guat. Selected Specimens Examined. Chis (Hardison 92A, UC). Hgo (Nava P. s.n., FCME). Oax (Cedillo T. 919, MO; Mickel 5737, ENCB, NY, UC). Pue (Ramos & Ferna´ndez R-123, CHAPA, NY; Ventura A. 8655, ENCB, LL). Ver (Lyonnet 550500011, MEXU; Mu¨ller 32, NY). Unverified, Doubtful, or Mistaken Reports. Gro (Lorea 1990, 4007, 4558b, cited by Lorea-Herna´ndez & Vela´zquez M., 1998, but not verified). Me´x (reported by Mickel & Beitel, 1988, but not verified; Tejero-Dı´ez 2377, IZTA, cited by Tejero-Dı´ez & Arreguı´n-Sa´nchez, 2004, but the specimen, although juvenile and sterile, is most likely M. pilosissima, and not M. leptostoma).

The glabrous pinna margins, blade surfaces, and sori of M. leptostoma, as well as the thinner-textured, more ascending segments and blackish costae abaxially, readily separate it from M. xiphopteroides. From M. pilosissima, which has similarly ascending segments, M. leptostoma is easily distinguished by its glabrous blades. See additional discussion under M. xiphopteroides.

5. MELPOMENE MELANOSTICTA Melpomene melanosticta (Kunze ex Klotzsch) A. R. Sm. & R. C. Moran, Novon 2: 430. 1992. Figs. 189H, J. Polypodium melanostictum Kunze ex Klotzsch, Linnaea 9: 44. 1834. Ctenopteris melanosticta (Kunze ex Klotzsch) Copel., Philipp. J. Sci. 84: 385. 1955 [1956]. Grammitis melanosticta (Kunze ex Klotzsch) F. Seym., Phytologia 31: 179. 1975. Type. Peru. Poeppig s.n. (LZ, destroyed).

Rhizomes short-creeping to usually suberect, compact, 2–3 mm diam.; rhizome scales dark brown to blackish, 2–3 mm, appressed, acute at tips; fronds clumped; stipes dark brown, 0–1 cm long, 1⁄20–1⁄10 the frond length, lacking setae on both sides, hairy abaxially, the hairs pale reddish, 0.1–0.2 mm long, generally branched, lax; blades 10–20 ⫻ 0.6–1(–1.5) cm, linear-elliptic, gradually tapering at both ends; rachises lacking setae on both sides, sparsely hairy to glabrescent abaxially, the hairs 0.1–0.2 mm long, generally branched, pale reddish, lax; pinnae 22–50 pairs, oblong, to 7 mm long, 1.5–2.5 times as long as wide, mostly spreading, rounded at tips; costae not visible abaxially; indument

MELPOMENE PILOSISSIMA

393

absent abaxially; sori mostly 2–4 pairs per segment, lacking setae among the sporangia.

Distribution. Epiphytic in montane rain forests; 800–1000 m. Mexico; Guat, CR; Cuba, Hisp; Col, Ven, Ec, Peru, Braz, Bol. Specimens Examined. Chis (Breedlove 29900, DS p.p., 29902, 33014, DS; Martı´nez S. 18679, MEXU, UAMIZ).

This species most resembles M. moniliformis but is found at lower elevations, is always epiphytic (vs. usually terrestrial or epipetric), has suberect (vs. creeping) rhizomes, and has blades tapering gradually at the base, nearly or quite to the rhizomes. The blade texture of M. melanosticta is also less coriaceous.

6. MELPOMENE MONILIFORMIS Melpomene moniliformis (Lag. ex Sw.) A. R. Sm. & R. C. Moran, Novon 2: 430. 1992. Figs. 190E, F. Polypodium moniliforme Lag. ex Sw., Syn. Fil. 33. 1806. Ctenopteris moniliformis (Lag. ex Sw.) J. Sm., Hist. Fil. 184. 1875. Grammitis moniliformis (Lag. ex Sw.) Proctor, Brit. Fern Gaz. 9: 219. 1965. Type. Peru. s. coll. [Ruiz & Pavo´n?] s.n. (S not seen). Polypodium moniliforme Lag. ex Sw. var. major Liebm., Mexic. Bregn. 196 (reprint 44). 1849. Lectotype (chosen by Smith, 1981: 123). Mexico. Veracruz: Pico de Orizaba, Liebmann s.n. [Pl. Mex. 2533, Fl. Mex. 183] (C!).

Rhizomes moderately short- to long-creeping, 1–3 mm diam.; rhizome scales dark red-brown, 3–4 mm long, appressed, acute at tips; fronds somewhat distant to clumped; stipes dark brown to blackish, 1–5 cm long, 1⁄8–1⁄4 the frond length, lacking setae but with abundant, pale, branched hairs 0.2–0.3 mm long; blades linear, 3–15 ⫻ 0.5–1 cm, slightly reduced proximally, gradually tapering distally; rachises lacking setae, with a few scattered small glandular hairs abaxially, glabrous adaxially; pinnae 22–40 pairs, semicircular to deltate, 2–4 mm long, often broader than long, spreading to very slightly ascending, obtuse at tips, alternate and often appearing zig-zag; costae not visible abaxially; indument abaxially absent or of very sparse short hairs on costae; sori 1– 2(–3) pairs per pinna, lacking setae among the sporangia, rarely with 1–3(–10) setae; 2n⫽ca. 148 (Jam).

Distribution. Terrestrial and on mossy rocks at high elevations, fir forests; 2150–4000 m. Mexico; Guat, Hond, Salv, CR, Pan; Jam, Hisp; Col, Ven, Ec, Peru, s Braz, Bol. Cited from Nicaragua by Smith (1981: 124), but this is on the basis of the citation by Go´mez (1976: 47); we have been unable to verify the occurrence in Nicaragua. Selected Specimens Examined. Chis (Matuda 2380, K, MEXU, US). DF (Lyonnet 3244, MEXU). Gro (Hinton 14294, ARIZ, LL, MO, NY, TEX). Hgo (Lyonnet 510900006, CAS, MEXU). Jal (McVaugh 11649, MEXU). Mor (Sa´nchez 463, MEXU). Mich (Zamudio 3917, ENCB, IEB). Oax (Pringle 6194, MO, NY, UC). Tlax (Acosta P. & Sa´nchez 2970, CIB). Ver (Mu¨ller 29, NY). Unverified, Doubtful, or Mistaken Reports. Me´x (reported by Mickel & Beitel, 1988, but not verified; Sandoval 131, UAMIZ, cited by Tejero-Dı´ez & Arreguı´n-Sa´nchez, 2004, from the Me´xico/Morelos border, but not verified).

This species, M. flabelliformis, and M. deltata form a species complex lacking hairs on the adaxial surface of the rachises and having creeping rhizomes. Melpomene flabelliformis also has pinnae perpendicular to rachises, but the numerous hairs in the sori, scattered hairs on the abaxial side of the rachises, and longer pinnae contrast with the 0–3 hairs per sorus, essentially glabrous abaxial surface of the rachises, and shorter pinnae of M. moniliformis. The strongly ascending, deltate segments and totally glabrous sori of M. deltata distinguish that taxon from the more common M. moniliformis. Breedlove 40337 (DS), collected on rocks, is a mixed collection of this species and M. deltata, and highlights their differences. Matuda 2889 (MEXU), from Volca´n Tacana´ in Chiapas, is aberrant in having 5–10 setae ca. 1–1.5 mm long within nearly all sori and also a few similar setae on the abaxial blade tissue of non-soriferous segments; in blade cutting, however, it is not unlike more typical specimens of M. moniliformis.

7. MELPOMENE PILOSISSIMA Melpomene pilosissima (M. Martens & Galeotti) A. R. Sm. & R. C. Moran, Novon 2: 431. 1992. Figs. 190J, K.

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MELPOMENE

Polypodium pilosissimum M. Martens & Galeotti, Me´m. Foug. Mexique 39, pl. 9, f. 2. 1842. Ctenopteris pilosissima (M. Martens & Galeotti) Copel., Philipp. J. Sci. 84: 390. 1955 [1956]. Grammitis pilosissima (M. Martens & Galeotti) C. V. Morton, Contr. U.S. Natl. Herb. 38: 114. 1967. Lectotype (chosen by Smith, 1981: 124). Mexico. Oaxaca: Tonaguia, Galeotti 6379 (BR!, photo US!; isolectotypes BR!, K!; frag. US!).

Rhizomes short-creeping or ascending, 1–2 mm diam.; rhizome scales blackish, 5–7 mm long, long-acuminate and often spreading at tips; fronds somewhat distant to clustered; stipes gray-brown to black, 2–9 cm long, 1⁄4–1⁄3 the frond length, with dense reddish setae 2–2.5 mm long, intermixed with pale, inconspicuous, multibranched hairs 0.3 mm long; blades elliptic, 7–15(–20) ⫻ 1–2(–3) cm, abruptly tapering proximally, gradually tapering distally; rachises with abundant reddish setae 1.5–2.5 mm long on abaxial and adaxial sides, short pale hairs few and very inconspicuous; pinnae ca. 25–50 pairs, linear-oblong, 7–14 mm long, 3–7 times longer than wide, usually ascending (more so than M. xiphopteroides), obtuse to usually acute at tips; costae barely visible and blackish abaxially; indument abaxially on costae, laminae, and pinna margins of abundant, reddish setae 1–2 mm long, also with sparse short pale hairs; sori 4–7 pairs per pinna, sporangia mixed with setae.

Distribution. Epiphytic and terrestrial among rocks in cloud forests; (2000–)2700–4400 m. Mexico; Guat, Hond, CR, Pan; Col, Ven, Sur, Ec, Peru, Bol (Kessler 7284, UC), Braz.

similar. In this character, they approach closely M. zempoaltepetlensis (which see), but they seem to have thicker blade texture and more strongly setose stipes and blades than the latter. Further study is needed. Stevens 11521 (UC), cited as this species from Nicaragua (in Davidse et al., 1995), is M. xiphopteroides.

8. MELPOMENE XIPHOPTEROIDES Melpomene xiphopteroides (Liebm.) A. R. Sm. & R. C. Moran, Novon 2: 431. 1992. Figs. 189K, L. Polypodium xiphopteroides Liebm., Mexic. Bregn. 196 (reprint. 44). 1849. Grammitis xiphopteroides (Liebm.) A. R. Sm., Amer. Fern J. 70: 26. 1980. Lectotype (chosen by Smith, 1981: 125). Mexico. Veracruz: “Hac. de Mirador,” Liebmann s.n. [Pl. Mex. 2548, Fl. Mex. 189] (C!). Polypodium rigens Maxon, Proc. U.S. Natl. Mus. 27: 741. 1904. Ctenopteris rigens (Maxon) Copel., Philipp. J. Sci. 84: 422. 1955 [1956]. Grammitis rigens (Maxon) Proctor, Brit. Fern Gaz. 9: 219. 1965. Type. Jamaica. John Crow Peak, Maxon 1346 (US!). Ctenopteris megaloura Copel., Philipp. J. Sci. 84: 391. 1955 [1956]. Type. Guatemala. Alta Verapaz: Coba´n, Tu¨rckheim II 1855 (US!).

Rhizomes short-creeping, 1–2 mm diam.; rhizome scales blackish, appressed, 4–7 mm long, long-acuminate at tips; fronds somewhat distant to clustered; stipes gray-brown to black, 2–9 cm long, 1⁄4–1⁄3 the frond length, with dense reddish brown setae 2–2.5 mm long, intermixed with pale, inconspicuous, multibranched hairs 0.3 mm long; blades linear-elliptic, 7–15(–20) ⫻ 1–2(–3) cm, abruptly tapering proximally, gradually tapering distally; rachises with abundant reddish setae 1.5–2.5 mm long on both sides, short pale hairs few and very inconspicuous; pinnae ca. 25–50 pairs, linear-oblong, 7–14 mm long, usually 3–7 times longer than wide, usually spreading, obtuse to acute at tips; costae barely visible and blackish abaxially; indument abaxially on costae, laminae, and pinna margins of sparse to numerous reddish setae 1–2 mm long, but setae often completely absent on sterile pinnae, axes abaxially also with sparse short pale hairs; sori 4–7 pairs per pinna, sporangia mixed with setae.

Selected Specimens Examined. Chis (Breedlove 32030, DS, 53604, CAS, ENCB, NY). DF (Rzedowski 37672, IEB). Gro (Hinton 14326, LL, MO, NY; Rzedowski 18602, ENCB, NY). Me´x (Lyonnet 1480, CAS, FCME, MO, UC, US; Pringle 4288, DS, MEXU, MO, NY, UC, US). Mich (Dı´az B. & Zamudio 5046, XAL). Oax (Mickel 4886, NY, UC; Smith 478, UC). Ver (Beaman 2173, MEXU; Mu¨ller 28, NY). Unverified, Doubtful, or Mistaken Reports. Jal (Wilbur 2000, MICH, cited by Va´squez G. et al., 1995, but not verified).

Melpomene pilosissima is most often confused with M. xiphopteroides. The former differs especially by the acute, strongly ascending segments, more spreading rhizome scales, presence of setae even on sterile segments abaxially, and occurrence at higher elevations. Specimens of M. pilosissima from Guatemala, Costa Rica, and Panama, and most (but not all) specimens from South America lack setae along the segment margins, but are otherwise very

Distribution. Epiphytic in wet montane forests; 1150–3050 m. Mexico; Guat, Hond, Salv, Nic (Stevens 11521, UC), CR, Pan; Gr Ant; Col, Ven, Ec, Peru, Bol (Kessler 7284, UC).

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395

Selected Specimens Examined. Chis (Hardison 92B, UC). Gro (Breedlove 65114, CAS; Lorea 1626, FCME, 2094, IEB; Rzedowski 18637, ENCB). Oax (Davidse et al. 30244, MEXU; Mickel 6704, UC; Pe´rez C. C46, MEXU). Ver (Lyonnet 541200020, MEXU; Purpus 213, US).

At least three of the four specimens cited from Veracruz as this species by Palacios-Rios (1992) are M. leptostoma. The presence of setae on the adaxial surface of the rachises and the clumped nature of the fronds distinguish M. xiphopteroides from the group of M. moniliformis, M. flabelliformis, and M. deltata, which have more distant fronds and no hairs on the rachises adaxially. Some plants of M. xiphopteroides become fertile at a very small size, and the narrow fronds are often mistaken for members of the M. moniliformis group. The glabrous pinna margins, abaxial blade surface, and sori of M. leptostoma readily separate that species from M. xiphopteroides. Melpomene pilosissima differs in having pinna margins with hairs over the entire length (vs. 0–4 hairs in distal 1⁄4 of segments in M. xiphopteroides), longer rhizome scales, and evenly spaced laminar hairs (vs. hairs clustered in the sori). Ctenopteris megaloura has generally been placed as a synonym of Melpomene pilosissima (e.g., by Stolze, 1981, and Lellinger, 1989), but the type is identical to the broad form of M. xiphopteroides.

9. MELPOMENE ZEMPOALTEPETLENSIS Melpomene zempoaltepetlensis (Mickel & Beitel) A. R. Sm., Novon 5: 21. 1995. Figs. 189A–D. Grammitis zempoaltepetlensis Mickel & Beitel, Mem. New York Bot. Gard 46: 205. 1988. Type. Mexico. Oaxaca: Dist. Mixe, NW slope of Cerro Zempoalte´petl, trail from Yacoche to Totontepec, Mickel 4638 (NY!).

Rhizomes short- to long-creeping, 1–2 mm diam.; rhizome scales red-brown, 2–4 mm long, appressed, acute at tips; fronds somewhat distant; stipes brown to red-brown, 2–4 cm long, 1⁄5– 1⁄3 the frond length, with red-brown setae 1–1.5 mm long; blades 10–15 ⫻ 1–2 cm, tapering gradually at both ends; rachises glabrescent or with scattered red-brown setae mostly 0.3–1 mm long on both sides; pinnae ca. 15–30 pairs, oblong to linear, slightly falcate, ascending, acute at tips; costae obscure abaxially; indument abaxially of sparse to frequent red-brown setae ca. 1–2 mm long, setae absent on pinna margins; sori 2–5 pairs per pinna, sporangia mixed with red-brown twisted hairs 1–1.5 mm long.

Distribution. Epiphytic in cloud forests; 2350–3600 m. Mexico. Selected Specimens Examined. Chis (Breedlove 22107, DS, 22928B, 24310, 29353, DS, MEXU, NY; Matuda 2379, MEXU, 2379A, NY, US; Mickel 1247b, NY). Oax (Hallberg 847b, US; Mickel 4597b, UC, 4598b, NY).

This species was treated as a synonym of M. pilosissima by Smith and Moran (1992), but mixed collections of the two (Mickel 4598b; Breedlove 22928B) and of M. moniliformis and M. zempoaltepetlensis (Mickel 4597b), show clear differences, even when the species grow together. Breedlove 22073, DS, and 22107 are atypical in lacking or nearly lacking setae on the rachises adaxially. They were cited as aberrant Grammitis xiphopteroides by Smith (1981).

77. METAXYA Metaxya C. Presl, Tent. Pterid. 59. 1836. Type: Metaxya rostrata (Kunth) C. Presl [⬅ Aspidium rostratum Kunth, a new name for Polypodium rostratum Willd. (1819), non Burm.f., 1768]. Amphidesmium J. Sm., Ferns Brit. For. 167. 1866 (Schott, Gen. Fil. sub Trichopteris, 1834, nom. nud.). Type: Amphidesmium blechnoides (Hook.) J. Sm. [⬅ Alsophila blechnoides Hook.] ⫽ Metaxya rostrata (Kunth) C. Presl.

Terrestrial; rhizomes stout, creeping, solenostelic, hairy (scales absent); fronds large, pinnate; costae grooved; veins free, closely parallel, occasionally forked; stomates with 3 subsidiary cells, one on the side of each guard cell and a third on the side of the smaller one; sori dorsal, one to several per vein, round or nearly so, exindusiate; receptacles flat or nearly so; sporangia numerous, maturing together, mixed with copious hairs, pedicel short, of 4 rows of cells; annuli slightly oblique; spores 64 per sporangium, Ⳳ globose with a trilete scar, with a granulate perine; x⫽ca. 94– 96. Metaxya is a genus of two species of the New World tropics. Its affinities are obscure, but it may be most closely related to Lophosoria, both genera being placed in the Cyatheaceae by Tryon (1970). Roy and Holttum (1965) suggested a relationship to Culcita or Thyrsopteris. Molecular data (Wolf et al., 1999; see

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citation under Lophosoria) show Metaxya to be at the base of the tree fern clade. References Riba, R. 1993. Pteridofitas. Familia Metaxyaceae. Flora de Me´xico 6(1): 17–19; Roy, S. K. & R. E. Holttum. 1965. Cytological and morphological observations on Metaxya rostrata (H.B.K.) C. Presl. Amer. Fern J. 55: 158–164; Smith, A. R., H. Tuomisto, K. M. Pryer, J. S. Hunt & P. G. Wolf. 2001. Metaxya lanosa, a second species in the genus and fern family Metaxyaceae. Syst. Bot. 26: 480– 486; Tryon, R. 1970. The classification of the Cyatheaceae. Contr. Gray Herb. 200: 3–53.

1. METAXYA ROSTRATA Metaxya rostrata (Kunth) C. Presl, Tent. Pterid. 60, pl. 1, f. 5. 1836. Figs. 191A, B. Polypodium rostratum Willd., Sp. Pl., ed. 4, 5(1): 193. 1810, hom. illeg., non Burm.f., 1768. Aspidium rostratum Kunth in Humboldt, Bonpland & Kunth, Nov. Gen. Sp. 1: 12. 1815. Alsophila rostrata (Kunth) Mart., Icon. Pl. Crypt. 64, pl. 39. 1834. Amphidesmium rostratum (Kunth) J. Sm., London J. Bot. 1: 668. 1842. Alsophila blechnoides Hook., Sp. Fil. 1: 35. 1844. Amphidesmium blechnoides (Hook.) J. Sm., Ferns Brit. For. 167. 1866. Type. Venezuela. Terr. Amazonas: near S. Antonio Javita, “in interioribus Americae meridionalis regionibus prope Jative,” Humboldt & Bonpland 966 (B-Willd. 19691!).

Fronds 1.5–2 m, with stipes to nearly 1 cm diam., pinnate to ca. 16 lateral pairs and a conform terminal one, basal one stalked to ca. 10 mm, to ca. 30 ⫻ 3.5 cm, cuneate at base, with cartilaginous entire to repand margins and serrate apices; laminar tissue subcoriaceous, glabrous; sori mostly nearer costa than margin; 2n⫽190–192 (Sur).

Distribution. Tropical rain forests, evergreen rain forests; 180– 300. Mexico; Guat, Bel, Hond, Nic, CR, Pan; Hisp, Cuba, L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, n Braz, Bol. Specimens Examined. Chis (Breedlove 34100, DS; Martı´nez S. 13956, MEXU, UAMIZ; Martı´nez & Riviere 2031, CAS).

7 8 . M I C R O GR A M M A Microgramma C. Presl, Suppl. Tent. Pterid. 213, t. 9, f. 7. 1836. Type: Microgramma persicariifolia (Schrad.) C. Presl [⬅ Polypodium persicariifolium Schrad.]. Craspedaria Link, Fil. Spec. 117. 1841. Type: Craspedaria vacciniifolia (Langsd. & Fisch.) Link [⬅ Polypodium vacciniifolium Langsd. & Fisch.] ⬅ Microgramma vacciniifolia (Langsd. & Fisch.) Copel. Anapeltis J. Sm., Cult. Ferns 5. 1857. Type: Anapeltis lycopodioides (L.) J. Sm. [⬅ Polypodium lycopodioides L.] ⬅ Microgramma lycopodioides (L.) Copel. Lopholepis (J. Sm.) J. Sm., London J. Bot. 1: 195. 1842, hom. illeg., non Decne., 1839 (Poaceae). Goniophlebium sect. Lopholepis J. Sm., J. Bot. (Hook.) 4: 56. 1841. Type: Lopholepis piloselloides (L.) J. Sm. [⬅ Polypodium piloselloides L.] ⬅ Microgramma piloselloides (L.) Copel.

Epiphytic, occasionally terrestrial; rhizomes long-creeping, often branched; rhizome scales concolorous to bicolorous, nonclathrate, imbricate, peltately attached, surfaces lacking hairs, margins entire to dentate; fronds simple, distant, monomorphic to dimorphic (fertile longer and narrower than sterile), stipitate, elliptic to oblong, articulate, phyllopodia obsolete to present, margins entire to undulate; blades chartaceous to subcoriaceous, glabrous or with filamentous or round (in M. percussa) scales; venation netted, with one to several free included veins in areoles; sori round to elongate, exindusiate, within costal areoles in one row on each side of costae, on end of free, included veins or at junction of several included veins (compital); paraphyses present, consisting of filiform scales or branched scales with filiform tips; sporangia glabrous; spores bilateral, verrucate to coarsely tuberculate or rugose; x⫽37. Microgramma is a segregate of Polypodium s.l. and comprises ca. 20 species in the New World tropics and 1 or 2 species in Africa. The genus is usually distinguished by the subdimorphic to dimorphic fronds, reticulate veins, peltate non-clathrate rhizome scales, and uniseriate sori with paraphyses of narrow scales. The generic limits of Microgramma, as well as the other segregate genera, are in need of careful study. Polypodium percussum Cav. is often placed in Pleopeltis. Smith (1981) considered it might be closer to Microgramma because of its dense, branched receptacular scales with hair-like segments, and it is considered a Microgramma in this flora, largely on the basis of the lack of peltate scales in the sorus. In addition, molecular data ally it with other species of Microgramma, and not with Pleopeltis (Schneider et al., unpubl. data). Copeland included Polypodium muenchii in Microgramma, and Smith (1981) included it in Pleopeltis, but it is aberrant in both genera; we place it in Polypodium s.l. in this treatment. It is sometimes treated as Microphlebodium muenchii (Christ) L. D. Go´mez.

Key to the Mexican Species of Microgramma 1. Blades essentially glabrous. 2. Rhizomes 1.5–2(–3) mm diam., scales 5–6(–10) mm long, concolorous; fronds monomorphic to subdimorphic, blades 3–12 ⫻ 0.4–1.6 cm; soral scales 0.3 mm long, hair-like, inconspicuous or absent. .................................................................................... 1. M. lycopodioides. 2. Rhizomes 3–6 mm diam., scales 6–10 mm long, bicolorous with whitish tips; fronds dimorphic, sterile blades 5–12(–20) ⫻ 2–4 cm, fertile blades 8–20 ⫻ (0.8–)1.4–1.8 cm; soral scales 1–1.3 mm long, 1–2 cells wide, sparse, longer than sporangia. .................. 2. M. nitida.

MICROGRAMMA NITIDA

397

1. Blades scaly with numerous lanceolate or orbicular scales. 3. Fronds 12–22 cm long, monomorphic, lanceolate; abaxial surfaces with round scales (0.3 mm wide), adaxial surfaces glabrous; rhizomes 1–2 mm diam.; sori with dense, short filiform scales branching from bases into several hair-like segments. ................. 3. M. percussa. 3. Fronds 0.8–8.5 cm long, dimorphic, sterile lanceolate or ovate to orbicular, fertile lanceolate; abaxial and adaxial surfaces with lanceolate scales 1–1.5 mm long with hair-like tips; rhizomes 0.3–1 mm diam.; sori with unbranched acicular scales. 4. Sterile blades 0.8–2 cm long, orbicular to ovate; rhizomes 0.3–0.5 mm diam., with spreading scales. ............................ 6. M. tecta var. nana. 4. Sterile blades to ca. 6 cm long, usually lanceolate to ovate; rhizomes 0.5–1 mm diam., with tightly to weakly appressed or somewhat spreading scales. 5. Fertile blades 5–10 mm wide; mature sori well contained within the margins; rhizome scales tightly appressed. ........... 4. M. piloselloides. 5. Fertile blades 2–3(–4.5) mm wide; mature sori commonly projecting beyond the margins; rhizome scales somewhat appressed to somewhat spreading. ....................................................................................................................................................... 5. M. reptans.

1. MICROGRAMMA LYCOPODIOIDES Microgramma lycopodioides (L.) Copel., Gen. Filic. 185. 1947. Fig. 192L. Polypodium lycopodioides L., Sp. Pl. 2: 1082. 1753. Lectotype (chosen by Proctor, 1977: 338). LINN 1251.2, locality unknown (see also Jackson, 1912).

Rhizomes long-creeping, 1.5–2(–3) mm diam.; rhizome scales brown, concolorous, becoming bicolorous with age, 5–6(–10) mm long, lance-attenuate with acuminate tips, subappressed, peltate, margins denticulate; fronds monomorphic to subdimorphic (the fertile slightly smaller and often a little narrower), distant, chartaceous to subcoriaceous; stipes absent or nearly so; blades lanceolate, linear-lanceolate or elliptic-lanceolate, 3–12 ⫻ 0.4–1.6 cm, apices acute or rarely obtuse, bases cuneate, glabrous, with rare hair-like blade scales 1 mm long on rachises; sori round, medial, sporangia mixed with narrow scales 0.3 mm long, these few and inconspicuous, or seemingly absent; 2n⫽74 (CR, Cuba, Jam, PR).

Microgramma lycopodioides is distinct by its glabrous blades and slender rhizomes with concolorous scales. Reports of this species from Africa, Madagascar, and Mauritius pertain to the closely related, but specifically different, M. mauritiana (Willd.) Tardieu [synonym: M. owariensis (Desv.) Alston].

2. MICROGRAMMA NITIDA Microgramma nitida (J. Sm.) A. R. Sm., Proc. Calif. Acad. Sci., ser. 4, 40: 230. 1975. Figs. 192E–G. Phlebodium nitidum J. Sm., Bot. Mag. 72(Compendium): 13. 1846, non Polypodium nitidum Kaulf., 1824. Neotype (chosen by Smith, 1981: 149). “Collect. Hort. Kew, 1851” in J. Smith’s hand (BM!). Polypodium palmeri Maxon, Contr. U.S. Natl. Herb. 17: 600. 1917. Type. Mexico. Tamaulipas: near Go´mez Farias, Palmer 308 (US!; isotypes K!, NY!, UC!, US!).

Rhizomes long-creeping, 3–6 mm diam.; rhizome scales 6–10 mm long, bicolorous, reddish brown with white margins on acroscopic portion, lance-attenuate with hair-like tips, appressed to spreading, peltate, margins erose; fronds dimorphic, rarely fertile, distant, coriaceous; stipes usually absent, rarely to 5 mm long, glabrous; sterile blades lanceolate to ovate-oblong, 5–12(–20) ⫻ 2–4 cm; fertile blades lanceolate to elliptic-lanceolate, 8–20 ⫻ (0.8–)1.4–2.5 cm, bases rounded to cuneate, apices obtuse to acute, rarely acuminate, glabrous; sori round, medial, soral scales 1–2 cells wide, hair-like, longer than sporangia, 1–1.3 mm long, sparse.

Distribution. Epiphyte on tree trunks, lower montane rain forests; 100–800 m. Mexico; Guat, Bel, Hond, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol. Moran (in Davidse et al., 1995) also reported this species from Paraguay. Selected Specimens Examined. Chis (Breedlove 22501, DS, 33040, DS, MEXU, NY; Martı´nez S. 15944, MO, 24986, MEXU; Rovirosa 1040, PH; Sohns 1682, MEXU, US). Oax (Wendt 3996, CHAPA, NY, UC). Ver (Dorantes 3913, IEB, XAL; Va´zquez et al. 912, XAL; Wendt et al. 2596, CHAPA, ENCB, MEXU, NY). Unverified, Doubtful, or Mistaken Reports. Tab (reported by Smith, 1981, and Magan˜a, 1992, but not verified).

Distribution. Epiphyte on tree trunks at usually low elevation; 0–900(–2500) m. Mexico; Guat, Bel, Hond, Nic, CR, Pan; Jam, Barbados; Trin.

398

MICROGRAMMA

Selected Specimens Examined. Camp (Sanders 9737, UC). Chis (Davidse 9434, MEXU, UC). Hgo (Clark 6890, MO). Oax (Mickel 5811, ENCB, UC). Pue (Ventura A. 16041, ENCB). QR (Ucan 1410, MEXU, UC). Qro (Carranza et al. 4528, IEB, MEXU). SLP (Pringle 3355, DS, ENCB, LL, MEXU, MO, UC, US). Tab (Matuda 3304, MEXU, MICH, US). Tam (Palmer 308, MO, UC). Ver (Purpus 3817, UC). Yuc (Ucan & Burgos 1169, MEXU, UC).

This species closely resembles M. lycopodioides in its glabrous blades, but is easily distinguished by its thicker rhizomes with larger bicolorous scales.

3. MICROGRAMMA PERCUSSA Microgramma percussa (Cav.) de la Sota, Physis (Buenos Aires), Secc. C, 44(106): 28. 1986. Figs. 192M–O. Polypodium percussum Cav., Descr. Pl. 243. 1801. Pleopeltis percussa (Cav.) Hook. & Grev., Icon. Filic. 1: t. 67. 1828. Type. Peru [incorrectly stated as Marianas by Cavanilles]. Ne´e s.n. (MA, seen and commented on by Christensen, Dansk Bot. Ark. 9(3): 11. 1937).

Rhizomes long-creeping, 1–2 mm diam.; rhizome scales brown, concolorous, 1–2 mm long, lanceolate, entire; fronds monomorphic or only slightly dimorphic, coriaceous; stipes stramineous, 1⁄10–1⁄3 the frond length, mostly 1–5 cm long; blades lanceolate, 12–22 ⫻ 0.8–2.2 cm, apices acute to long-acuminate, bases very gradually cuneate, midribs with scattered lanceolate scales 1–2 mm long, adaxially with the veins often readily visible; indument abaxially of sparse round scales 0.3 mm diam. and rare lanceolate scales 1 mm long, margins fimbriate, indument adaxially absent; sori round, slightly sunken, with dense filiform hairs (scales) branching from the bases into several reddish brown hair-like segments, nearly obscuring sporangia; 2n⫽74 (CR, Peru).

See discussion under generic description for comments on taxonomic position. Microgramma percussa is distinct by its sori with dense cushion of hair-like trichomes, small round abaxial blade scales, and coriaceous blades, usually with long-acuminate apices. The venation of M. percussa is often readily visible in dried fronds adaxially, contrasting with other scaly-bladed species of Microgramma.

4. MICROGRAMMA PILOSELLOIDES Microgramma piloselloides (L.) Copel., Gen. Filic. 185. 1947. Figs. 192A, B. Polypodium piloselloides L., Sp. Pl. 2: 1083. 1753. Type. “Habitat in America meridionali,” LINN 1251.3 is this species, but, according to Jackson (1912) was added after 1755; Plumier and Petiver illustrations also cited.

Differs from M. reptans by: rhizome scales tightly appressed and more broadly lanceolate and without long-attenuate tips; sterile blades more obviously scaly; fronds subdimorphic (vs. strongly dimorphic in M. reptans); fertile blades elliptic, mostly 5–10 mm wide and with sori not overlapping the blade margins; 2n⫽148 (Jam, PR).

Distribution. Epiphytic in montane rain forests; 1400 m. Mexico; Guat; Gr & L Ant; Ven. Reported from Nicaragua and Costa Rica by Moran (in Davidse et al.,1995), but identity of specimens should be verified. Also reported from Colombia, Guyana, French Guiana, Ecuador, Peru, Bolivia, and Brazil by R. Tryon and Stolze (1993) and by Moran (in Davidse et al., 1995), but because of confusion with M. reptans and M. acatallela Alston, this added range is suspect. Specimens Examined. Chis (Breedlove 25140, DS, MEXU).

Distribution. Epiphytic, or occasionally epipetric; 100–800 (–1200) m. Mexico; Guat, Bel, Hond, Nic, CR, Pan; Col, Ven, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol. Selected Specimens Examined. Chis (Breedlove 48254, 50869, 57861, CAS, ENCB). Oax (Herna´ndez G. 445, MEXU, NY; Wendt et al. 3721, 4739, NY). Tab (Cowan 2967, CAS; Zamudio 1469, IEB). Ver (Wendt et al. 2634, 3999, CHAPA, NY).

Unverified, Doubtful, or Mistaken Reports. Tab (reported by Magan˜a, 1992, based on a misidentification of M. reptans).

Tryon and Tryon (1982: 717) commented that M. reptans and several other scaly-bladed species were “poorly distinguished” from M. piloselloides. More recently, R. Tryon and Stolze (1993) separated these two and distinguished them by the more tightly appressed rhizome scales of M. piloselloides (vs. loosely appressed and with tips widely spreading in M. reptans) and the whitish

MICROGRAMMA TECTA

(vs. orangish to castaneous in M. reptans) scales on the adaxial laminar surfaces. However, they also included M. acatallela Alston in synonymy under M. piloselloides, and we believe the former is adequately distinct from both M. piloselloides and M. reptans. Moreover, they accorded a much broader range to M. piloselloides than we give it.

399

and M. tecta, which see. From the former it differs by its lanceolate to ovate sterile blades and slightly thicker rhizomes with more appressed scales; from the latter it differs chiefly by the narrower fertile blades with mature sori projecting beyond the blade margins.

6. MICROGRAMMA TECTA VAR. NANA 5. MICROGRAMMA REPTANS Microgramma reptans (Cav.) A. R. Sm., Proc. Calif. Acad. Sci., ser. 4, 40: 230. 1975. Figs. 192C, D. Acrostichum reptans Cav., Anales Hist. Nat. 1: 104. 1799. Type. Ecuador. Guayaquil, Ne´e s.n. (MA; seen and commented upon by Christensen, Dansk Bot. Ark. 9(3): 9. 1937. Polypodium ciliatum Willd., Sp. Pl., ed. 4, 5(1): 144. 1810. Microgramma ciliata (Willd.) Alston, Bull. Jard. Bot. E´tat 27: 56. 1957. Type. Brazil. Para´: Hoffmannsegg s.n. (B).

Rhizomes ca. 0.5–1 mm diam.; rhizome scales ferrugineous to stramineous, mostly 2–4 mm long, linear-lanceolate with longattenuate apices, entire, Ⳳ appressed but with spreading tips; fronds strongly dimorphic; stipes 0–4 mm long in sterile fronds, 0–10 mm in fertile fronds; sterile blades nearly exstipitate, elliptic to ovate, to ca. 6 ⫻ 1 cm, acute to cuneate at bases, acute to rounded at apices; fertile blades linear, ca. 5(–9) cm ⫻ 2–3(–4.5) mm; indument of scattered, ferrugineous, peltate-based scales on both sides of blades, scales with pale, fringed margins and a tail often longer than body of a scale; sori when mature usually projecting beyond margins of fertile blades, with numerous, tan acicular scales among the sporangia.

Distribution. Epiphytic in lowland and montane rain forests; 100–1400 m. Mexico; Guat, Bel, Hond, Nic, CR, Pan; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol. Specimens Examined. Chis (Breedlove 34129, DS, MEXU, MO; Martı´nez 15541, MEXU; Rovirosa 859, PH). Tab (Magan˜a et al. 1024, ENCB, MO; Mares et al. 53, MEXU).

This species was recorded for Cuba by Smith (1981) and Moran (in Davidse et al., 1995), but we are no longer able to verify that distribution. Microgramma reptans is most closely related to M. piloselloides

Microgramma tecta (Kaulf.) Alston var. nana (Liebm.) Mickel & Beitel, Mem. New York Bot. Gard. 46: 251. 1988. Figs. 192H, J, K. Acrostichum nanum Liebm., Mexic. Bregn. 171 (reprint 19). 1849. Type. Mexico. Oaxaca: [Dist. Miahuatla´n] between San Miguel Coatla´n and La Galera, Liebmann s.n. [Pl. Me´x. 2655, Fl. Me´x. 36] (C!). Polypodium blandulum Christ, Bull. Herb. Boissier, se´r. 2, 7: 259. 1907. Type. Costa Rica. Espavey, Rı´o Grande, 1906, Werckle´ s.n. (P!).

Rhizomes slender, 0.3–0.5 mm diam.; rhizome scales 2–3 mm long, concolorous, light brown, narrow, peltate, somewhat spreading, entire; fronds dimorphic, thin; stipes 0–12 mm long in sterile fronds, 1–10 mm in fertile fronds; sterile blades ovate to orbicular or elliptic, 8–20 ⫻ 6–13 mm, apices obtuse, bases cordate to truncate; fertile blades lanceolate, 10–40 ⫻ 2.5–5 mm, apices obtuse, bases cuneate; indument of numerous narrow, brownish scales, 1–1.5 mm long on both blade surfaces, scales with orbicular fimbriate bases and long hair-like tips; sori round, with narrow scales (like laminar scales) among the sporangia.

Distribution. Epiphytic, mixed forest of pines and hardwoods; 1100–2200 m. Mexico; CR; Col, Ven, Guy (Jansen-Jacobs 4962, UC), Sur, Fr Gui, Ec, Peru, Bol. Also reported from Honduras by Moran (in Davidse et al., 1995). Specimens Examined. Chis (Gilly & Herna´ndez X. 162, US p.p.). Oax (Anderson 13205, CAS, CHAPA, IEB, MO, NY; Bashor 1971, NY; Campos V. 1011, MEXU; Conzatti & Reko 3033, 3298, MEXU, US).

The small blade size with distinctly dimorphic fronds, lanceolate blade scales, and similar soral scales distinguish this species from congeners in Mexico. Variety tecta, from southeastern Brazil (based on Polypodium tectum Kaulf., Enum. Filic. 87. 1824. Type. Brazil. Ilha de Santa Catarina, Chamisso s.n., LE), is distinguished primarily by the broader scales of the fertile blades

400

MICROPOLYPODIUM

(mostly 0.3–0.6 mm wide at bases vs. 0.1–0.25 mm wide in var. nana) and the longer sterile blades (length 2–4 times width, acute at tips vs. length equalling to 1.5 times width, obtuse at tips in var. nana).

7 9. M IC R O PO L YPO D I U M Micropolypodium Hayata, Bot. Mag. Tokyo 42: 341. 1928. Type: Micropolypodium pseudotrichomanoides (Hayata) Hayata [⬅ Polypodium pseudotrichomanoides Hayata] ⫽ M. okuboi (Yatabe) Hayata. Usually epiphytic, rarely epipetric; rhizomes suberect or erect, radially symmetrical; rhizome scales golden, orangish, or yellowto orange-brown, often lustrous, non-clathrate or weakly clathrate, entire or irregularly toothed, lacking setulae or sparsely glandular to setulose on the margins, setulae concolorous or often darker than scale body; fronds monomorphic or nearly so, determinate; stipes not articulate, 0.2–0.5 mm diam.; blades linear, commonly 3–15(–25) ⫻ 0.4–8(–1.2) cm, shallowly to deeply pinnatifid, pinnatisect, or barely pinnate, the segments (pinnae) oblong to deltate, decurrent, sometimes with an acroscopic hump at the base (i.e., segments gibbous); hydathodes present, round to clavate, usually conspicuous adaxially; veins simple or with only a single acroscopic fertile vein branch, hidden and not darkened abaxially; indument of reddish to dark red-brown stiff setae often 1–3 mm long on stipes, rachises, costae, and sometimes laminae, also of simple or often branched, septate, hyaline,

sometimes glandular hairs; sori round, superficial, one per pinna, usually adjacent to rachises at bases of costae or on the acroscopic veinlets, exindusiate; soral paraphyses absent; sporangia glabrous; spores tetrahedral-globose; x⫽37. Micropolypodium comprises about 30 species from the Antilles and southern Mexico to Bolivia and southern Brazil; a few species occur in eastern Asia and Malesia (Smith, 1992). The genus differs from all other neotropical genera of Grammitidaceae by the combination of radially symmetric, erect rhizomes, often golden brown, tumid, non-clathrate rhizome scales, linear blades usually less than 10 mm wide, prominent vein tips adaxially, and unisoriate segments (pinnae) each with a single unbranched or acroscopically 1-forked vein. The type of the genus is from eastern Asia, but by far the greatest diversity in the genus is in the neotropics, particularly in the Antilles and Mesoamerica. Preliminary molecular data on three neotropical species of Micropolypodium indicate that it is allied to very dissected (2pinnate) species of Terpsichore (T. achilleifolia (Kaulf.) A. R. Sm. and T. longisetosa (Hook.) A. R. Sm.), and that the Asian (and type) species M. okuboi may not be congeneric (Ranker et al., 2004). If upheld, these findings have implications for the classification of Micropolypodium, as well as for Terpsichore. References Maxon, W. R. 1916. Studies of tropical American ferns–No. 6. Contr. U.S. Natl. Herb. 17: 541–608; Smith, A. R. 1992. A review of the fern genus Micropolypodium (Grammitidaceae). Novon 2: 419–425.

Key to the Mexican Species of Micropolypodium 1. Rhizome scales brown-tinged toward tips, with castaneous setulae along margins; blades, at least proximally, incised to rachises, proximal segments not strongly decurrent. ............................................................................................................................................... 2. M. taenifolium. 1. Rhizome scales golden or orangish tan throughout, lacking setulae; blades deeply pinnatifid, segments connected by a narrow wing; proximal segments usually strongly decurrent. 2. Segments not or only slightly gibbous, ascending 60–75⬚ with respect to rachises on which they are borne; stipes usually distinct. .............................................................................................................................................................................................. 1. M. basiattenuatum. 2. Segments conspicuously gibbous (with a hump or lobe on acroscopic margin of each segment), spreading or ascending to ca. 70⬚; stipes essentially absent. ............................................................................................................................................................ 3. M. trichomanoides.

1. MICROPOLYPODIUM BASIATTENUATUM Micropolypodium basiattenuatum (Jenman) A. R. Sm., Novon 2: 422. 1992. Figs. 193D, E. Polypodium basiattenuatum Jenman, Bull. Bot. Dept. Jamaica, n.s. 4: 114. 1897. Xiphopteris basiattenuata (Jenman) Copel., Amer. Fern J. 42: 104. 1952. Grammitis basiattenuata (Jenman) Proctor, Bull. Inst. Jamaica, Sci. Ser. 5: 32. 1953. Type. Jamaica. Blue Mt. Peak, Jenman s.n. (NY!). Grammitis basiattenuata (Jenman) Proctor var. valens Mickel & Beitel, Mem. New York Bot. Gard. 46: 197. 1988. Type. Mexico. Oaxaca: Distrito Ixtla´n, 2 km E of Natividad, on lower part of trail from San Pedro Nolasco to the Llano Verde, Mickel 3824 (NY!).

Rhizome scales golden to orangish tan, 1–2 mm long, with 1–2 irregular teeth along margins, not setulose, near tips with 1(–2) glandular hairs ca. 0.05 mm projecting away from plane of scale; fronds 6–20 cm long, usually straight; stipes brownish, 1–1.5(–3) cm long, narrowly winged, ca. 1⁄5 the frond length or less, with red-brown setae 1–2 mm long; blades 7–10(–17) cm ⫻ 7–8 mm, deeply pinnatifid, gradually tapering proximally to a narrow wing; segments ascending ca. 60–75⬚ to rachis, 15–25(–45) pairs, short-oblong, 3.5–4.5 ⫻ 2–2.5 mm, more decurrent, distant, and ascending proximally, not gibbous acroscopically, obtuse apically; indument on rachises, costae, veins, and laminar tissue (both sides of blades) of red-brown setae 1.5–2.5 mm long and also, abaxially, of twisted, branched hairs 0.3 mm long; veins usually 1-forked, at least in fertile segments, or sometimes simple in sterile segments; sori ca. 1.5–2 mm diam.

MICROPOLYPODIUM TRICHOMANOIDES

401

hairs; veins usually 1-forked, at least in fertile segments, or sometimes simple in sterile segments; sori ca. 1–1.5 mm diam.; 2n⫽74 (Trin).

Distribution. Epiphytic in montane cloud forests; 2050–2500 m. Mexico; Guat, Hond (Moran 5580, MO, UC), Salv (Seiler 1133, NY); Jam; w Ven. Specimens Examined. Chis (Breedlove 25421, DS p.p.; Breedlove 30160, DS, LL, MEXU, MO, NY; Pe´rez F. 252, UAMIZ). Oax (Mickel 3824, NY).

Distribution. Epiphytic in wet montane forests; 900–1600 m. Mexico; Guat, Bel (Holst et al. 5248, UC), Hond, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec.

From M. taenifolium, M. basiattenuatum differs by the golden or orangish rhizome scales lacking marginal setulae, the less incised blades, and the more strongly decurrent segments. Micropolypodium basiattenuatum differs from M. trichomanoides by the non-gibbous segments and distinct stipes. Jamaican specimens, including the type, have shorter stipes (5–10 mm) and shorter pinnae (2.5–3 ⫻ 1.5–2 mm at mid-segment), but otherwise do not differ significantly from Mexican and Central American specimens.

Selected Specimens Examined. Chis (Breedlove 30809, MEXU, TEX, 31531, 32119, 38855, 38906, DS, 42074, DS, MEXU, MO). Oax (Hallberg 1728, NY; Wendt et al. 6751, TEX). Ver (Nee 24971, XAL; Va´zquez et al. 4108, XAL).

2. MICROPOLYPODIUM TAENIFOLIUM Micropolypodium taenifolium (Jenman) A. R. Sm., Novon 2: 423. 1992. Figs. 193F, G. Polypodium taenifolium Jenman, Bull. Bot. Dept. Jamaica, n.s. 4: 114. 1897. Xiphopteris taenifolia (Jenman) Copel., Amer. Fern J. 42: 109. 1952. Grammitis taenifolia (Jenman) Proctor, Bull. Inst. Jamaica, Sci. Ser. 5: 35. 1953. Type. Jamaica. St. Andrew: Near Mt. Moses, Syme s.n. (J.P. 197, p.p.) (isotype IJ). Polypodium blepharodes Maxon, Contr. U.S. Natl. Herb. 17: 407. 1914. Xiphopteris blepharodes (Maxon) Copel., Amer. Fern J. 42: 109. 1952. Grammitis blepharodes (Maxon) F. Seym., Phytologia 31: 173. 1975. Type. Costa Rica. La Palma, Maxon 406 (US!; isotype NY!).

Rhizome scales orangish brown, often castaneous at tips, 1.5– 2 mm long, with setulose margins, setulae darker than scale body, ascending, 0.1–0.3 mm long; fronds mostly 7–20(–25) cm long, usually straight; stipes gray-brown, 0.5–2.5 cm long, ca. 1⁄10 the frond length, with abundant, dark red-brown or maroon setae 1–2.5 mm long; blades 7–10 ⫻ 0.5–0.7 cm, pinnatisect, gradually tapering proximally; segments (pinnae) spreading nearly perpendicularly (80–90⬚) to rachis, 35–60⫹ pairs, 2–3 ⫻ 1–1.5 mm, bases broadly adnate, slightly decurrent, gibbous (especially when fertile) or not acroscopically, apices obtuse; indument on rachises, costae, veins, and laminar tissue (both sides of blades) of dark red-brown setae 1–2.5 mm long, lacking septate hyaline

Ibarra (1983) cited a specimen as Grammitis aff. blepharodes from Michoaca´n, but we have not seen the specimen in question (see Dı´az-Barriga & Palacios-Rios, 1992: 46). In Mexico and Mesoamerica, this species has often gone under the name Grammitis blepharodes, but we are unable to distinguish adequately that species from the Antillean and northern South American G. taenifolia, the older name. Micropolypodium taenifolium differs from M. basiattenuatum and M. trichomanoides in having setulose rhizome scales, pinnatisect or even pinnate blades with shorter, narrower segments, more decidedly setose stipes, and darker rachises. In general, it can be also be distinguished from its congeners in Mexico by the blades being darker green adaxially and distinctly lighter green abaxially. Pichi Sermolli (Webbia 37: 118–125. 1983), Proctor (1985), and Lellinger (1989) applied the name Grammitis flabelliformis (Poir.) C. V. Morton to this species, but Bishop (Taxon 38: 91–95. 1989) argued convincingly that one must accept the first lectotypification of the name, by Morton (Contr. U.S., Natl. Herb. 38: 29–83. 1967); if that is done, the name is applied to the species here treated as Melpomene flabelliformis (which see).

3. MICROPOLYPODIUM TRICHOMANOIDES Micropolypodium trichomanoides (Sw.) A. R. Sm., Novon 2: 423. 1992. Figs. 193A–C. Polypodium trichomanoides Sw., Prodr. 131. 1788. Grammitis trichomanoides (Sw.) Ching, Bull. Fan Mem. Inst. Biol. 10: 16. 1940. Xiphopteris trichomanoides (Sw.) Copel., Gen. Fil. 215. 1947. Type. Jamaica. Swartz s.n. (S?; isotypes

402

MILDELLA

B-Herb. Willd. 19670—icrofiche UC!, FI–photo US, UPS-Thunb. 24583, microfiche UC!).

Rhizome scales golden to orangish tan, 2–4 mm long, not setulose, near tips with 1 or 2 glandular hairs ca. 0.05 mm projecting away from plane of scales; fronds mostly 6–12 cm long, often curved; stipes essentially absent; blades to ca. 12 ⫻ 0.5–0.7 cm, deeply pinnatifid, gradually to subabruptly tapering proximally to a narrow wing; segments spreading to ascending ca. 70⬚ to rachis, connected along rachis by a narrow wing, to ca. 45 pairs, 2–3 ⫻ 1–1.5 mm, bases broadly adnate, fertile ones often conspicuously gibbous at the acroscopic base of each segment, rounded to subacute at apices; indument on rachises, costae, veins, and laminar tissue (both sides of blades) of scattered reddish brown to dark maroon setae to ca. 2 mm long, also, abaxially, of a few twisted, branched hairs 0.1–0.3 mm; veins usually 1-forked, at least in fertile segments, or sometimes simple in sterile segments; sori 1–1.5 mm diam.; 2n⫽72 (Braz), probably erroneous.

Distribution. Epiphytic in montane rain forests, pine-oakLiquidambar forests; 1200–1800 m. Mexico; Guat, Hond; Cuba, Jam; Ven, Fr Gui. Specimens Examined. Chis (Breedlove 29554B, DS, 32225A, DS, ENCB, NY, 38886, DS, 53455, CAS). Oax (Ishiki 1259, MEXU p.p.; Rzedowski 33769B, ENCB). Ver (Lira 28, ENCB, MEXU, UAMIZ; Ramı´rez & Palma 1049, XAL).

In Mexico, this species is most similar to M. basiattenuatum but has the fertile segments usually conspicuously gibbous and more spreading. Doubtful Species Polypodium gibbosum Fe´e, Me´m. Foug. 6: 8, t. 2, f. 2. 1854. Type. Mexico. Oaxaca: 2400–2600 m, specimen received from Galeotti under the name Polypodium delicatulum. This name has been of uncertain application since the taxon was described, no doubt because no one has been able to locate the type (not mentioned by Windisch, 1982, Amer. Fern J. 72: 56–60, in a paper on Fe´e’s types in RB). Maxon (1916, p. 556) discussed its identity and concluded that it belonged to the group of Polypodium trichomanoides (⬅ Micropolypodium trichomanoides), an opinion with which we concur. If the type really is from Mexico,

it is most likely Micropolypodium trichomanoides or M. taenifolium; if it is the same as the latter, Polypodium gibbosum has priority.

80. MILDELLA Mildella Trevis., Rendiconti Reale Ist. Lombardo Sci., Cl. Sci. Mat., ser. 2, 9: 810. 1876. Type: Mildella intramarginalis (Kaulf. ex Link) Trevis. [⬅ Pteris intramarginalis Kaulf. ex Link] Terrestrial or epipetric; rhizomes short- to long-creeping, short-ascending, with bicolorous scales; fronds small, clumped, monomorphic; stipes stramineous, reddish brown or black, adaxially grooved, lustrous, glabrous except for clavate hairs adaxially; blades pinnate-pinnatifid to bipinnate, proximal pinnae basiscopically exaggerated, laminae subcoriaceous; veins free, simple or forked; sori essentially marginal, indusia just back from margin (inframarginal), recurved, entire, erose or ciliate; spores roughened; x⫽30. Mildella, as usually construed, comprises a genus of nine species, six of southern and eastern Asia, one in Haiti, and two widespread in Mexico and Central America at middle elevations, in mesic habitats. The false margins projecting beyond the false indusia distinguish Mildella. The genus is doubtfully distinct from Cheilanthes. Molecular data show Mildella to be nested within Cheilanthes (Gastony & Rollo, 1995) and also that Mildella sensu Hall and Lellinger (1967) is polyphyletic (Cranfill, unpubl.) with the Asian species belonging to Aleuritopteris. New World species are probably related to the Cheilanthes angustifolia complex, having distinct hydathodes, prominulous veins, and grooved rachises and stipes, but the stipes and rachises in Mildella are hirsute adaxially (vs. glabrous in C. angustifolia and allies). References Hall, C. C. & D. B. Lellinger. 1967. A revision of the fern genus Mildella. Amer. Fern J. 57: 113-134; Nesom, G. L. 1993. Mildella intramarginalis var. serratifolia (Adiantaceae) elevated to species rank. Phytologia 75: 383-384. 1993.

Key to the Mexican Species of Mildella 1. Blade margins entire to slightly serrate; rhizome scales castaneous to atropurpureous, pale margins nearly as wide as the central streak; indusia entire to rarely erosecrenate. .................................................................... 2. M. intramarginalis. 1. Blade margins strongly serrate; rhizome scales blackcentered, with very narrow pale margins; indusia ciliate, hairs 0.3–0.5 mm long. ........................................................ 1. M. fallax.

1. MILDELLA FALLAX Mildella fallax (M. Martens & Galeotti) Nesom, Phytologia 75: 384. 1993. Figs. 194A–D.

NEPHROLEPIS Pteris fallax M. Martens & Galeotti, Me´m. Foug. Mexique 53, pl. 14, f. 2. 1842. Pellaea intramarginalis (Kaulf. ex Link) J. Sm. var. ß serratifolia Hook. & Baker, Syn. Fil. 149. 1867. Mildella intramarginalis (Kaulf. ex Link) Trevis. var. serratifolia (Hook. & Baker) C. C. Hall & Lellinger, Amer. Fern J. 57: 124. 1967. Type. Mexico. Oaxaca: “Pre`s de Tanetze, Talea et Llano-Verde,” Galeotti 6467 (BR!, photo BM; isotype BR!).

Similar to M. intramarginalis, differing from that in having strongly serrate blade margins, very narrowly margined rhizome scales, and ciliate indusia, hairs 0.3–0.5 mm long; n⫽2n⫽90 (Oax).

Distribution. Rocky slopes in moist pine-oak woods; (950–) 1350–3400 m. Mexico; Guat. Selected Specimens Examined. Chih (Fishbein 1823, ARIZ). Chis (Ghiesbreght 239, NY). DF (Lyonnet 234, NY). Gro (Hinton 9490, NY). Gto (Harris 5894, NY). Hgo (Chase 7371, NY). Jal (McVaugh 16256, MICH). Me´x (Quiroz 5111, MEXU). Mich (Dı´az B. 2038, ENCB). Mor (Orcutt 4232, US). NL (Pringle 1986, NY). Oax (Mickel 5034, NY). Pue (Nicolas 5095, NY). Qro (Argu¨elles 958, MEXU). Sin (Breedlove 44988, CAS). SLP (Limon 56, MEXU). Tam (Stanford 2071, NY). Tlax (Sohns 646, MEXU). Ver (Nee 33083, NY). Unverified, Doubtful, or Mistaken Reports. Ags (Garcı´a 3161, HUAA, and one other collection, cited by Siqueiros-Delgado & Gonza´lez-Adame, 2004 [as Mildella intramarginalis var. serratifolia], but not verified).

This species has strongly ciliate indusia and serrate blade margins, and generally occurs at higher elevations than does M. intramarginalis.

2. MILDELLA INTRAMARGINALIS Mildella intramarginalis (Kaulf. ex Link) Trevis., Rendiconti Reale Ist. Lombardo Sci., Cl. Sci. Mat., ser. 2, 9: 810. 1876. Figs. 194E, F. Pteris intramarginalis Kaulf. ex Link, Hort. Berol., ed. 2, 2: 34. 1833. Cheilanthes intramarginalis (Kaulf. ex Link) Hook., Sp. Fil. 2: 112. 1852. Pellaea intramarginalis (Kaulf. ex Link) J. Sm., Cat. Kew Ferns 4. 1856. Lectotype (chosen by C. C. Hall & Lellinger, 1967: 119). Cultivated specimen labelled “Pteris intramarginalis Klf.. . . .Hort. berol.” (B; isolectotype LE!).

Rhizomes short, horizontal, to ascending; rhizome scales castaneous to atropurpureous, thick, with thin brown margins nearly as broad as the central streak, 2–4 ⫻ 0.3–0.8 mm; fronds

403

to 39 cm tall, clumped; stipes 1⁄2–2⁄3 the frond length, castaneous, glabrous, puberulous in grooves with short, clavate hairs; blades pinnate-pinnatifid to bipinnate, segments broadly adnate, linear, 3–5 pinna pairs divided, distal pinnae undivided; costae and larger costules dark; veins prominulous, diverging from costules at narrow angles (similar to C. angustifolia), with short hairs 0.1 mm; adaxial laminar surfaces glabrous except for puberulous grooves of rachises and costae, and adjoining laminar tissue; abaxial laminar surfaces glabrous; blade margins entire to slightly serrate; indusia distinct, 0.8 mm wide, well modified, thin, entire or erose-crenate, cells whitened; spores gray-brown to black.

Distribution. Rocky slopes in moist pine-oak forests; 800– 2200 m. Mexico; Guat, Hond, Salv, Nic, CR, Pan; Galapagos. Selected Specimens Examined. Chis (Breedlove 22684, NY). Gro (Rzedowski & McVaugh 49, NY). Hgo (Rzedowski 27654, NY). Mich (Hinton 15544, NY). Mor (Hinton 17459, NY). Oax (Mickel 5068, NY). Pue (Ventura A. 1394, NY). Qro (Ferna´ndez N. 3778, NY). Sin (Gonza´lez Ortega 5305, MEXU). SLP (Rzedowski 10052, ENCB). Tam (Breedlove 63695, CAS). Ver (Seaton 40, NY).

Mildella intramarginalis is distinct by its entire to slightly serrate blade margins, entire to erose-crenate indusial margins, and general occurrence at lower elevations.

81. NEPHROLEPIS Nephrolepis Schott, Gen. Fil., no. 3. 1834. Lectotype: Uncertain, regarded by J. Smith (Hist. Fil. 226. 1875) and R. Tryon (Contr. Gray Herb. 194: 225. 1964) as Nephrolepis exaltata (L.) Schott, but by Christensen (Index Filic. XXVII. 1906) as N. biserrata (Sw.) Schott. Terrestrial, on rocks, or epiphytic; rhizomes decumbent or commonly erect, weakly developed, dictyostelic, perennial to probably annual, scaly at the apices, usually stoloniferous, with stolons mostly aboveground and wiry, sometimes producing young plants along their length, a few species bearing underground, perennating tubers; fronds clumped in a tight cluster, erect to arching to pendent in some species, monomorphic, polystichous, non-articulate, medium-sized to large; stipes short, scaly to glabrescent, stramineous to brown; blades 1-pinnate,

404

NEPHROLEPIS

slowly determinate to seemingly indeterminate with a loose bud formed of undeveloped pinnae at each blade apex, glabrous, hairy, or with lanceolate scales; pinnae numerous, thin to subcoriaceous, entire or shallowly lobed to serrulate or crenulate, sometimes auriculate at the superior base, inequilateral with greater development on acroscopic side, articulate to rachises, often with lime-dotted hydathodes on vein tips adaxially; veins free, 1–4-forked; sori terminal on the veins, sometimes confined to the distal pinnae, medial to often supramedial or submarginal, round to slightly elongate, uniseriate along both sides of costae, lacking paraphyses; indusia persistent, round to semicircular or reniform (lunate), entire; spores ellipsoid, bilateral, perispores rugose to tuberculate; x⫽41.

teroid genera, but the exact relationships are still uncertain (Hasebe et al., 1995). Numerous cultivars exist within several of the native and naturalized species. These are discussed under the appropriate species below. One additional cultivar, not known to have escaped in Mexico, is often also grown in many cities: Nephrolepis falcata (Cav.) C. Chr. cv. Furcans, represented by specimens from Veracruz (Nevling & Go´mez-Pompa 297, MEXU) and Guerrero (Calzada & Salinas 17700, UAMIZ); we have seen it growing in the city of Xalapa, and also in Veracruz, and Magan˜a (1992) cited it from Tabasco. This species is believed to be native to Malesia and Polynesia. The cultivar can be readily recognized by the rather large, usually 1-forked pinnae, and hence has the common name “Fishtail fern.”

Nephrolepis is a genus of 20–25 species of tropical regions of the world, 7 native in America, one species naturalized from Asia. The genus is very distinctive, distinguished by the wiry stolons, linear, 1-pinnate fronds, and articulate pinnae. It is sometimes placed in its own family, Nephrolepidaceae; by others, it is placed in Davalliaceae or Dryopteridaceae. In molecular analyses, Nephrolepis is imbedded within a large group of dryop-

References Morton, C. V. 1958. Observations on cultivated ferns. V. The species forms of Nephrolepis. Amer. Fern J. 48: 18–27; Nauman, C. E. 1981. The genus Nephrolepis in Florida. Amer. Fern J. 71: 65–70; Nauman, C. E. 1985. A systematic revision of the neotropical species of Nephrolepis Schott. Ph.D. dissertation. Univ. Tennessee, Knoxville; Pichi Sermolli, R. E. G. 1968. Taxonomical notes on Nephrolepis cordifolia (Linnaeus) C. Presl and related species. Estratto Dagli Ann. Mus. Civico Storia Nat. Genova 77: 270–277.

Key to the Mexican Species of Nephrolepis 1. Bases of mature stipes with dark brown or blackish, appressed scales with narrow pale margins. ..................................................... 4. N. hirsutula. 1. Bases of mature stipes often with a few lax, reddish to light brown scales, or seemingly without scales. 2. Indusia orbicular with sinuses almost closed to orbicular-reniform with sinuses narrow to U-shaped, sporangia spreading in all directions; leaf tissue with sparse to dense fibrillose scales and/or catenate hairs abaxially. 3. Costae adaxially with dense, catenate hairs 0.2–0.6 mm long. ..................................................................................................... 1. N. biserrata. 3. Costae adaxially with sparse, fibrillose scales, or sometimes glabrous. 4. Blades 5–8.5 cm wide; pinnae with basiscopic bases rounded to auriculate; indusia suborbicular to reniform, pale brown with dark brown center, 1.1–1.3 mm wide; rhizome scales obscurely bicolorous, mostly pale brown, margins entire; scales on stipes, rachises, and pinnae pale brown. ..................................................................................................................... 3. N. exaltata. 4. Blades (7–)8.5–11 cm wide; pinnae with basiscopic side cuneate; indusia round-reniform, dark reddish brown, 0.8–1 mm wide; rhizome scales distinctly bicolorous, with lustrous, reddish brown central stripe and pale brown, long-ciliate margins; scales on stipes, rachises, and pinnae castaneous. ..................................................................................................... 7. N. rivularis. 2. Indusia reniform to lunate, sinuses broad or absent, mature sporangia projecting on open side, mostly facing pinna apices; leaf tissue glabrous abaxially. 5. Pinna bases cuneate basiscopically, Ⳳ perpendicular to rachis; rhizome scales weakly bicolorous; rhizomes erect, lacking tubers. .................................................................................................................................................................................................... 5. N. pectinata. 5. Pinna bases rounded to cordate basiscopically; rhizome scales concolorous; rhizomes well or poorly developed, tubers present or absent. 6. Stipes stramineous; plants with poorly developed rhizomes, often bearing scaly tubers; pinnae acuminate at apices, tapering gradually from bases; indusia 1.5–2 mm in greatest dimension. ............................................................................. 8. N. undulata. 6. Stipes light brown to brown; plants with well developed rhizomes, often forming erect caudices, bearing tubers or not; pinnae rounded to acute at apices, usually parallel-sided or slightly tapering; indusia 1–1.5 mm in greatest dimension. 7. Rhizome scales orange to tan, entire; stipes and rachises persistently fibrillose-scaly, dull light brown; rachis scales bicolorous (pale with dark point of attachment); tubers present or absent. .................................................................. 2. N. cordifolia. 7. Rhizome scales castaneous, lustrous, with long-ciliate margins; stipes brown, lustrous, with a few scales at bases; rachises glabrous except at pinna bases; rachis scales concolorous; tubers lacking. ........................................................ 6. N. pendula.

1. NEPHROLEPIS BISERRATA Nephrolepis biserrata (Sw.) Schott, Gen. Fil., t. 3. 1834. Figs. 196H, J–M. Aspidium biserratum Sw., J. Bot. (Schrader) 1800(2): 32. 1801. Type. Mauritius. Gro¨ndal s.n. (S).

Nephrolepis mollis Rosenst., Repert. Spec. Nov. Regni Veg. 22: 13. 1925. Type. Costa Rica. Brade 141 [Rosenst. exs. 289] (S?; isotypes NY!, UC!).

Rhizomes suberect to erect, stout, lacking tubers; rhizome scales concolorous, pale brown to orange, lanceolate, 5–7 mm long; stipes light brown, 20–30 cm long, filiform-scaly to glabrescent; blades firm-herbaceous, linear-elliptic, 75–100(–200) ⫻ (11–)16–24 cm; rachises stramineous to tan, with dense, fibrillose

NEPHROLEPIS CORDIFOLIA

scales, 1–2 mm long, adaxial surfaces with catenate hairs; pinnae short-stalked, Ⳳ equally cuneate to equally auriculate at bases, (5–)8.5–20 ⫻ 0.9–2 cm, apices of longest pinnae acuminate, margins finely serrulate (sterile blades) to crenulate or bicrenulate (fertile blades); indument abaxially, especially on costae, of fine, often rather dense catenate hairs 0.2–0.6 mm long and scattered to dense, tan, fibrillose scales, especially on costae, adaxial surfaces with numerous catenate hairs 0.1–0.3 mm long on costae; hydathodes often lime-dotted on adaxial blade surfaces; indusia brown, round-reniform, 0.8–1 mm diam., with a few hairs 0.1– 0.3 mm long, each indusium with a narrow sinus (mature sporangia projecting on all sides); 2n⫽82 (Jam, PR, Trin, Africa, India).

405

in this presumed complex must await more critical examination of specimens throughout the range.

2. NEPHROLEPIS CORDIFOLIA Nephrolepis cordifolia (L.) C. Presl, Suppl. Tent. Pterid. 79. 1836. Figs. 196D–G. Polypodium cordifolium L., Sp. Pl. 2: 1089. 1753. Type. Petiver, Pter. Amer., t.1, f. 11, 1712, which is a transposed copy of Plumier, Traite´ Foug. Ame´r., pl. 71, 1705, illustrating a plant from Hispaniola.

Rhizomes erect, stoloniferous, often with tubers; rhizome scales spreading, concolorous, linear, orange to tan, 1–3 mm long, margins entire; stipes 9–18 cm long, ca. 1⁄6 the frond length, light brown, grooved, often persistently fibrillose-scaly; blades firm, herbaceous, linear-elliptic, 30–80 ⫻ 5–7 cm; rachises light brown, fibrillose-scaly, scales bicolorous (pale with dark point of attachment); pinnae narrowly deltate, apices acute, bases unequal, lobed with basiscopic lobe rounded to cordate, not overlapping rachises, acroscopic lobe larger, auriculate, slightly overlapping the rachis, margins crenate; veins indistinct, glabrous; hydathodes often lime-dotted on adaxial blade surfaces; indusia tan, reniform to lunate, 1–1.3 mm in longest dimension, mostly opening toward pinna apices, attached along broad, shallow sinuses, opening toward pinna apices; 2n⫽82 (India).

Distribution. Terrestrial, epipetric or epiphytic on matorral, palm plantations (Tabasco), evergreen rain forests; 100–1000 m. USA (Fla); Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Bah, Gr & L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol; Africa, Asia, Polynesia. Selected Specimens Examined. Camp (Tun G. et al. 67, MEXU). Chis (Martı´nez S. 9505, MEXU, UAMIZ, XAL, 10496, MEXU, MO, UAMIZ). Nay (Gonza´lez Ortega 73, US). Oax (Herna´ndez G. & Gonza´lez 1692, CAS, MO, NY, UAMIZ). Pue (Ventura 576, ENCB, NY). QR (Torres 47, UAMIZ). Tab (Valdivia 2032, MO, XAL). Ver (Orcutt 6625, DS, NY, UC; Ventura A. 14976, UAMIZ, XAL). Unverified, Doubtful, or Mistaken Reports. Jal (Martı´nez s.n., May 1989, IBUG, cited by Mickel, 1992, but not verified). Yuc (Tun G. 262, UADY, cited by Palacios-Rios, 2002b, but not verified).

The rotund-reniform indusia with occasional hairs, pinnae with hairs on the costae adaxially and laminar tissue abaxially, and concolorous, spreading rhizome and stipe base scales distinguish this species. See key for comparison with N. hirsutula, seemingly a recent introduction. In Florida and Jamaica N. biserrata hybridizes with N. exaltata to produce an abortive-spored hybrid (N. ⫻averyi), described by Nauman (Amer. Fern J. 69: 69. 1979). The hybrid grows with its two parents, is larger than N. exaltata, and has sparse hairs adaxially. Paleotropical material under this name in herbaria, when not misidentified (which is often the case), seems to diverge significantly in a number of characters, including pubescence and pinna shape. Most likely, the name, as currently applied, encompasses more than one species, but a better delimitation of species

Distribution. Terrestrial on montane cloud forest; 750–1500 (–2200) m. USA (Fla); Mexico; Guat, Hond, Pan; Gr & L Ant; Peru, Braz; Old World tropics. Often attributed a much broader range in South America, by virtue of inclusion of N. pendula. Selected Specimens Examined. Chis (Breedlove 16064, 23670, DS). DF (Ferna´ndez N. 2, ENCB, a cultivar; Montes R. 7, UAMIZ). Me´x (Tejero-Dı´ez 2213, IZTA). Oax (Schultes 756, US). Pue (Knobloch 2165, US; Rzedowski 29996, ENCB, NY). Ver (Pringle 7995, MO; Provance 3339, UC; Purpus 253, BM; Valdivia Q. 1972, XAL; Ventura A. 884, ENCB, NY). Unverified, Doubtful, or Mistaken Reports. Tab (reported by Magan˜a, 1992, probably based on a misidentification).

This species is probably not native in Mexico (or Central America either), but is relatively recently introduced and natu-

406

NEPHROLEPIS

ralized in a few localities. The two Chiapas collections cited are from cultivated plants in gardens in and around San Cristo´bal Las Casas, and the two specimens from Distrito Federal are also from cultivated plants in or near Mexico City. Arse`ne 6690 (MEXU), cited by Dı´az-Barriga and Palacios-Rios (1992) from Michoaca´n, has not been found, but is apparently of garden origin (Palacios-Rios, pers. comm.); they cited this same collection, from the same herbarium, as N. pectinata. Rzedowski 29996 is from very disturbed oak forests. Most of the specimens in herbaria that have been identified as N. cordifolia have proven to be other species, particularly N. pectinata and N. pendula. See N. pendula, N. exaltata, and key for differences and discussion. Mexia 9274 has been determined as this species in some herbaria (e.g., NY), by Nauman, 1979, but apparently he later changed either his mind or species concepts, as other specimens of the same gathering (e.g., in UC, 1984) are determined by him as N. pectinata. All specimens of this number appear to be the same taxon, and we concur with this more recent determination. Nephrolepis cordifolia ‘Duffii’, a form with small orbicular pinnae, is sometimes cultivated in Mexico, but it is always vegetative and not likely escaped. However, a specimen has been seen from Tabasco, Acosta 42 (UAMIZ), “de abundancia regular en selva alta perennifolia en el cerro del madrigal.” This taxon was cited and illustrated from Tabasco by Magan˜a (1992) as Jamesonia alstonii!

3. NEPHROLEPIS EXALTATA Nephrolepis exaltata (L.) Schott, Gen. Fil., t. 3. 1834. Figs. 196S–W. Polypodium exaltatum L., Syst. Nat., ed. 10, 2: 1326. 1759. Type. Sloane, Nat. Hist. Jam., t. 31. 1707 [representing a plant collected by Harlow on Jamaica; see Jenman, J. Bot. 24: 34. 1886].

Rhizomes suberect, stoloniferous, lacking tubers; rhizome scales obscurely bicolorous, linear-lanceolate to filiform, ferruginous to pale brown, 6–7 ⫻ 0.5–0.8 mm, entire; stipes 10–20 cm long, pale brown, deciduously fibrillose-scaly; blades firmherbaceous, linear, 40–80 ⫻ 5–8.5 cm, narrowed at bases; rachises stramineous to pale brown, with dense to scattered fibrillose scales; pinnae subsessile to short-stalked, contiguous or subimbricate at bases, mostly subfalcate, 3–4.5 ⫻ 0.8–1.2 cm at the middle, usually rather unequal at the bases, with basiscopic lobe rounded to auriculate, acroscopic lobe auriculate, apices obtuse to acute, margins subentire to crenulate; indument abaxially of pale brown fibrillose scales 1–1.5 mm long, adaxially glabrous or laminae with few fibrillose scales; hydathodes rarely lime-dotted on adaxial blade surfaces; indusia pale brown with darker center, toward pinna bases orbicular to orbicular-reniform and each with a narrow sinus, to usually reniform or lunate and lacking a distinct sinus toward pinna apices, 1.1–1.3 mm in longest dimension; 2n⫽82 (Jam).

Distribution. Terrestrial, epiphytic or epipetric; ca. 1000 m. USA (Fla); Mexico; Pan; Gr Ant; Fr Gui, Braz. Native range uncertain, perhaps widely distributed in Old World tropics, but introduced there. Specimens Examined. Ver (Mu¨ller s.n., in 1853, NY, 441, K, NY; Purpus 3812, BM, MICH, MO, NY, UC, US). Unverified, Doubtful, or Mistaken Reports. Mich (reported by Mickel & Beitel, 1988, but not verified). Tab (reported by Magan˜a, 1992, but not verified).

Nephrolepis exaltata is usually considered pantropical, but many specimens labelled as this species are misidentified, and in particular it has been confused with N. hirsutula (also recently introduced), which Proctor (1981; as N. multiflora) reported to have supplanted N. exaltata in open or disturbed situations in Jamaica. The two cultivated specimens cited by Smith (1981: 152) as N. exaltata from Chiapas are now determined as N. cordifolia. Many cultivars of this species are known, but the most common ones seen in Mexico appear to belong to the Nephrolepis exaltata ‘Whitmanii’ complex, which has been found in Chiapas growing in the sitios of San Cristo´bal las Casas (Breedlove 16061, DS) and Chapa de Corzo (Breedlove 22862, DS). Also in Chiapas, and probably elsewhere, the cultivar ‘Bostoniensis’ has been found in sitios of Ocozocoautla (Breedlove 23496, DS).

4. NEPHROLEPIS HIRSUTULA Nephrolepis hirsutula (G. Forst.) C. Presl, Suppl. Tent. Pterid. 79. 1836. Figs. 195D–F, 196N–R. Polypodium hirsutulum G. Forst., Fl. Ins. Austr. 81. 1786. Type. Probably Polynesia, Society Islands, Tahiti, Forster s.n. (holotype or isotype BM, Morton photo 6876). For discussion and comments, see Morton, Contr. U.S. Natl. Herb. 38: 345. 1974. Davallia multiflora Roxb., Calcutta J. Nat. Hist. 4: 515, t. 31 [left]. 1844. Nephrolepis multiflora (Roxb.) Jarrett ex C. V. Morton, Contr. U.S. Natl. Herb. 38: 309. 1974. Lectotype (chosen by Morton, Contr. U.S. Natl. Herb. 38: 309. 1974). India. Roxburgh s.n. (BR!, frag. and photo US!).

Rhizomes erect, well developed, stolons stout, lacking tubers; rhizome scales bicolorous, 2–3 ⫻ 0.8–1 mm, dark reddish brown to blackish, lustrous, central portion sclerotic, margins whitish

NEPHROLEPIS PECTINATA

fimbriate-ciliate, appressed; stipes 5–25 cm long, ca. 1⁄5 the frond length, stramineous to light brown, with moderate to dense, lanceolate, appressed, bicolorous scales with dark sclerotic center and narrow, membranous, sometimes fibrillose margins; blades linear-elliptic, 30–70 ⫻ 6–15 cm; rachises with dense, appressed, white, fibrillose scales; pinnae narrowly deltate, obtuse to acute, bases slightly unequal, acroscopic auricle narrow, acute, basiscopic side rounded to short-auriculate; indument abaxially, especially on costae, of whitish fibrillose scales, adaxially the costae with dense hairs 0.1–0.3 mm long and scattered fibrillose scales; hydathodes sometimes lime-dotted on adaxial blade surfaces; indusia dark reddish brown or pale with dark center, roundreniform, 0.8–1 mm diam., each with a narrow sinus (mature sporangia projecting on all sides); 2n⫽82 (India).

407

the almost certain conspecificity of N. multiflora with the widespread N. hirsutula. R. Tryon and Stolze (1993: 52) distinguished these species in Peru by N. hirsutula having the costae adaxially “moderately to densely invested with dissected scales, rather than pubescent as in N. multiflora,” but we find dissected scales on all specimens so identified as N. multiflora, in both the Neotropics and Paleotropics. Thus, we are unable to distinguish these two species in herbarium holdings, and so apply the oldest name. A cultivar of this species, N. hirsutula ‘Superba’, is occasionally grown, e.g., in the sitios of San Cristo´bal las Casas (Chiapas): Breedlove 16031, DS.

5. NEPHROLEPIS PECTINATA Nephrolepis pectinata (Willd.) Schott, Gen. Fil., t. 3. 1834. Figs. 195G, H, J–L. Aspidium pectinatum Willd., Sp. Pl., ed. 4, 5(1): 223. 1810. Type. s. coll. s.n. (B-Willd. 19753!; photo GH, microfiche UC!).

Distribution. Terrestrial along roadsides and banks, lowland, and lower montane rain forests; 0–500(–1650) m. USA (Fla); Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Bah, Gr & L Ant; Col, Ven, Trin, Guy, Ec, Peru, Braz, Bol; tropical and subtropical Asia and the Pacific Basin (Malesia, Polynesia).

Rhizomes erect, well developed, stoloniferous, lacking tubers; rhizome scales weakly bicolorous, linear, reddish brown, 1–3 mm long, margins pale ciliate-dentate; stipes 4–17 cm long, 1⁄10–1⁄5 the frond length, tan, glabrous, with rare hair-like scales; blades firm-herbaceous, 18–60 ⫻ (2.5–)3–5(–6.5) cm, linear-elliptic; rachises glabrous except for matted, linear scales at pinna bases; pinnae usually oblong, parallel-sided to near apex, apices obtuse to acuminate, bases unequal, auriculate acroscopically, cuneate basiscopically, margins crenulate; indument on both blade surfaces generally absent or very sparse; hydathodes lime-dotted on adaxial blade surfaces; indusia tan, reniform or lunate, 1–1.5 mm wide, attached along broad, shallow sinuses mostly opening toward pinna apices; 2n⫽82 (Chis), 164 (Jam).

Selected Specimens Examined. Chis (Breedlove 26449, DS, NY, 34482, DS, ENCB, MO, NY; Martı´nez S. 10816, MO, UAMIZ). Oax (Martı´nez C. 848, CAS, NY; Mickel 1461, ENCB, NY, US). QR (Calzada et al. 6893, UC). Tab (Magan˜a et al. 1149, XAL). Ver (Lira 15, ENCB, IEB, MEXU, UAMIZ, XAL; Martı´nez C. 2115, CAS, ENCB, MEXU, MO, NY, XAL). Yuc (Castillo Rivero 90, XAL).

This is a fairly recent introduction to the New World and is rapidly becoming naturalized in many countries. About 35 collections have now been seen from Mexico, especially from eastern Chiapas and southern Veracruz, and nearly all have been collected since 1970. In the Greater Antilles, N. hirsutula is now one of the dominant species on roadbanks. Although often misidentified as N. exaltata or N. biserrata, the dense adaxial hairs on the costae, bicolorous (black-centered) appressed rhizome and stipe scales with fimbriate margins, lack of septate hairs on the blades abaxially and adaxially, and narrow whitish scales on the costae abaxially distinguish this taxon. In floristic accounts for New World areas, N. multiflora is usually the name applied to this species, but Bell (in McCarthy, 1998: 444), citing Du Puy in Fl. Australia 50: 568, 1993, noted

Distribution. Terrestrial, epipetric or epiphytic in lowland and montane forests, pine-oak-Liquidambar forests, pine forests, and oak forests; 150–1200(–1700) m. Mexico; Guat, Hond, Nic, CR, Pan; Jam, Cuba; Col, Ven, Ec, Peru, Braz, Bol. Selected Specimens Examined. Chis (Breedlove 21804, DS, 24881, DS, NY; Martı´nez S. 17602, MEXU, NY, UAMIZ). Oax (Mexia 9274,

408

NEPHROLEPIS

CAS, GH, MICH, NY, UC, US; Mickel 4801, 7256, NY, UC). Pue (Campos V. et al. 352, IEB, MEXU). Ver (Palacios-Rios 2307, XAL; Ventura A. 12653, IEB, XAL, 17045, IEB, MEXU, XAL). Unverified, Doubtful, or Mistaken Reports. Sin (reported by Mickel & Beitel, 1988, with a query, but not verified). Tab (reported by Magan˜a, 1992, probably based on a misidentification).

The lunate indusia, pinna bases that are cuneate basiscopically, and essentially parallel-sided pinnae distinguish this species from its most similar congeners (N. pendula and N. undulata) in Mexico. Dı´az-Barriga and Palacios-Rios (1992: 48) cited Arse`ne 6690 (MEXU), from Michoaca´n, as N. pectinata and also as N. cordifolia, but the specimen has not been found at MEXU (March 1998); most likely, it is not N. pectinata.

6. NEPHROLEPIS PENDULA Nephrolepis pendula (Raddi) J. Sm., J. Bot. (Hooker) 4: 197. 1841. Figs. 196A–C. Aspidium pendulum Raddi, Opusc. Sci. 3: 289. 1819. Type. Brazil. Raddi s.n. (FI; isotype K!).

Rhizomes short-creeping to suberect, well developed, lacking tubers; rhizome scales concolorous, linear, castaneous, 2–3 ⫻ 0.5 mm, margins whitish-ciliate; stipes brown, lustrous, glabrous or glabrescent but with a few persistent scales at bases; blades often pendent, 30–100⫹ ⫻ 3.5–7 cm; costae brownish, glabrous except for matted, linear scales at pinna bases; pinnae sessile to shortstalked, bases unequally lobed with basiscopic lobe small and overlapping rachis, acroscopic lobe larger, acute to obtuse, overlapping rachis, pinna apices usually obtuse, margins crenate; indument on both surfaces generally absent; hydathodes often limedotted on adaxial blade surfaces; indusia tan, reniform to semicircular, 1.3–1.5 ⫻ 0.2–0.3 mm, attached along broad, shallow sinuses, mostly opening toward pinna apices; 2n⫽ca. 82 (Galapagos).

Distribution. Epiphytic (rarely terrestrial), pine-oakLiquidambar forests, evergreen forests; 150–550(–1500) m. Mexico; Guat, Bel, Hond, Nic, CR, Pan; Cuba, Hisp, PR; Col, Ven, Guy, Sur, Fr, Gui, Ec, Peru, Braz, Bol.

Selected Specimens Examined. Chis (Breedlove 34481, DS, MO, NY, 48362, CAS, ENCB; Martı´nez S. 15505, MEXU, XAL). Oax (Herna´ndez G. 719, CAS, MO, NY; Williams 9534, GH). Tab (Matuda 3332, MEXU, NY, US; Zamudio 327, ENCB, IEB). Ver (Brigada Dorantes 2733, MEXU, UAMIZ, XAL, 2768, MEXU, MO, UAMIZ, UC; Brigada Va´zquez 699, ENCB, MEXU, MO, XAL).

This species is sometimes treated as a synonym of N. cordifolia, e.g., by R. Tryon (1964: 230), Stolze (1981: 311), and R. Tryon and Stolze (1993: 52), but others regard N. pendula as distinct (e.g., Nauman in Davidse et al., 1995: 288). The two species seem amply different to us, with N. pendula differing in the moderately to densely scaly petioles (proximal third or more), scalier rachises adaxially, glabrous pinnae abaxially, nonoverlapping pinnae, rounded pinna tips, more prominent hydathodes (with lime dots) adaxially, and lack of subterranean tubers. Moreover, the two have distinctly different ranges, with N. cordifolia being essentially absent or only sparingly naturalized in Mexico, Central America, and South America, N. pendula being common in these areas.

7. NEPHROLEPIS RIVULARIS Nephrolepis rivularis (Vahl) Mett. ex Krug in Urban, Bot. Jahrb. Syst. 24: 122. 1897. Figs. 195M–O. Polypodium rivulare Vahl, Eclog. Amer. 3: 51. 1807. Type. Montserrat. Ryan [11] (C!–3 sheets). Nephrolepis neglecta Kunze, Linnaea 13: 149. 1839. Type. Mexico. [Veracruz: ?]. “In rupibus hacienda de la laguna (cueva grande),” Schiede s.n. (LZ, destroyed; probable isotype NY!).

Rhizomes erect, well developed, stoloniferous, lacking tubers; rhizome scales bicolorous, dark reddish brown, centers lustrous, margins tan, ciliate; stipes 8–30 cm long, brown, persistently scaly at bases; blades firm-herbaceous, linear, 50–200 ⫻ (7–)8.5–11 cm, somewhat narrowed at bases; rachises scaly with dense, castaneous, fibrillose scales 2–3 mm long; pinnae many, (8–)9–10 mm wide at bases, (6–)8–9 mm wide at mid-pinna, apices acute to acuminate, bases unequal, basiscopic sides cuneate, acroscopic sides each with an acute basal auricle, margins serrulate toward base, crenate-serrate toward tip; indument on both surfaces lacking or abaxially of sparse castaneous fibrillose scales 0.3–1 mm long; hydathodes lime-dotted on adaxial blade surfaces; indusia dark reddish brown, round-reniform, 0.8–1 mm diam., with distinct sinuses (mature sporangia projecting on all sides); 2n⫽82 (Trin).

NEURODIUM

409

broad, shallow sinuses, mostly opening toward pinna apices; 2n⫽ca. 90, probably erroneous (Nay), 82 (Afr).

Distribution. Terrestrial or epiphytic on hardwood forest; 400– 900 m. Mexico; Guat, Bel (Davidse 36876, UC), Hond, CR, Pan; Gr & L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol. Specimens Examined. Chis (Collins & Doyle 92, US). Oax (Hallberg 1313, 1589, NY, US; Mickel 7162, NY, UC; Vera Santos 3423, ENCB, MICH, US). Ver (Schiede s.n., NY).

The sparse, fibrillose scales on the abaxial blade surfaces, glabrous adaxial surfaces, cuneate pinnae basiscopically, bicolorous, ciliate rhizome scales, and densely scaly rachises distinguish this taxon. Mickel 6460 (NY), cited by Mickel and Beitel (1988: 256) as this species from Oaxaca, is N. pendula.

8. NEPHROLEPIS UNDULATA Nephrolepis undulata (Afzel. ex Sw.) J. Sm., Bot. Mag. 72(Compendium): 35 bis. 1846. Figs. 195A–C. Aspidium undulatum Afzel. ex Sw., J. Bot. (Schrader) 1800(2): 31. 1801. Type. Sierra Leone. Afzelius s.n. (BM!). Nephrolepis occidentalis Kunze, Linnaea 18: 343. 1844. Type. Mexico. Leibold 27 [127] (holotype LZ, destroyed; presumed isotype B!). For a discussion of typification, see Pichi Sermolli (1968). Nephrolepis intermedia Fe´e, Me´m. Foug. 9: 32. 1857. Type. Mexico. [Veracruz:] Orizaba, Schaffner 447 (isotype K!).

Rhizomes erect, almost non-existent, stoloniferous, with oval tubers; rhizome scales concolorous, linear, orange to pale brown, 1.5–3 ⫻ 0.5–1 mm; stipes 4–11 cm long, 1⁄4–1⁄6 the frond length, stramineous, glabrous or with scattered linear scales, grooved; blades thinly herbaceous (veins distinct), linear-elliptic, 22–50 ⫻ 3.5–7 cm; rachises stramineous, glabrous except for sparse concolorous, pale brown, linear to fimbriate hair-like scales 1–1.5 mm long, these mostly at pinna bases; pinnae 15–30(–38) pairs, sessile, narrowly deltate, 2–4.5 ⫻ 0.8–1.2 cm at bases, apices acuminate, bases unequally lobed with basiscopic lobe small, rounded to subcordate, and rarely overlapping rachis adaxially, acroscopic lobe much larger and overlapping rachis abaxially, margins crenate, especially on the fertile fronds, on sterile ones only slightly so; indument on both blade surfaces absent; hydathodes rarely lime-dotted on adaxial blade surfaces; indusia pale brown, lunate, 1.5–2 mm in greatest dimension, attached along

Distribution. Mostly terrestrial in mesic areas on cliffs and along banks and roadsides, occasionally epiphytic on oaks, in pine-oak forests, volcanic areas with clay soils; (400–)800–2100 m. Mexico; Guat, Hond, Salv, Nic, CR, Pan; Gr Ant; Col, Ven, Ec, Peru, Braz, Bol; Africa, se Asia. Selected Specimens Examined. Chis (Purpus 6866, MO, UC, US). Col (Sanders et al. 8277, UC). Dgo (Gonza´lez Ortega 4459, US). Gro (Hinton et al. 9341, MO, NY, US). Jal (Pringle 1835, DS, NY, UC, US). Me´x (Rzedowski 20663, DS, ENCB). Mich (Lyonnet 2772, MEXU, US). Mor (Pringle 15717, CAS, MO, UC, US). Nay (Mexia 707, CAS, DS, MO, NY, UC, US). Oax (Mickel 4727, ENCB, NY, UC). Sin (Breedlove 44956, CAS, ENCB). Tam (Sharp et al. 50158, US). Ver (Pe´rez 1016, UAMIZ).

Nephrolepis undulata is the most widespread member of its genus in Mexico, and is distinct in its weakly developed rhizomes, presence of tubers below ground, entire rhizome scales, thin-textured pinnae with distinct veins, small number of pinnae (usually fewer than 35 pairs), pinnae tapering from bases to acute apices, and large lunate indusia 1.5–2 mm wide. The presence of perennating, underground tubers enables this species to survive burning and helps perpetuate an annual flush of growth with the advent of the rainy season; in times of drought, the plants disappear aboveground. This species is usually treated under the name N. occidentalis (e.g., by Smith, 1981; Mickel & Beitel, 1988), but we find no essential differences between the paleotropical and neotropical specimens of this very widespread species. R. Tryon and Stolze (1993: 53) believed that there exists intergradation between N. undulata (as occidentalis) and N. pectinata in South America, but we see no evidence of such intermediates in Mexico and find the two taxa amply distinct throughout their ranges. Breedlove 28698 (DS), from Chiapas, is an aberrant (dissected) form of the species.

82. NEURODIUM Neurodium Fe´e, Me´m. Foug. 3: 11, 28. 1852. [“Nevrodium”]. Type: Neurodium lanceolatum (L.) Fe´e [⬅ Pteris lanceolata L.].

410

NIPHIDIUM

Paltonium C. Presl, Epimel. Bot. 156. 1849 [1851]. Type: Paltonium lanceolatum (L.) C. Presl [⬅ Pteris lanceolata L.]. See Pichi-Sermolli (Webbia 9: 361–366. 1953) for a discussion of publication dates.

Epiphytic; rhizomes short-creeping, dictyostelic, scaly; fronds approximate, articulate to rhizomes, short-stalked, simple, entire-margined; veins immersed, not easily seen, reticulate with simple or hamate excurrent and recurrent included veinlets; leaf tissue glabrous; sori exindusiate; sporangia submarginal, coenosoral, confined to distal part of blades, which are usually somewhat contracted; paraphyses none or filamentous; spores bilateral, hyaline, minutely tuberculate; x⫽37. Neurodium is distinct among Mexican polypodioid ferns by its undivided blades with marginal sori. Traditionally it is treated as monotypic, but Pleopeltis repanda A. R. Sm. from Venezuela is certainly congeneric and probably the closest ally, even if Neurodium is accepted as a good genus. Molecular studies (Schneider et al., submitted) have shown Neurodium to be sister to Dicranoglossum, and both are nested in the Pleopeltis clade (scaly bladed polypods).

Distribution. Epiphyte in mangrove swamps, tropical rain forests, lower montane rain forests; 0–300 m. USA (Fla); Mexico; Guat, Bel, Hond, Nic, CR; Bah, Gr & L Ant; Fr Gui. Specimens Examined. Chis (Breedlove 29761, DS, 49135, CAS; Martı´nez S. 19100, MEXU; Mell 2047, US). Tab (Cowan & Magan˜a 3258, CAS, CSAT, ENCB, MEXU, NY; Rovirosa 979, NY).

Reference Schneider, H., A. R. Smith, R. Cranfill, T. J. Hildebrand, C. H. Haufler & T. A. Ranker. 2004. Unraveling the phylogeny of the polygrammoid ferns (Polypodiaceae & Grammitidaceae): Exploring aspects of the diversification of epiphytic plants. Molec. Phylogen. Evol. (in press).

1. NEURODIUM LANCEOLATUM Neurodium lanceolatum (L.) Fe´e, Me´m. Foug. 3: 28. 1852. Figs. 197A–D. Pteris lanceolata L., Sp. Pl. 2: 1073. 1753. Taenitis lanceolata (L.) Kaulf., Enum. Filic. 130. 1824. Paltonium lanceolatum (L.) C. Presl, Epimel. Bot. 156. 1849 [1851]. Lectotype (chosen by Proctor 1977: 346). Plumier, Descr. Pl. Ame´r., pl. 40. 1693, illustrating a plant from Hispaniola.

Roots brown, in dense mass, often covering stems, densely clothed with root hairs; rhizome scales imbricate, ovate, entire, dark brown, clathrate; fronds numerous, rigidly ascending, erectarching, 20–40 cm long; stipes 1–5 cm long, tan, narrowly alate distally; blades 1.5–3(–4) cm wide at middle, coriaceous, longattenuate at both ends, distal half or third sometimes fertile; costae prominent abaxially, stramineous; margins entire to repand; leaf tissue coriaceous, bright green, lustrous, without scales; sori 2–3 mm wide, extending almost to the tip; 2n⫽74 (Jam).

8 3. N I P H I D I U M Niphidium J. Sm., Hist. Fil. 99. 1875. Type: Niphidium americanum (Hook.) J. Sm. [⬅ Polypodium americanum Hook.] ⫽ Niphidium longifolium (Cav.) C. Morton & Lellinger. Anaxetum Schott, Gen. Fil., t. 1. 1834, hom. illeg., non Anaxeton Gaertner, 1791. Pessopteris Underw. & Maxon, Contr. U.S. Natl. Herb. 10: 485. 1908. Type: Anaxetum crassifolium (L.) Schott [⬅ Polypodium crassifolium L.] ⬅ Niphidium crassifolium (L.) Lellinger

Epiphytic or terrestrial; rhizomes short- to long-creeping; rhizome scales concolorous to bicolorous, clathrate, attached near base or peltate, surfaces glabrous, margins entire (in ours) to toothed; fronds simple, approximate to distant, monomorphic, stipitate, narrowly lanceolate to oblong, attenuate at bases and apices, articulate, margins entire; blades coriaceous, glabrous, often glaucous, sometimes with a few scales along the costae (in ours); venation areolate, inconspicuous transverse areoles with minor areoles containing free veins; sori round to slightly oblong, exindusiate, apparently at the junction of veins (actually veins forming an irregular ring beneath the sori), solitary in major areoles, usually in one row between main lateral veins; paraphyses (abortive sporangia) present (in ours) or not; sporangia with multicellular hairs (in ours) or glabrous; spores bilateral, smooth; x⫽37. Niphidium is a neotropical genus of ten species within the polypodioid ferns and is most closely related to Campyloneurum (Schneider et al., 2004). Most species have restricted ranges in South America; one species occurs in Mexico. Lellinger (1972) united Niphidium and Pessopteris, genera previously regarded as distinct. Niphidium is characterized by the large, strap-like fronds with a single row of large, round sori between the major lateral veins and intricate venation with multiple veins to each sorus.

NOTHOLAENA Reference Lellinger D. B. 1972. [1973]. A revision of the fern genus Niphidium. Amer. Fern J. 62: 101–120.

1. NIPHIDIUM CRASSIFOLIUM Niphidium crassifolium (L.) Lellinger, Amer. Fern J. 62: 106. 1972 [1973]. Figs. 198A–F. Polypodium crassifolium L., Sp. Pl. 2: 1083. 1753. Pessopteris crassifolia (L.) Underw. & Maxon, Contr. U.S. Natl. Herb. 10: 485. 1908. Lectotype (chosen by Lellinger, 1972: 106). Petiver, Pterid. Amer., t. 6, f. 1. 1712, which is a redrawing of Plumier, Traite´ Foug. Ame´r., pl. 142, 1705, illustrating a plant from Martinique.

Rhizomes short-creeping, 7–10 mm diam.; rhizome scales 5–7 ⫻ 1.5–2 mm, clathrate, margins membranous, pale, entire; fronds

35–125 cm long, clumped; stipes absent to 30 cm long; blades narrowly elliptic to oblanceolate, bases gradually narrowed, apices acuminate, 4.5–11(–14) cm wide, glabrous, coriaceous, rachises with a few scattered scales 2–3 mm long; main lateral veins prominent to blade margins, minor veins netted, crossveins obscure; sori large, round, mostly ca. 3(2–5) mm diam., solitary in major areoles, forming single rows of 6–10 sori between main veins; sporangia setose (several short hairs per sporangium), paraphyses usually common among sporangia. 2n⫽148 (Jam).

411

Selected Specimens Examined. Chis (Breedlove 22426, DS, MEXU, NY; Matuda 2770, US). Gro (Lorea 2995, FCME). Hgo (Sa´nchezMejorada 921, MEXU). Oax (Mickel 4236, ENCB, NY; Wendt et al. 5133, CAS, CHAPA, NY). Pue (Lo´pez-Forment 226, MEXU; Sa´nchezMejorada 607, US). Qro (Arse`ne 10656, US). Ver (Lira 133, MEXU, UAMIZ).

84. NOTHOLAENA Notholaena R. Br., Prodr. 145. 1810. Lectotype (chosen by J. Smith, Hist. Fil. 278. 1875): Notholaena trichomanoides (L.) Desv. [⬅ Pteris trichomanoides L.]. For details of typification, see Yatskievych & Smith (2003). Notholaena sect. Chrysochosma J. Sm., Hist. Fil. 279. 1875. Chrysochosma (J. Sm.) Ku¨mmerle, Magyar Bot. Lapok 13: 35. 1914. Type: Notholaena sulphurea (Cav.) J. Sm. [⬅ Pteris sulphurea Cav.]

Terrestrial or epipetric; rhizomes compact to short-creeping, scaly; roots wiry; fronds monomorphic, small to rarely mediumsized, clumped; blades 1-pinnate-pinnatifid to 3-pinnate, generally farinose abaxially (the farina being a white to yellow waxy substance produced by short capitate glandular hairs), occasionally also bearing hairs and/or scales; veins free; sori at vein ends, forming marginal bands, exindusiate; blade margin only slightly curved, undifferentiated (somewhat curved and thin in N. rosei); spores tetrahedral-globose; gametophytes glandular-farinose; x⫽30. Of the approximately 30 species in the genus, 24 occur in Mexico, on or among rocks, usually in warm, seasonally dry regions. Notholaena has been treated in various ways in the past, most notably to include what we now treat as Argyrochosma (R. Tryon, 1956; Windham, 1986), or included within Cheilanthes (Mickel & Beitel, 1988; Mickel, 1991). Molecular data (Gastony & Rollo, 1995) suggest Notholaena is monophyletic with Cheiloplecton closely related. Cheilanthes aurea and C. aurantiaca may belong in Notholaena. References

Distribution. Epiphytic or epipetric, montane rain forests, pine-oak and Liquidambar forests, tropical rain forests, seasonal evergreen forests; 120–1800 m. Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol, Parag.

Hall, C. C. 1950. Notholaena copelandii, a newly recognized species of the Texano-Mexican region. Amer. Fern J. 40: 178–187; Seigler, D. S. & E. Wollenweber. 1983. Chemical variation in Notholaena standleyi. Amer. J. Bot. 70: 790–798; Tryon, R. M. 1956. A revision of the American species of Notholaena. Contr. Gray Herb. 179: 1–106; Windham, M. D. 1986. [Abstract] Reassessment of the phylogenetic relationships of Notholaena. Amer. J. Bot. 73: 742; Wollenweber, E. 1984. Exudate flavonoids of Mexican ferns as chemotaxonomic markers. Rev. Latinoamer. Quim. 15: 3–11; Yatskievych, G. & A. R. Smith. 2003. Typification of Notholaena R. Br. (Pteridaceae). Taxon 52: 331–336.

Key to the Mexican Species of Notholaena 1. Blade indument of farina and of scales and/or hairs, the hairs occasionally limited to the adaxial surfaces of the stipes and rachises. 2. Abaxial surfaces of blades and rachises with large, brown, stiff hairs or with only farina, not with both farina and hairs or scales. 3. Short hairs limited to adaxial surfaces of stipes and rachises. ............................................................................................... 17. N. meridionalis. 3. Bristle-like hairs on abaxial blade surfaces, stipes, and rachises. 4. Blades bipinnate-pinnatifid; blade scales ciliate-denticulate (like those of rhizomes) on the rachises and other blade axes and veins, often reduced to bristle-like hairs. ............................................................................................................. 21. N. schaffneri.

412

NOTHOLAENA 4. Blades pinnate-pinnatifid; blade scales reduced, bristle-like, sparse to scattered, not toothed except toward bases of fronds. 5. Stipes 1⁄5–1⁄4 the frond length; blades 0.9–1.4 cm wide, linear; pinnae (16–)20–35 pairs; rachises puberulent with erect, stalked, glandular hairs 0.1 mm or less, especially adaxially, also with bristle-like scales. ................................. 1. N. affinis. 5. Stipes 1⁄30–1⁄10 the frond length; blades 2–3.3 cm wide, narrowly elliptic; pinnae(11–)14–25 pairs; rachises lacking stalked glandular hairs (other than farina), with or without short non-glandular hairs adaxially. 6. Farina white; pinnae with 3–5 pairs of lobes; blades with (11–)17–25 pinna pairs. ........................................ 6. N. brevistipes. 6. Farina pale yellow; pinnae with 5–7 pairs of lobes; blades with 14–16 pinna pairs. .................................... 5. N. brachycaulis. 2. Abaxial surfaces of blades and rachises with farina and scales, sometimes also with hairs (in N. weatherbiana, the tomentum more evident than the few scales). 7. Scales on abaxial blade surfaces long-ciliate or dissected. 8. Adaxial blade surfaces with stellate hairs. ...................................................................................................................... 3. N. aschenborniana. 8. Adaxial blade surfaces with sparse, weak, simple hairs. ........................................................................................................ 11. N. galeottii. 7. Scales on abaxial blade surfaces entire or nearly so. 9. Farina orange; scales on abaxial blade surfaces large, broadly ovate; blades pentagonal, the basal pinnae much the largest and strongly inequilateral. ......................................................................................................................................... 4. N. aurantiaca. 9. Farina white to yellow; scales on abaxial blade surfaces lanceolate or narrower; blades much longer than broad, the proximal pinnae not much the largest or strongly inequilateral. 10. Adaxial blade surfaces somewhat farinose and with orange-tan scales on blade axes; hairs absent. .......................... 12. N. grayi. 10. Adaxial blade surfaces with sparse hairs, slightly or not farinose, lacking scales. 11. Abaxial blade surfaces scaly (many scales bristle-like but not forming a tomentum); adaxial surfaces sparsely villous and sparingly farinose; rachises scaly, with the few hairs not forming a tomentum. .............................. 2. N. aliena. 11. Abaxial blade surfaces with many curly white hairs, with only a few scales; adaxial surfaces persistently (although thinly with age) lanose with curly, white hairs, non-farinose; rachises scaly and tomentose. ............................................................................................................................................................................. 24. N. weatherbiana.

1. Blade indument of farina only. 12. Basal pinnae not exaggerated basiscopically; blades linear to narrowly deltate, much longer than broad; pinnae (4–)8–18 pairs. 13. Stipes and rachises terete. 14. Rhizome scales entire, glandular-margined, reddish brown to black with age, with tan margins. ................................ 16. N. leonina. 14. Rhizome scales ciliate-denticulate, black, sclerotic. ................................................................................................................. 19. N. rigida. 13. Stipes and rachises grooved adaxially. 15. Segments lobed, margins plane; blade apices pinnatifid. 16. Blades pinnate-pinnatifid to bipinnate; stipes and rachises light brown to black. ................................................ 15. N. lemmonii. 16. Blades bipinnate-pinnatifid to tripinnate; stipes and rachises stramineous. .................................................................. 20. N. rosei. 15. Segments entire, ovate-oblong, margins recurved; blade apices pinnate. 17. Rhizome scales with black sclerotic central band and pale erose-denticulate margins, those of stipes lanceolateovate; blades about equalling stipes in length; farina white; Dgo, Chih, Coah. ........................................................ 13. N. greggii. 17. Rhizome scales subconcolorous bright orange-brown with subsclerotic central band and entire margins, those of stipes linear-lanceolate; blades longer than stipes; farina pale yellow or cream-colored; Coah, NL. .................... 7. N. bryopoda. 12. Basal pinnae exaggerated basiscopically; blades sometimes broadly pentagonal to deltate pentagonal, 2⁄3–3 times as long as broad; pinnae 4–9 pairs. 18. Blades 11⁄2–3 times as long as broad. 19. Basal pinnae each with more than one exaggerated basiscopic pinnule, the basiscopic pinnules on basal pinna pair gradually decreasing distally. ..................................................................................................................................................... 9. N. candida. 19. Basal pinnae with a single exaggerated basiscopic pinnule. 20. Blade apices abruptly narrowed, apices resembling a more or less discrete terminal pinna; blades 4–7.5 cm wide; farina white; segments 4–8 mm long; proximal pinna pairs nearly touching; Coah, Hgo, NL, Tam. ............. 10. N. copelandii. 20. Blade apices gradually narrowed distally, blades 2.2–4 cm wide; farina pale yellow or white; segments 2–3.5 mm long; proximal pinna pairs widely spaced; Hgo, SLP. ................................................................................................ 14. N. jacalensis. 18. Blades 2⁄3–1 1⁄4 times as long as broad. 21. Blades above proximal pinnae usually pinnatifid, the first suprabasal pinna pair usually shortened. .......................... 22. N. standleyi. 21. Blades above proximal pinnae fully pinnate to pinnate-pinnatifid, the pinnae decreasing gradually in size upward. 22. Stipes and rachises reddish brown to dark brown; BC, Son, Guad. ....................................................................... 8. N. californica. 22. Stipes and rachises blackish; not BC. 23. Ultimate segments fully adnate. ............................................................................................................................ 23. N. sulphurea. 23. Ultimate segments constricted at bases, narrowly adnate to petiolulate. .......................................................... 18. N. neglecta.

NOTHOLAENA ASCHENBORNIANA

413

1. NOTHOLAENA AFFINIS

2. NOTHOLAENA ALIENA

Notholaena affinis (Mett.) Hook. ex T. Moore, Index Fil. 233. 1861. Figs. 204H, J, K.

Notholaena aliena Maxon, Contr. U.S. Natl. Herb. 17: 605. 1916. Figs. 206J–M.

Cheilanthes affinis Mett., Abh. Senckenberg. Naturf. Ges. 3: 63. 1859. Aleuritopteris affinis (Mett.) E. Fourn., Mexic. Pl. 1: 21. 1872. Chrysochosma affinis (Mett.) Pic. Serm., Webbia 43: 307. Type. Mexico. [Morelos?:] Aschenborn s.n. (B, photos GH!, US!). Notholaena angusta R. M. Tryon, Rhodora 63: 80. 1961. Chrysochosma angusta (R. M. Tryon) Pic. Serm., Webbia 43: 307. 1989. Type. Mexico. Hidalgo: ca. 1 km S of Tasquillo Bridge, Frye & Frye 3145 (US!; isotypes DS!, NY!, UC!, WASH!).

Rhizomes compact, horizontal, 1–2 mm diam.; rhizome scales linear-lanceolate, 1.5–2 mm long, reddish black, concolorous, margins ciliate-denticulate; fronds 3.4–24 ⫻ 0.9–1.4 cm, clumped; stipes 1/5–1⁄4 the frond length, reddish brown to atropurpureous, terete, with scattered to sparse linear-lanceolate scales 1–1.5 mm long, short sharp-pointed hairs 0.2–0.5 mm long, and glandular hairs 0.1 mm long, farina absent; rachises with hairs concentrated adaxially, long hairs/reduced scales pointed, 1 mm, shorter hairs gland-tipped, 0.2 mm, very short glandular hairs 0.1 mm or less; blades linear, 9–14 mm broad, pinnate-pinnatifid; pinnae (16–)20–35 pairs, equilateral, stalked 0.4–0.7 mm, articulate, pigment entering the pinnae; adaxial surfaces with scattered glands; abaxial surfaces with white farina and sparse dark hairs; laminar margins slightly curved, undifferentiated; spores black or tan.

Chrysochosma aliena (Maxon) Pic. Serm., Webbia 43: 307. 1989. Type. Mexico. Coahuila: Soledad, 25 mi SW from Monclova, Palmer 1389 (US! photo GH; isotypes GH!, K!, NY!).

Rhizomes compact, horizontal, 1–2 mm diam.; rhizome scales linear, attenuate at apices, concolorous, dark brown to black, sclerotic, to weakly bicolorous, margins with weak pale hairs, 1.5–2.5 mm long; fronds 14–22 ⫻ 1–1.5 cm, clumped; stipes ca. 1⁄3 the frond length, terete (rachises grooved), castaneous, with scattered scales like those of rhizomes, lax pale hairs to 1 mm long, and minute glandular hairs; blades linear-oblong, pinnatepinnatifid; pinnae 8–12 pairs, pinna lobes 4–5 pairs; adaxial blade surfaces minutely farinose and sparsely villous with lax hairs; abaxial blade surfaces densely pale yellow-farinose, with scattered to dense orange-tan linear scales and hairs 1–1.5 mm long; blade margins slightly curved; sporangia 16-spored; spores tan to black.

Distribution. Rocky slopes and cliffs and ledges, usually on igneous rocks; 700–1200 m in USA. USA (Tex); Mexico. Specimens Examined. Chih (Burgess 5216, ARIZ; Pringle 463, GH p.p.; Reeves R57232, ASC). Coah (only the type collection). Tam (Bartlett 10999, MICH, US).

Distribution. Among basaltic rocks; 1500–2450 m. Mexico. Selected Specimens Examined. Gto (Carranza G. 3732, NY). Hgo (Broun s.n., PH, photo NY; Hitchcock & Stanford 7249, US; Matuda 23571, MEXU, US; Pray 3104, LAM, NY). Qro (Dı´az B. & Carranza 7460, NY; Go´mez 487, 5806, IEB; Labbat 2603, IEB). Tam (Johnston 4948H, TEX).

Dı´az B. & Carranza 7640, NY, from Quere´taro has more scales and glandular hairs on the rachis and lacks white farina. Plants called N. affinis from further south are herein referred to N. meridionalis. Notholaena affinis is distinguished from N. meridionalis by the linear blades and more numerous pinnae.

Notholaena aliena is closely related to N. grayi but is readily distinguished in having the adaxial surfaces of the blades distinctly villous with sparse, lax, whitish, tortuous hairs. Notholaena grayi is only sparsely farinose on the adaxial blade surfaces and is devoid of hairs. Notholaena aliena differs from the closely allied N. weatherbiana by the latter having darker brown stipes, scales and hairs on the stipes, inconspicuous glands on the adaxial surfaces of the pinnules, and finely white-villous blades on both sides.

3. NOTHOLAENA ASCHENBORNIANA Notholaena aschenborniana Klotzsch, Linnaea 20: 417. 1847. Figs. 201K–N.

414

NOTHOLAENA

Cheilanthes aschenborniana (Klotzsch) Mett., Abh. Senckenberg. Naturf. Ges. 3: 21. 1859. Chrysochosma aschenborniana (Klotzsch) Pic. Serm., Webbia 43: 307. 1989. Type. Mexico. Distrito Federal: Chapultepec, Aschenborn 593 (B!; isotypes P!, photo GH, US!). Notholaena bipinnata Liebm., Mexic. Bregn. 214 (reprint 62). 1849. Type. Mexico. Puebla: Tehuaca´n, Liebmann 2455 (not found at C; isotype US!).

Rhizomes compact, horizontal, 2–3 mm diam.; rhizome scales linear-lanceolate, ca. 3 mm long, black, concolorous, sclerotic, lustrous, margins ciliate-denticulate; fronds 21–28 ⫻ 2–6 cm, clumped; stipes ca. 1⁄4 the frond length, black, terete, with scattered, dark brown, linear-lanceolate, ciliate-denticulate scales 3 mm long (these especially abundant near the base of the stipe), and brown to whitish highly dissected scales (resembling stellate hairs); blades narrowly lanceolate, bipinnate to bipinnatepinnatifid, not strongly narrowed proximally; pinnae 12–20 pairs, equilateral, the segments oblong, obtuse; adaxial surfaces with dense, stellate, white hairs; abaxial surfaces with a slight farina covering, densely clothed with brown, linear-lanceolate, long-ciliate scales 0.5–1 mm long, and pale cilia covering the surfaces and sori; blade margins somewhat curved; sporangia 32spored; spores orange-tan; n⫽2n⫽90, apogamous (NL, SLP, Tam, USA).

Distribution. In oak woodlands, rocky slopes and cliffs, mostly on limestone, less commonly on basalt; 900–3000 m. USA (s Ariz, w Tex); Mexico. Selected Specimens Examined. Ags (McVaugh 16682, NY). Chih (Pringle 469, US). Coah (Hinton 16561, NY). Dgo (Sa´nchez 737, MEXU). Hgo (Edwards 892, US). Mor (Pacheco 3439, UAMIZ). NL (Aguirre s.n., 6 Dec 1976, NY). Oax (Lorence 3359, MEXU). Pue (Chiang et al. 2103, MEXU). Qro (Dı´az B. 3774, IEB). SLP (Purpus 5486, GH). Son (White 533, MICH). Tam (Stanford et al. 1983, NY). Zac (Johnston et al. 10468a, MEXU).

Notholaena aschenborniana is sometimes described as nonfarinose, but the abaxial glandular farina characteristic of all species of Notholaena is present beneath the dense covering of scales. Windham (in FNA Ed. Comm., 1993) reported this species to comprise mostly apogamous triploids that apparently arose through autopolyploidy; it is possible that 64-spored, diploid populations of N. aschenborniana may exist. This species is similar to N. galeottii but is distinct in the stellate hairs on the adaxial blade surfaces.

4. NOTHOLAENA AURANTIACA Notholaena aurantiaca D. C. Eaton, Proc. Amer. Acad. Arts 22: 462. 1887. Figs. 205A–G. Chrysochosma aurantiaca (D. C. Eaton) Pic. Serm., Webbia 43: 307. 1989. Cheilanthes aurantiaca (D. C. Eaton) Mickel in McVaugh, Fl. Nov.-Gal. 17: 217. 1992, hom. illeg., non (Cav.) T. Moore, 1857. Type. Mexico. Jalisco: Barrancas near Guadalajara, Palmer 83 (YU!, photo GH; isotypes MEXU!, NY!, US!).

Rhizomes compact, horizontal, 1.5–2 mm diam.; rhizome scales lanceolate-deltate, 2–3 mm long, concolorous, dark reddish brown, sclerotic, lustrous, entire; fronds 8–16 cm long, often 3– 5 mm distant; stipes ca. 2⁄3–4/5 the frond length, atropurpureous, terete, with scattered to sparse, linear-lanceolate, sclerotic redbrown scales, these grading distally to thin, tan scales 1–2 mm long; blades pentagonal, 3–6.5 cm wide, 2–3-pinnatifid, the proximal pair of pinnae exaggerated basiscopically; pinnae 7–12 pairs, linear-lanceolate, 2–3 ⫻ 1–1.5 mm; adaxial surfaces with orange farina and dense, large, broadly ovate scales, tan with sclerotic red-brown base; abaxial surface with scattered, jointed hairs 0.5–1 mm long; blade margins plane, undifferentiated; spores gray.

Distribution. On basaltic rocks in low deciduous woods; 1200– 1400 m. Mexico. Specimens Examined. Jal (Pray 1928, LAM, 3130a, LAM; Pringle 1840, UC, 2587, UC, US, 5405, MEXU). Nay (Tenorio L. 16163, MEXU). Unverified, Doubtful, or Mistaken Reports. Ags (Siqueiros 4294, HUAA, and two other collections, cited by Siqueiros-Delgado & Gonza´lez-Adame, 2004 [as Cheilanthes aurantia], but not verified).

Cheilanthes aurantiaca is distinct by the pentagonal, pinnatepinnatifid blades, gold-orange farina, and broad scales on the abaxial surfaces of the pinnae.

5. NOTHOLAENA BRACHYCAULIS Notholaena brachycaulis Mickel, sp. nov. Type. Mexico. Nuevo Leo´n: North side of Cerro Sombreretillo, 26⬚19'N, 99⬚59'W, 610 m, 27 Apr 1949, Clausen 7619 (holotype MEXU!; isotypes MEXU!, US!). Figs. 204L–N.

NOTHOLAENA BRYOPODA A N. brevistipes paribus pinnarum 14–16 (non (11–)17–25), paribus lobis pinnarum 5–7 (non 3–5), et farina aurantiaca distincta. (Gr., brachys, short, caulis, foot, alluding to the short stipes.)

415

petiolate 0.5–1 mm, pinna lobes 3–5 pairs; adaxial surfaces slightly farinose; abaxial surfaces white-farinose; laminar margins slightly curved, undifferentiated; spores tan.

Rhizomes compact, 3–4 mm diam.; rhizome scales linearlanceolate, ca. 3 mm long, orange-tan with narrow dark reddish brown ciliate margins, to concolorous dark reddish brown with age; fronds 10–20 cm long, clumped; stipes absent to 1/10 the frond length, atropurpureous, terete, with scattered scales like those of rhizomes; blades narrowly elliptic, pinnate-pinnatifid, 2–3 cm wide; rachises terete, slightly farinose, lacking hairs but with sparse bristles or toothed scales; pinnae 14–16 pairs, equilateral, short-petiolate ca. 0.5 mm, pinna lobes 5–7 pairs; adaxial blade surfaces slightly farinose; abaxial blade surfaces with pale yellow farina; laminar margins slightly curved, undifferentiated; spores tan or black.

Distribution. Dry, rocky slopes; 230–610 m. Mexico. Specimen Examined. Tam (Martı´nez Martı´nez & Luyando F-1782, TEX).

See comments under N. brachycaulis for comparison with that species.

7. NOTHOLAENA BRYOPODA

Distribution. Sandstone crevices along stream; 610 m. Known only from the type collection. Notholaena brachycaulis closely resembles N. brevistipes in the very short stipes, but the former differs in having 14–16 pinna pairs (vs. (11–)17–25), 5–7 pairs of pinna lobes (vs. 3–5), and yellowish farina (vs. white).

6. NOTHOLAENA BREVISTIPES Notholaena brevistipes Mickel, sp. nov. Type. Mexico. Tamaulipas: Hwy 180, 8.5 km N of Aldama, ca. 0.5 km N of the dirt road marked “La Coma,” Mayfield, Hemple & Jack 786 (holotype UC!; isotypes LL!, TEX!). Figs. 199D–H, J. A N. affinis pilis et squamis reductis abstans. (L., brevis, short, stipes, stipe, alluding to the short stipes.)

Rhizomes compact, horizontal, ca. 3 mm diam.; rhizome scales linear-lanceolate, 3–4 mm long, dark reddish brown, concolorous, sclerotic, lustrous, margins stiffly ciliate-denticulate; fronds (14–)20–27 cm long, clumped; stipes 1⁄30–1/12 the frond length, atropurpureous to castaneous, terete, with scattered scales like those of rhizomes; blades narrowly elliptic, pinnate-pinnatifid, 2.2–3.3 cm wide, rachises terete, puberulent on adaxial surfaces with short, erect brown hairs 0.3–0.6(–1) mm long, also with scattered, stiff, brown, spreading hairs or narrow scales 0.8–1.7 mm long; pinnae (11–)17–25 pairs, equilateral, very short-

Notholaena bryopoda Maxon, Proc. Biol. Soc. Wash. 15: 205. 1905. Type. Mexico. Nuevo Leon: Sierra de San Lorenzo, Pringle 8802 (US!; isotypes B, BM, CAS!, F!, G, GH!, K!, MEXU!, MO!, NY!, P!, PR!, UC!, US!). Figs. 202A–F. Rhizomes compact, horizontal, ca. 2 mm diam.; rhizome scales linear-lanceolate, 5–9 mm long, orange-brown, concolorous to weakly bicolorous, the center castaneous to nearly black with age, lustrous, entire; fronds 10–18 ⫻ 2–3 cm broad, clumped; stipes 1⁄3 the frond length, brown, grooved, with scattered to sparse linear-lanceolate scales 1–1.5 mm, lacking farina; blades lanceolate to narrowly deltate, 2-pinnate, coriaceous; pinnae 5– 11(–13) pairs, equilateral; adaxial surfaces glabrous; abaxial surfaces with very pale yellow or cream-colored farina; blade margins curved, covering about half the abaxial surfaces, undifferentiated; spores brown to black; 2n⫽60 (NL).

416

NOTHOLAENA

Distribution. On gypsum outcrops, calcite, limestone gravel, slopes; 1100–2100 m. Mexico. Selected Specimens Examined. Coah (Hinton 16543, NY; Johnston et al. 12156, NY; Stewart 839, US). NL (Aguirre s.n., 23 Apr 1976, ENCB; Dorr 2552, ARIZ, 2612, NY; Nesom 4270, ENCB; Sa´nchez 3872, MEXU; Villareal 4362, ASC, NY; Yatskievych 83-308, ARIZ).

See N. greggii for comparison with that species.

8. NOTHOLAENA CALIFORNICA Notholaena californica D.C. Eaton, Bull. Torrey Bot. Club 10: 27. 1883. Cheilanthes desertii Mickel, Phytologia 41: 434. 1979, non Cheilanthes californica Mett., 1859. Chrysochosma californica (D.C. Eaton) Pic. Serm., Webbia 43: 307. 1989. Lectotype (chosen by R. Tryon, 1956: 73). U.S.A. California: San Diego Co., Spring Valley, in 1878, Burbeck s.n. (YU!, photo GH!).

Rhizomes compact, horizontal, 1.5–2.5 mm diam.; rhizome scales linear-lanceolate, dark reddish brown to black, sclerotic, 2–3 mm long, concolorous or with narrow pale margins, weakly short-ciliate-denticulate; fronds 4–14 ⫻ 1.5–4 cm, clumped; stipes 1 ⁄2–3⁄4 the frond length, reddish brown to dark brown, terete, rachises grooved, with scattered scales at bases like those of rhizomes, otherwise glabrous; blades pentagonal, 3-pinnate at base, 2-pinnate distally, coriaceous; pinnae ca. 5 pairs of divided pinnae, equilateral except the basal pair strongly exaggerated basiscopically, segments more or less ovate, somewhat bead-like, narrowly adnate to free; adaxial surfaces with scattered white or pale to bright yellow farina; abaxial surfaces with dense white or pale to bright yellow farina; blade margins curved; sporangia 32- or 64-spored; spores tan to black.

Distribution. Rocky slopes and cliffs, usually on volcanic rocks in desert scrub; 100–1100 m. USA (Ariz, Calif); Mexico. Selected Specimens Examined. BCN (Orcutt s.n., 25 Jan 1889, US; Wiggins & Thomas 123, ENCB). BCS (Wiggins 17831, DS p.p.). Guad (Kipping 307, CAS). Son (Felger 16799, ARIZ p.p., 17923, 89-18, 20227, ARIZ, 19689, 20008, ENCB).

8b. Notholaena californica subsp. leucophylla Windham, Contr. Univ. Michigan Herb. 19: 35. 1993. Type. Mexico. Baja California: 4 mi N of rancho Mesquital, Wiggins 16187 (US!; isotypes DS!, MICH!, RSA). Blades with white farina on abaxial surfaces. Sporangia 64spored; sexual diploids, triploids, and perhaps tetraploids.

Distribution. Epipetric on rocky slopes and cliffs; 100–1100 m. USA (Ariz, Calif); Mexico. Key to the Mexican Subspecies of Notholaena californica 1. Farina yellow. .......................................................... 8a. subsp. californica. 1. Farina white. ...........................................................8b. subsp. leucophylla.

8a. Notholaena californica subsp. californica Figs. 202O–R. Notholaena californica D. C. Eaton subsp. nigrescens Ewan, Amer. Fern J. 32: 93, t. 10, f. b. 1942. Type. U.S.A. California: Los Angeles Co., San Gabriel Canyon, Wheeler 878 (COLO; isotypes LAM, NO). Additional synonymy in R. Tryon (1956: 73).

Blades with pale to bright yellow farina on abaxial surfaces; n⫽2n⫽150, apogamous (USA).

Distribution. Rocky slopes and cliffs, usually on granitic or volcanic substrates; 100–1100 m. USA (Calif); Mexico. Selected Specimens Examined. BCN (Moran 7240, ENCB; Wiggins 15081, DS). BCS (Johnston 4007, US; Moran 9604, US; Turner & Burgess 6227, MEXU; Wiggins 16129, US). Cedros Island (Mason 2001, CAS, US; Palmer 748, US; Rose 16852, US). Son (Palmer 552, US).

9. NOTHOLAENA CANDIDA Notholaena candida (M. Martens & Galeotti) Hook., Sp. Fil. 5: 110. 1864. Figs. 203A–D. Cheilanthes candida M. Martens & Galeotti, Me´m. Foug. Mexique 73, pl. 20, f. 1b. 1842. Aleuritopteris candida (M. Martens & Galeotti) Fe´e, Me´m.

NOTHOLAENA GALEOTTII Foug. 5: 154. 1852. Chrysochosma candida (M. Martens & Galeotti) Ku¨mmerle, Magyar Bot. Lapok 13: 40, 42. 1914. Type. Mexico. Jalisco: N of Guadalajara, Galeotti 6442 (BR!, photo BM!; isotype BR!).

Rhizomes compact, horizontal, ca. 2 mm diam.; rhizome scales lanceolate, 2 mm long, bicolorous, with castaneous to black central streak and pale margins, entire to suberose; fronds 10–34 cm long; stipes 1⁄2–2⁄3 the frond length, atropurpureous to black, terete, with scattered ovate-lanceolate scales at bases or to basal pinnae, also with sparse white farina; blades ovate-pentagonal to deltate-pentagonal, 4–7(–10) cm wide, pinnate-pinnatifid except for the basal pair of pinnae which are 2-pinnate-pinnatifid and exaggerated basiscopically (basal pinnule of suprabasal pinnae separated from next pinnule on same pinna by dark pinna rachis), the basiscopic pinnule on basal pinna pair gradually decreasing distally; pinnae 4–8 pinnatifid pairs above basal pinnae; adaxial surfaces glabrous or with a few scattered farina glands; abaxial surfaces white-farinose, rarely pale yellow; blade margins slightly curved; sporangia 64-spored; spores tan or black; 2n⫽60 (Mor, Oax, Pue).

417

Notholaena candida (M. Martens & Galeotti) Hook. var. copelandii (C.C. Hall) R. M. Tryon, Contr. Gray Herb. 179: 68. 1956. Cheilanthes candida M. Martens & Galeotti var. copelandii (C. C. Hall) Mickel, Phytologia 41: 433. 1979. Chrysochosma candida (M. Martens & Galeotti) Ku¨mmerle var. copelandii (C. C. Hall) Pic. Serm., Webbia 43: 307. 1989. Type. Mexico. Nuevo Leo´n: Sierra Madre near Monterrey, Pringle 2038 (isotype NY!).

Similar to N. candida, differing in: blade apices abruptly contracted to a fairly well defined terminal pinna, and the basal pinnae with a single basiscopically exaggerated pinnule; 2n⫽60 (NL, Tam).

Distribution. Rocky slopes and cliffs, apparently confined to limestone; 300–1500 m. USA (Tex); Mexico. Selected Specimens Examined. Coah (Johnston 7205, US; Sa´nchez 784, MEXU). Hgo (Hinton 21139, UC). NL (Dorr 2586, NY; Pennell 16949, MEXU, NY; Windham 468, NY). Tam (Bartlett 10586, MICH; King 4528, NY; Palmer 191, NY; Stanford et al., 2137, NY). Unverified, Doubtful, or Mistaken Reports. SLP (Purpus 5484, F, US, 4876, C, 4877, F, UC, US, all cited by Hall, 1950, but not verified).

Notholaena candida closely resembles Cheilanthes farinosa but lacks a false indusium. Farina in N. candida is usually white but occasionally pale yellow. There is a question of the distinctness of Notholaena copelandii, which see for discussion.

Notholaena copelandii often has been treated as a variety of N. candida. The two taxa show differences in chemical composition of the farinose indument (Wollenweber, 1984), and isozyme data indicate that Notholaena copelandii and N. candida are distinct (Windham, in FNA Ed. Comm., 1993); however, few intermediate specimens occur in Coahuila. Several specimens in other parts of the country (Chiapas, Oaxaca, Guerrero, Nayarit, Zacatecas) resemble Notholaena copelandii in having a single exaggerated basiscopic pinnule and somewhat abruptly narrowed blade apices, suggesting that there is some variability in these characters. However, the basiscopic basal pinnules in these specimens tend to be larger than those of N. copelandii and the blade apices are not as abruptly narrowed. Probably these specimens from southern and western states are aberrant specimens of N. candida. Notholaena copelandii differs from N. jacalensis in its wider blades: (3–)4–7.5 cm wide (vs. 2.5–4 cm in N. jacalensis).

10. NOTHOLAENA COPELANDII

11. NOTHOLAENA GALEOTTII

Distribution. Dry rock scrubby hillsides; 450–1700 m. Mexico; Guat, Hond, Salv, Nic. Selected Specimens Examined. Chih (Palmer 82, NY). Chis (Breedlove 21577, NY). Col (McVaugh 15539, MICH). Dgo (Ibana Garcı´a 650, US). Gro (Hinton 9628, NY). Gto (Ventura & Lo´pez 8971, IEB). Hgo (Fisher 37107, US). Jal (Pringle 11786, B, F, GH, K, P, US). Me´x (Hinton 7395, NY). Mor (Lyonnet 2676, US). Nay (Feddema 792, NY). Oax (Mickel 5000, NY). Pue (Purpus 3145, F). Qro (Breedlove 66612, CAS). SLP (Sohns 1247, US). Sin (Palmer 1575, NY). Son (Gentry 11370, US). Ver (Purpus 2171, NY). Zac (Pray 3138, LAM, NY). Unverified, Doubtful, or Mistaken Reports. Ags (De la Cerda & Garcı´a 276, HUAA, cited by Siqueiros-Delgado & Gonza´lez-Adame, 2004 [as Cheilanthes candida], but not verified). BCS (reported by Wiggins, 1980, but not verified).

Notholaena copelandii C. C. Hall, Amer. Fern J. 40: 181, pl. 16. 1950. Fig. 203E.

Notholaena galeottii Fe´e, Me´m. Foug. 5: 159. 1852. Figs. 201A–E.

418

NOTHOLAENA

Cheilanthes galeottii (Fe´e) Mickel & Beitel, Mem. New York Bot. Gard. 46: 113. 1988. Chrysochosma galeottii (Fe´e) Pic. Serm., Webbia 43: 308. 1989. Type. Mexico. Oaxaca: Caputalpan [Capulalpan], Galeotti 6565 (P?, not found; isotypes BM!, BR!–2 sheets, K!). Notholaena arsenii Christ, Notul. Syst. (Paris) 1: 232. 1910. Lectotype (chosen by R. Tryon, 1956: 54). Mexico. Puebla: Tetele, Arse`ne 2046 (P). Notholaena hyalina Maxon, Amer. Fern J. 5: 4. 1915. Type. Mexico. San Luis Potosı´: San Jose´ Pass, Pringle 3297 (US!; isotypes GH!, NY!).

Rhizomes compact, horizontal, ca. 2 mm diam.; rhizome scales linear-lanceolate, 2–3 mm long, black, sclerotic, lustrous, margins ciliate-denticulate; fronds 15–54 cm long but generally only ca. 15–20 cm long, clumped; stipes ca. 1⁄4 the frond length, black, terete, clothed at bases with dark brown, ciliate-denticulate scales 2–3 mm long, becoming light brown at scale bases and distally on rachises totally light brown, stipes and rachises also with a tomentum of much reduced, dissected, appressed scales, some appearing like stellate hairs; blades narrowly elliptic, 3.5–8 cm wide, only slightly narrowed proximally, 2-pinnate to 2-pinnatepinnatifid, subcoriaceous; pinnae 16–30 pairs, equilateral; adaxial surfaces with sparse white to orange simple hairs 0.5–1 mm long; abaxial surfaces densely clothed with rusty, long-ciliate scales 1 mm long, covering the white farina; blade margins somewhat curved; spores tan to black; n⫽2n⫽90, apogamous (Oax).

12. NOTHOLAENA GRAYI Notholaena grayi Davenp., Bull. Torrey Bot. Club 7: 50, pl. 4. 1880. Figs. 206N–Q. Cheilanthes grayi (Davenp.) Domin, Biblioth. Bot. 20(85): 133. 1913. Chrysochosma grayi (Davenp.) Pic. Serm., Webbia 43: 308. 1989. Type. U.S.A. Arizona: mts of SE Arizona, Feb-Mar 1880, Courtis s.n. (GH, photo MO). Notholaena hypoleuca Goodd., Muhlenbergia 8: 94. 1912, hom. illeg., non Kunze, 1834. Type. U.S.A. Arizona: Mule Mts., Goodding 1004 (US!, photo GH). Notholaena grayi Davenp. subsp. sonorensis Windham, Contr. Univ. Michigan Herb. 19: 36. 1993. Type. Mexico. Sonora: La Mina Verde, 31 km W of Cumpas, Wiggins 7400 (US!; isotypes ARIZ, DS, MICH, UC).

Rhizomes compact, horizontal, 1.5–2.5 mm diam.; rhizome scales linear-lanceolate, 2–4 mm long, concolorous, sclerotic, dark reddish brown (to black with age), apices pale, flexuous, capillary-tipped, margins with pale weak hairs in the proximal two-thirds of the scale and short, dark, stiff teeth distally; fronds (8–)12–23(–35) cm long, clumped; stipes 1/5–1⁄3(–1⁄2) the frond length, light brown to dark castaneous, terete, farinose-glandular and with sparse, linear-lanceolate, reddish brown scales 2–3 mm long, these with weak pale hairs and occasional dark, short-stiff teeth distally; blades narrowly oblong to lanceolate, 1.5–2.5 cm wide, bipinnate to bipinnate-pinnatifid; pinnae (6–)8–10 pairs, sessile, the segments nearly equilateral, narrowly to broadly deltate, the basiscopic pinnules slightly longer than the acroscopic; pinnules 1–3 pairs per pinna, oblong, obtuse, sessile; segments not articulate, the color of the costae passing into the bases of the segments; adaxial blade surfaces sparsely farinose; abaxial surfaces densely white-farinose with scattered, orange-tan, linearlanceolate, entire scales 1–1.5 mm long; laminar margins somewhat recurved; spores brown; 2n⫽60, n⫽2n⫽90 (USA).

Distribution. Dry, rocky slopes in scrub, oak, or pine-oak woods; 1050–2100 m. Mexico; Guat. Selected Specimens Examined. Chis (Breedlove 41207, NY). DF (Schaffner 84, NY). Gro (Martı´nez S. 1215, NY). Gto (Ventura & Lo´pez 9004, IEB). Hgo (Chase 7412, NY). Jal (Jones 533, NY). Me´x (TejeroDı´ez 2170a, IZTA). Mich (Martı´nez 1817, MEXU). Mor (Copeland herb. 98a, US). Oax (Mickel 6315, NY). Pue (Rose 10126, US). Qro (Gonza´lez 171, IEB). SLP (Pringle 3297, NY). Tlax (Acosta P. 2168, XAL). Unverified, Doubtful, or Mistaken Reports. Reports by Mickel & Beitel, 1988, of occurrence in BC, Chih, Dgo, NL, and Ver have not been verified.

Notholaena galeottii is distinct in having a dense indument of laminar scales overlying farina. The species can be distinguished from N. aschenborniana by having stiff simple hairs adaxially, and rachises clothed primarily with scales (vs. spreading, stellate hairs adaxially and rachises clothed primarily with stellate hairs).

Distribution. Rocky slopes and cliffs, on a variety of substrates, including both granite and limestone; 350–1300 m. USA (Ariz, N Mex, Tex); Mexico. Selected Specimens Examined. Chih (Correll & Johnston 21759, MSC, NY; Pringle 463, NY) Coah (Palmer 1388, NY). Dgo (Gonza´lez 1276, CHAPA, ENCB). Hgo (Pray 3106, LAM). Jal (Pringle 5373, GH). NL (Herna´ndez I. s.n., 25 Aug 1976, ENCB). Sin (White 4046, ARIZ). Son (Wiggins 7400, US). Zac (Pray 3137, LAM).

NOTHOLAENA JACALENSIS

Notholaena grayi resembles Argyrochosma pallens but is distinguished by the orange blade scales, sessile pinnae, light brown stipes, and weakly ciliate scales of rhizomes and stipes. It is also close to N. aliena (which see) and superficially to N. schaffneri, but the latter has ciliate-denticulate rhizome scales and bristlelike laminar scales rather than flat scales, and darker stipe color. Notholaena grayi includes two cytotypes, which were treated as subspecies by Windham (1993). Sexually reproducing diploid populations (N. grayi subsp. sonorensis) occur in southern Arizona and western Mexico. The two differ only in chromosome number and the associated spore size and number of spores per sporangium and are not formally recognized here. In the diploid, most sporangia contain 32 spores, and the spores generally measure less than 55 ␮m. In the apogamous triploid, most sporangia contain 16 spores, and the spores generally measure more than 55 ␮m.

13. NOTHOLAENA GREGGII Notholaena greggii (Mett. ex Kuhn) Maxon, Contr. U.S. Natl. Herb. 17: 606. 1916. Figs. 202G, H, J. Pellaea greggii Mett. ex Kuhn, Linnaea 36: 86. 1869; Allosorus greggii (Mett. ex Kuhn) Kuntze, Revis. Gen. Pl. 2: 806. 1891. Cheilanthes greggii (Mett. ex Kuhn) Mickel, Phytologia 41: 434. 1979. Chrysochosma greggii (Mett. ex Kuhn), Pic. Serm., Webbia 43: 308. 1989. Type. Mexico. Durango: Gregg 467 (B, frag. US!, photo GH; isotypes G, GH, MO). Notholaena pringlei Davenp., Bull. Torrey Bot. Club 13: 132, t. 58. 1886. Cheilanthes davenportii Domin, Biblioth. Bot. 20(85): 133. 1910, non Cheilanthes pringlei Davenp. Lectotype (chosen by Maxon, Contr. U.S. Natl. Herb. 17: 607. 1916). Mexico. Chihuahua: Santa Eulalia Mts, Pringle 441 (GH!, photo MO; isotypes GH!, NY!, P!, US!).

Rhizomes compact, horizontal, 2–3 mm diam.; rhizome scales linear-lanceolate, bicolorous, ca. 3 mm long, each with black sclerotic center and brown, erose-denticulate margins, attenuate to a pale, weak, flexuous, hair-like apex; fronds 6–20 cm long, clumped; stipes 1⁄3–3/5 the frond length, brown, slightly grooved, with a few lanceolate-ovate scales and scattered farina glands; blades narrowly lanceolate, 2–3-pinnate, 1–2.5 cm wide, grayish green, subcoriaceous; pinnae slightly ascending, 6–12 pairs, with segments ovate to oblong, somewhat bead-like, slightly longer on basiscopic side, segments deltate-oblong to suborbicular; adaxial surfaces more than half covered with white farina; abaxial surfaces densely covered with white farina; laminar margins strongly recurved; sporangia 64-spored; spores tan to black; 2n⫽60 (USA).

419

Distribution. Calcareous slopes and ledges, usually on limestone or gypsum; 1000–2840 m. USA (Tex); Mexico. Selected Specimens Examined. Chih (Chiang et al. 9758, NY; Knobloch 221, MSC; Pringle 1567, MEXU, NY). Coah (Henrickson 7942, ASC; Hinton 16562, GH, NY, 16870, ARIZ; Palmer 1382, NY; Wynd 747, MSC, NY). Dgo (Correll & Johnston 20023, NY; Orcutt 3476, DS). Unverified, Doubtful, or Mistaken Reports. NL (Aguirre s.n., ENCB, cited by Arreguı´n Sa´nchez & Aguirre-Clavera´n, 1985, but not verified).

Notholaena greggii is most closely related to N. bryopoda Maxon, an endemic gypsophile in Coahuila and Nuevo Leo´n. The latter can be distinguished by subconcolorous, bright brown rhizome scales, linear stipe scales (vs. lance-ovate), and a more easterly distribution.

14. NOTHOLAENA JACALENSIS Notholaena jacalensis Pray, Amer. Fern J. 57: 101. 1967. Type. Mexico. Hidalgo: 1 mi S of Jacala on Hwy 85, Pray 3095 (US!; isotypes GH!, NY!, UC!). Figs. 203F, G. Rhizomes compact, horizontal, 1–2 mm diam.; rhizome scales linear, ca. 2 mm long, bicolorous, each with black center and brown margins, lustrous, entire; fronds 12–20 cm long, clumped; stipes 1⁄2 the frond length, atropurpureous to black, terete to flattened adaxially, at base with sparse lance-deltate, weakly bicolorous, brown scales, farina absent; blades lanceolate, pinnatepinnatifid, 2.2–4 cm wide, proximal pinna pairs widely spread, apices gradually narrowed distally; pinnae 5–9 divided pairs, equilateral above the basal pinna pair, mostly 5–6 mm wide, basal pinna pair with one basiscopically exaggerated pinnatifid pinnule, segments constricted at bases, 2–3.5 mm long, entire, rachises shallowly grooved; adaxial surfaces glabrous; abaxial surfaces with pale yellow or white farina; laminar margins curved; sporangia 64-spored; spores tan or black.

420

NOTHOLAENA

Distribution. Gypsum slopes, rocky roadsides; 1100–1450 m. Mexico. Specimens Examined. Hgo (Fisher s.n., 12 Aug 1937, NY, 3 Aug 1946, CAS; Medrano et al. 9503, MEXU). SLP (Chiang et al. 8178C, LL; Purpus 5484, NY, US; Zamudio 3796, NY).

Notholaena jacalensis is similar to N. copelandii in having a single basiscopic pinnule per basal pinna, but it has narrower blades, gradually narrowed blade apices, widely spaced proximal pinna pairs, pale yellow farina, and entire segments.

15. NOTHOLAENA LEMMONII Notholaena lemmonii D. C. Eaton, Bull. Torrey Bot. Club 7: 63. 1880. Cheilanthes lemmonii (D. C. Eaton) Domin, Biblioth. Bot. 20(85): 133. 1913. Chrysochosma lemmonii (D. C. Eaton) Pic. Serm., Webbia 43: 308. 1989. Type. U.S.A. Arizona: Sta. Catarina Mts, near Ft. Lowell and Tucson, 8 Apr 1880, Lemmon s.n. (YU!; isotypes GH!–3 sheets, NY!).

Distribution. Epipetric on rocky slopes in tropical deciduous forests; 75–1450 m. Mexico. Notholaenea lemmonii is distinct from N. grayi by its lack of scales on the fronds. It is similar to Notholaena meridionalis except that it has grooved stipes and rachises. Key to the Mexican Varieties of Notholaena lemmonii 1. Mature stipes and rachises dark castaneous to black; rhizome scales 2 mm long or more; BCS to Jal. ... 15a. var. lemmonii. 1. Mature stipes and rachises light brown to brown; rhizome scales ca. 1 mm long; Gro, Me´x, Mich, Oax, Pue. ................................................................................. 15b. var. australis.

15a. Notholaena lemmonii var. lemmonii

Figs. 200J, K.

Differing from var. australis by its frond axes dark castaneous to black, larger rhizome scales, and more northern distribution; 2n⫽60 (USA).

Rhizomes compact, horizontal, 1.5–2.5 mm diam.; rhizome scales linear-lanceolate, bicolorous, dark brown to black with very narrow tan margins, weakly ciliolate, lustrous, apices flexuous, hair-like; fronds 16–35 cm long, clumped; stipes ca. 1⁄3 the frond length, shallowly grooved, lustrous, reddish brown to black, with a few cordate lance-ovate scales proximally; blades linear to narrowly oblong, 1.3–5 cm wide; rachises grooved, pinnate-pinnatifid to bipinnate, subcoriaceous; pinnae 10–17 pairs, deltate to lanceolate, short-stalked (1 mm), proximal ones somewhat reduced; segments linear to oblong, broadly adnate; adaxial surfaces glabrous; abaxial surfaces densely white- to pale yellow-farinose; laminar margins curved; sporangia 64-spored; spores tan to dark brown. 2n⫽60 (USA).

Distribution. Epipetric on granitic hillsides, in rock crevices, on steep seepy slopes, with leguminous shrubs in thorn forests, and in tropical deciduous forests; 75–1450 m. USA; Mexico. Selected Specimens Examined. BCS (Brandegee 659, NY; Jones 24564, NY; Pray 1779, NY; Wiggins 5465, MICH, NY). Chih (Knobloch 1340, MSC; Palmer 453, NY, US). Jal (Palmer 701, NY p.p.). Sin (Keil & Canne 8766, NY). Son (Palmer 266, NY; Wiggins & Rollins 303, NY).

NOTHOLAENA MERIDIONALIS

421

15b. Notholaena lemmonii var. australis R. M. Tryon, Contr. Gray Herb. 179: 62. 1956. Figs. 200L–P. Cheilanthes lemmonii (D. C. Eaton) Domin var. australis (R. M. Tryon) Mickel & Beitel, Mem. New York. Bot. Gard. 46: 116. 1988. Chrysochosma lemmonii (D. C. Eaton) Pic. Serm. var. australis (R. M. Tryon) Pic. Serm., Webbia 43: 308. 1989. Type. Mexico. Puebla: Coxcatla´n, Purpus 4178 (US!; isotype UC!).

Differing from var. lemmonii by its light brown to brown frond axes, smaller rhizome scales, and more southern distribution.

Distribution. Submontane oak woods; ca. 800 m. Mexico. Specimens Examined. NL (Rodrı´guez L. 420, TEX, 433, IEB, NY).

This species is distinct by its small size, bipinnate (to tripinnate proximally) deltate blades, dense white farina abaxially (to yellowish with age), and relatively few pinnae.

17. NOTHOLAENA MERIDIONALIS

Distribution. Epipetric on open, rocky slopes; 900 m. Mexico. Specimens Examined. Gro (Campos R. 932, MEXU, NY). Me´x (Tejero-Dı´ez 3064, IZTA). Mich (McVaugh 17984, MICH). Oax (Hitchcock s.n., 14 Aug 1910, US; Mickel 844, ISC, US; Rose 10093, US). Pue (Guı´zar 1373, MEXU; Smith et al. 3718, US).

The rhizome scales have pale, weak, easily detached hairs on the margins. The typical variety has only slightly longer rhizome scales and black to dark castaneous stipes.

16. NOTHOLAENA LEONINA Notholaena leonina Maxon, Contr. U.S. Natl. Herb. 16: 56. 1912. Figs. 200E–H. Chrysochosma leonina (Maxon) Pic. Serm., Webbia 43: 308. 1989. Type. Mexico. Nuevo Leo´n: Monterrey, Palmer 1381 (US!; isotypes GH!–3 sheets, NY!, P!, YU!).

Rhizomes compact, horizontal, 1–2 mm diam.; rhizome scales linear, ca. 3 mm long, bicolorous, the center dark reddish brown to black with age, margins tan, lustrous, entire, but with glandular hairs when young; fronds 5–14 cm long, clumped; stipes 1⁄2 the frond length, atropurpureous, terete (rachises grooved), with scattered to sparse, lance-deltate scales 1.5–2.5 mm long and scattered farina; blades bipinnate, tripinnate at base, narrowly deltate, 2–2.5 cm wide; pinnae 4–7 pairs, equilateral; segments oblong, entire, subcoriaceous; abaxial surfaces with dense farina, white to pale yellow with age; laminar margins curved; spores black.

Notholaena meridionalis Mickel, sp. nov. Type. Mexico. Oaxaca: Tehuantepec, Orcutt 3315 (holotype NY!; isotypes GH!, K, MEXU, MO, US!). Figs. 199A–C. A N. lemmonii stipitibus non sulcatis et squamis rhizomatum concoloribus nigrescentibus distincta. (L., meridionalis, south, alluding to the more southerly distribution than its close relative N. affinis.)

Rhizomes compact, horizontal, 2 mm diam.; rhizome scales linear-lanceolate, ca. 3 mm long, concolorous, dark reddish brown, sclerotic, lustrous, margins short-ciliate-denticulate; fronds 7–23 cm long, clumped; stipes ca. 1/6 the frond length, atropurpureous to black, terete, with narrowly deltate, bicolorous scales (dark reddish brown with pale center) toward the base; blades narrowly oblong, pinnate-pinnatifid, 1.5–3 cm wide; rachises puberulent adaxially with short brown hairs less than 0.1 mm arranged in two rows; pinnae 8–18 pairs, equilateral, shortpetiolate (less than 1 mm), costae dark abaxially, pinnae mostly cut 2⁄3–4/5 to the midvein; adaxial blade surfaces glabrous; abaxial blade surfaces pale yellow-farinose; laminar margins slightly curved; spores tan to black.

422

NOTHOLAENA

Distribution. Dry, rocky slopes; 300–700 m. Mexico; Guat, Salv, Nic, CR.

Distribution. Rocky slopes and cliffs, apparently confined to limestone; 750–2150 m. USA (Ariz, Tex); Mexico.

Specimens Examined. Oax (Matuda 205, MEXU, US; Mickel 4214, NY; Torres C. 385, 479, MEXU). Guat (Maxon & Hay 3417, NY). Salv (Seiler 88, NY). Nic (Stevens 14893, NY). CR (Boyle 6015, NY; Lems 64090803, NY).

Selected Specimens Examined. Chih (Johnston et al. 11389B, MEXU; Pringle 452, MICH, NY, P; Wendt et al. 9889A, MEXU). Coah (Cowan 3804, UC; Johnston & Muller 822, US; Jones 520, MO, US; Pray 3018, NY; Villareal 5201, IEB). NL (Yatskievych 83-102, 83-295, ARIZ).

Notholaena meridionalis resembles N. lemmonii in its narrow, farinose blades, but can be distinguished by its non-sulcate stipes and rachises. Plants of N. meridionalis have been called Notholaena affinis (Tryon, 1956), but a photograph of the type of N. affinis shows that it has more pinnae and is the same as N. angusta (see comments under N. affinis). Thus, plants treated as Notholaena affinis by Stolze (1981), Mickel and Beitel (1988, as Cheilanthes affinis), and Yatskievych (in Davidse et al., 1995) need a name.

18. NOTHOLAENA NEGLECTA Notholaena neglecta Maxon, Contr. U.S. Natl. Herb. 17: 602. 1916. Figs. 202K–N. Cheilanthes neglecta (Maxon) Mickel, Phytologia 41: 435. 1979. Chrysochosma neglecta (Maxon) Pic. Serm., Webbia 43: 308. 1989. Type. Mexico. Coahuila: vicinity of Saltillo, Palmer 324 (US!, photos GH, MICH; isotypes BM, GH,! MO, NY!).

Rhizomes compact, horizontal, ca. 3 mm diam.; rhizome scales linear-lanceolate, bicolorous, 2.5–3.5 mm long, dark brown to black with brown margins, erose-denticulate, long-attenuate, apices twisted; fronds (5–)10–15(–25) cm long, clumped; stipes 1⁄2– 2 ⁄3 the frond length, atropurpureous to black, terete, with a few scales at base like those of rhizomes; rachises similar but shallowly grooved; blades 3-pinnate to 3-pinnate-pinnatifid (to 4-pinnate at base), elongate-pentagonal, 2–6.5 cm wide, herbaceous to subcoriaceous; ultimate segments constricted at bases, narrowly adnate to petiolulate; pinnae 4–8 pairs, with basal pair exaggerated basiscopically; adaxial surfaces sparsely pale yellow-farinose; abaxial surfaces densely pale yellow-farinose; laminar margins strongly curved (covering half of lower surface); spores tan or black; 2n⫽60 (NL, USA), n⫽2n⫽90 (USA).

Notholaena neglecta resembles small specimens of N. candida, but the blades are more dissected and pentagonal and have revolute margins.

19. NOTHOLAENA RIGIDA Notholaena rigida Davenp., Gard. & Forest 4: 519, f. 80. 1891. Figs. 199K–M. Cheilanthes rigida (Davenp.) Domin, Biblioth. Bot. 20(85): 133. 1913. Chrysochosma rigida (Davenp.) Pic. Serm., Webbia 43: 308. 1989. Type. Mexico. Nuevo Leo´n: Sierra de la Silla, Pringle 2599 (GH; isotypes B, BM, F, G, K, MO, NY!, P, US!, YU!).

Rhizomes compact, horizontal, 1.5–2.5 mm diam.; rhizome scales linear-lanceolate, ca. 3 mm long, black, concolorous, sclerotic, lustrous, margins ciliate-denticulate; fronds 30–52 cm long (12–30 cm in Chis), clumped; stipes 1/6–1⁄3 the frond length, castaneous to atropurpureous, terete, bases with scattered to sparse, lanceolate scales 1–1.5 mm long, these denticulate to subentire, short-glandular to white-farinose-glandular; blades narrowly elliptic, bipinnate, 4.5–7.5 cm wide; pinnae (10–)13–16 pairs, equilateral to slightly exaggerated basiscopically, the segments shallowly lobed, chartaceous; adaxial surface glabrescent medially, glandular near margins; abaxial surfaces densely whitefarinose; laminar margins not or only slightly curved; spores tan or black; 2n⫽60 (Tam).

NOTHOLAENA SCHAFFNERI

Distribution. Limestone ledges; 320 m. Mexico. Specimens Examined. Chis (Breedlove 53647, CAS, ENCB). NL (Pringle 2599, NY, 15617, US). Tam (Knobloch 2244, MSC; Palmer 142, NY, UC, US; Van Devender & Burgess s.n., 20 Aug 1978, ARIZ; Yatskievych 83–109, 83–318, ARIZ).

This species has an unusual disjunct distribution. Chiapas material differs only in being slightly smaller (12–30 cm vs. 30–52 cm long), having pinnae equilateral (vs. most slightly inequilateral), and lacking farina (but with glands).

20. NOTHOLAENA ROSEI Notholaena rosei Maxon, Contr. U.S. Natl. Herb. 16: 59. 1912. Figs. 200A–D, 206F–H. Cheilanthes rosei (Maxon) Mickel & Beitel, Mem. New York Bot. Gard. 46: 121. 1988. Chrysochosma rosei (Maxon) Pic. Serm., Webbia 43: 308. 1989. Type. Mexico. Jalisco: Chapala, Rose & Painter 7665 (US!, photo GH, frag. NY!). Notholaena lemmonii (D. C. Eaton) Domin var. straminea Davenp., Gard. & Forest 4: 519. 1891. Type. Mexico. Jalisco: Guadalajara, Pringle 2830 (GH). Cheilanthes ⫻coruscans Mickel & Beitel, Mem. New York Bot. Gard. 46: 110. 1988. Type. Mexico. Oaxaca: Dist. Jamiltepec, 40 km SE of Pinotepa Nacional on Rte 200, Mickel 6896 (NY!).

Rhizomes short-creeping, horizontal, 2–3 mm diam.; rhizome scales linear-lanceolate, 3 mm long, black, concolorous, sclerotic, lustrous, margins often very narrow, tan, weakly ciliate; fronds (8–)18–53 cm long, clumped; stipes 1⁄4–1⁄3 the frond length, stramineous (dark brown at bases), terete to slightly flattened on adaxially; blades narrowly elliptic, (2–)5–12 cm wide, slightly narrowed proximally, bipinnate-pinnatifid to tripinnate, subcoriaceous; rachises grooved, with short glandular hairs adaxially, white-farinose abaxially; pinnae 10–15 pairs, slightly inequilateral; adaxial surfaces with sparse white farina or distinct glands; abaxial surfaces densely white-farinose, occasionally only with dense, minute, glandular hairs (glands clear to white or pale yellow in type of C. ⫻coruscans); laminar margins curved, somewhat thinner and erose; spores tan to black; 2n⫽60 (Oax).

423

Distribution. Roadside and woody stream banks; 800–1050 m. Mexico; Hond, Nic. Specimens Examined. Jal (Palmer 701, GH, US; Pringle 2030, GH; Rzedowski 29352, NY). Oax (Mickel 6240, 6896, NY).

Notholaena rosei is distinct by the stramineous stipes and rachises, grooved rachises, and somewhat differentiated fertile blade margins. Cheilanthes ⫻coruscans was described from a very small plant that appeared to be distinct, but agrees with N. rosei in the pale sulcate rachises, bipinnate-pinnatifid blade dissection, somewhat differentiated margins, poorly developed farina, bicolorous rhizome scales, and the lesser axes darker (reddish brown) than the stramineous rachises.

21. NOTHOLAENA SCHAFFNERI Notholaena schaffneri (E. Fourn.) Underw. ex Davenp., Gard. & Forest 4: 519. 1891. Figs. 201F–H, J. Aleuritopteris schaffneri E. Fourn., Bull. Soc. Bot. France 27: 328. 1880. Cheilanthes schaffneri (E. Fourn.) Domin, Biblioth. Bot. 20(85): 133. 1913. Chrysochosma schaffneri (E. Fourn.) Pic. Serm., Webbia 43: 308. 1989. Cheilanthes potosina Mickel in McVaugh, Fl. Nov.-Gal. 17: 241. 1992, non C. schaffneri (E. Fourn.) Domin, 1913. Type. Mexico. San Luis Potosı´: San Miguelito Mts., Schaffner 2, in 1877 (P!, photos GH!, US; isotypes GH!, NY!). Notholaena nealleyi Seaton ex Coulter, Contr. U.S. Natl. Herb. 1: 61. 1890. Cheilanthes nealleyi (Coulter) Domin, Biblioth. Bot. 20(85): 133. 1913. Notholaena schaffneri (E. Fourn.) Underw. ex Davenp. var. nealleyi (Coulter) Weath., J. Arnold Arbor. 24: 315. 1943. Chrysochosma schaffneri (E. Fourn.) Pic. Serm. var. nealleyi (Coulter) Pic. Serm., Webbia 43: 308. 1989. Type. U.S.A. Texas: Jeff Davis Co., Limpia Canyon, Nealley 560, US!, photo GH [Nealley 894, from Chinati [Chenate] Mts. in Presidio Co. in protologue, perhaps in error]. Notholaena nealleyi Seaton ex Coulter var. mexicana Davenp., Bot. Gaz. 16: 54. 1891. Notholaena schaffneri (E. Fourn.) Underw. ex Davenp. var. mexicana (Davenp.) Davenp., Gard. & Forest 4: 519. 1891. Type. Mexico. Jalisco: near Guadalajara, Pringle 1864, (GH!–4 sheets; isotypes NY!, P!, UC!, US!).

Rhizomes compact, horizontal, ca. 2 mm diam.; rhizome scales linear-lanceolate, 2–3 mm long, concolorous, black, sclerotic, lustrous, margins ciliate-denticulate; fronds 8–26 cm long, clumped; stipes 1⁄8–1⁄6 the frond length, atropurpureous, with linear-lanceolate, castaneous, ciliate-denticulate scales 1.5–3 mm, bristle-like hairs ca. 1 mm long in addition to farina; blades narrowly elliptic, 2–4 cm wide, reduced proximally, bipinnate-

424

NOTHOLAENA

pinnatifid, the segments cut only 1⁄2 way to the costae, coriaceous; pinnae to 24 pairs; adaxial surfaces with sparse glands; abaxial surfaces densely covered with white farina glands and long, bristle-like, castaneous hairs on costae and costules; laminar margins somewhat curved; spores tan to black; 2n⫽60 (Oax), 120 (NL).

Notholaena candida (M. Martens & Galeotti) Hook. var. quinquifidopalmata Hook., Sp. Fil. 5: 111. 1864. Notholaena hookeri D. C. Eaton in Wheeler, Rept. U.S. Geogr. Surv. W. 100th Merid. 6: 308, t. 30. 1879, nom. et stat. nov., hom. illeg., non Lowe, 1856. Chrysochosma hookeri Ku¨mmerle, Magyar Bot. Lapok 13: 40–41. 1914, nom. nov. Cheilanthes hookeri (Ku¨mmerle) Domin, Biblioth. Bot. 20(85): 133. 1915. Notholaena sulphurea (Cav.) J. Sm. var. quinquifidopalmata (Hook.) Farw., Amer. Midl. Naturalist 12: 284. 1931. Type. U.S.A. New Mexico: La Cuesta, Bigelow s.n. (K, photo GH).

Rhizomes compact, horizontal, 1–1.5 mm diam.; rhizome scales lanceolate, 2–3 mm long, bicolorous with broad dark brown to black, sclerotic, lustrous central streak and pale entire margins; fronds 9–23 cm long, clumped; stipes ca. 3⁄4 the frond length, castaneous, terete, lustrous, glabrous except for a few broadly lanceolate scales at bases (2 ⫻ 1 mm); blades broadly pentagonal, 2.5–7 cm wide, pinnatifid above basal pinnae, with the suprabasal pinnae usually shortened, basal pinnae pinnatifid with one basiscopically exaggerated pinnatifid basal pinnule; adaxial surfaces glabrous; abaxial surfaces densely pale yellow-to deep yellow-farinose; laminar margins slightly curved; sporangia 32- or 16-spored; spores tan to black; 2n⫽60 (Oax, USA). Distribution. Dry, rocky slopes, among shaded rocks, wooded ravines, shrub-covered hills, shaded ledges; (300–)1450–2100 m. USA (Tex); Mexico; Jam. Selected Specimens Examined. Coah (Johnston & Muller 205, US). DF (Schaffner 30, in 1875, NY). Dgo (Sa´nchez S. 754, MEXU, US). Gro (Frye & Frye 3145, US). Gto (Galva´n 2664, MEXU). Jal (McVaugh 17317, NY). Mich (Torres & Ramı´rez 13601, IEB). NL (Orcutt 1304, US). Oax (Mickel 4084, NY). Pue (Purpus 3146, NY). SLP (Schaffner 962, GH, NY). Tam (Bartlett 11007, DS, US). Ver (Purpus 6199, NY). Zac (Rose 2662, US). Unverified, Doubtful, or Mistaken Reports. Ags (Siqueiros 4291, HUAA, and one other collection, cited by Siqueiros-Delgado & Gonza´lez-Adame, 2004 [as Cheilanthes potosina], but not verified). Mor (Dixon 61, MEXU, cited by Riba et al., 1996, but not verified).

This species can be distinguished from N. galeottii by having farina on both surfaces of the pinnae and scattered, brown, terete, hair-like scales on the costae and veins (glabrous to hirsute adaxial pinna surfaces, abaxial pinna surfaces with broad, pinnatifid, brown scales in N. galeottii). Notholaena nealleyi appears to be only a diminutive form of N. schaffneri. Notholaena nealleyi is said to differ by the pinnae having only 1–3 pairs of free pinnules per pinna, vs. 4–6 in true N. schaffneri. However, Windham (in FNA Ed. Comm., 1993) treated Notholaena nealleyi as distinct and reported that N. nealleyi may represent a tetraploid hybrid between N. schaffneri and an unknown taxon. More work is needed.

22. NOTHOLAENA STANDLEYI Notholaena standleyi Maxon, Amer. Fern J. 5: 1. 1915. Type. U.S.A. Texas: w Texas to El Paso, New Mexico, Wright 821 (YU; isotypes K!, US!). Figs. 204A–D.

Distribution. Calcareous or noncalcareous cliffs, igneous rocks, sandstone, rock crevices, among large boulders and cliffs, limestone outcrops; 380–2200(–3100) m. Sw USA; Mexico. Selected Specimens Examined. BCN (Moran 18714, ENCB). BCS (Carter 2049, MEXU, US). Chih (Pringle 467, NY). Coah (Palmer 1379, NY). Dgo (Purpus 4632, US). Hgo (Rose 9107, US). NL (Rodrı´guez L. & Sauceda M. 412, ANSM). Oax (Mickel 4490, NY). Pue (Purpus 4027, NY). Sin (Pitelka 286B, UC). SLP (Parry 992, NY). Son (Lumholtz Exped. 361, NY). Tam (Bartlett 10708, US).

The farina varies in color geographically. In Sonora, Chihuahua, and part of Coahuila the farina is very pale yellow, to nearly white in some Chihuahuan specimens. In part of Coahuila and in San Luis Potosı´, the farina is deep yellow, and in Puebla and Oaxaca pale yellow. Windham (in FNA Ed. Comm., 1993) reported that the low number of spores per sporangium in N. standleyi is apparently not associated with apogamy since all plants analyzed were sexual diploids. Siegler and Wollenweber (1983) identified three chemotypes in this species; these correlate with substrate specificity, range, and variations in farina color.

ODONTOSORIA

23. NOTHOLAENA SULPHUREA Notholaena sulphurea (Cav.) J. Sm. in Seeman, Bot. Voy. Herald 1: 233. 1854. Figs. 204E–G. Pteris sulphurea Cav., Descr. Pl. 269. 1801. Cheilanthes sulphurea (Cav.) Mickel & Beitel, Mem. New York Bot. Gard. 46: 123. 1988. Chrysochosma sulphurea (Cav.) Ku¨mmerle, Magyar Bot. Lapok 13: 40, 41. 1914. Type. Mexico. Hidalgo: Chimapan [Zimapan], Ne´e s.n. (MA, seen and verified by C. Chr., Dansk Bot. Ark. 9(3): 23. 1937; isotypes F, G). Notholaena cretacea Liebm., Mexic. Bregn. 216 (reprint 64). 1849. Aleuritopteris cretacea (Liebm.) E. Fourn., Mexic. Pl. 1: 121. 1872. Type. Mexico. Puebla: Tehuaca´n, San Lorenzo, Liebmann s.n. (C!, frags. GH, US!; isotypes K, frag. NY!, P!). For additional synonymy, see R. Tryon (1956).

Rhizomes compact, horizontal, 1–2 mm diam.; rhizome scales linear-lanceolate, 2–3 mm long, bicolorous with black sclerotic central streak and narrow light brown glandular-erose margins (at least at rhizome apices); fronds 9–14(–20 cm) long, clumped; stipes ca. 3⁄4 the frond length, black to atropurpureous, terete, with broadly lanceolate, glandular-ciliate scales at bases (2 ⫻ 1 mm); blades pentagonal, 3.2–7 cm wide, pinnate to pinnatepinnatifid above basal pinna pair, the basal pinna pair pinnate to pinnate-pinnatifid, exaggerated basiscopically, 1–4 pairs of pinnatifid pinnae above basal pinnae, subcoriaceous; adaxial surfaces glabrous or sparsely farinose; abaxial surfaces densely whiteto yellow-farinose; laminar margins slightly curved; spores tan to black; 2n⫽60 (Tam, white form).

425

yellow in Quere´taro, San Luis Potosı´, and Nuevo Leo´n, and deep yellow in Puebla and Oaxaca.

24. NOTHOLAENA WEATHERBIANA Notholaena weatherbiana R. M. Tryon, Contr. Gray Herb. 179: 60. 1956. Type. Mexico. Southwestern Chihuahua: Palmer 215 (MO; isotypes NY!, GH!, US p.p.). Figs. 206A–E. Rhizomes compact, horizontal, ca. 2 mm diam.; rhizome scales linear-lanceolate, dark reddish brown to black, younger ones with white edges, or all white, margins with sparse ephemeral teeth; fronds 6–20 cm long, clumped; stipes 1⁄3–1⁄2 the frond length, castaneous, terete or flattened adaxially, lanose with curly hairs and narrow whitish scales; blades oblong, 1.5–3.3 cm wide, only slightly narrowed proximally, bipinnate to bipinnatepinnatifid, subcoriaceous; pinnae 5–12 or more pairs, equilateral; adaxial surfaces lanose with curly white hairs; abaxial surfaces with scales on rachises and costae, segments with many curly white hairs and white to cream-colored farina; laminar margins only slightly curved; spores tan to black.

Distribution. No habitat or elevation data given. Mexico. Specimens Examined. Chih (Knobloch 535, DS, 551, MSC, UC; Pray 3184, LAM).

Distribution. Dry, rocky slopes; 1400–2460 m. Mexico; Guat, Nic, CR, Pan; Col, Peru, Chile. Selected Specimens Examined. Coah (Palmer 1381, PH). Gto (Pray 1960, LAM). Hgo (Mickel 1636, ISC). Mich (Dı´az B. & Pe´rez C., 24 Jul 1992, NY). NL (Chase 7769, F). Oax (Conzatti & Go´mez 2360, P). Pue (Purpus 3142, NY, US). Qro (Sohmer 9236, MEXU, NY). SLP (Pringle 3398, US). Tam (Stanford et al. 2635, NY).

The distinctly pinnate blades above the basal pinnae distinguish this taxon from N. standleyi, which is pinnatifid above the basal pinnae. See key for differences from N. candida. This species displays variation of farina color across its range: white in Hidalgo, San Luis Potosı´, Tamaulipas, and Puebla, pale

This species is distinct by its abundant curly hairs mixed with farina on the blades and mixed with narrow whitish scales on the stipes.

8 5 . O D O N T O SO R I A Odontosoria Fe´e, Me´m. Foug. 5: 325, t. 27 B, f. 1. 1852. Type: Odontosoria uncinella (Kunze) Fe´e [⬅ Davallia uncinella Kunze]. Terrestrial; rhizomes short-creeping, slender, scaly; fronds large, scandent, narrowly deltate to linear, 1–6 m long; stipes smooth to spiny, lustrous, stramineous to castaneous; blades 3–5 times pinnate, pinnae and subdivisions at right angles to

426

ODONTOSORIA

their subtending axes; segments linear to flabellate, chartaceous to subcoriaceous; sori marginal, in outward facing pouches; indusia attached on 3 sides, with only ca. 3 sporangia per sorus; spores tetrahedral-globose, without perispore; x⫽96 (from x⫽48?). Odontosoria is a tropical genus of 22 species, of which 12 occur in America and only two in Mexico. It is probably most closely related to Sphenomeris, and a member of the lindsaeoid group of genera. The genus is distinct by the sori in small pockets at segment apices and the rhizome indument of scales. References Kramer, K. U. 1972. The lindsaeoid ferns of the Old World–IX. Africa and its islands. Bull. Jard. Bot. Natl. Belgique 42: 305–345; Maxon, W. R. 1913. Studies of tropical American Ferns. No. 4. Contr. U.S. Natl. Herb. 17: 133–179.

Key to the Mexican Species of Odontosoria 1. Ultimate divisions almost capillary, divaricate, 0.3–0.5 mm wide, linear, or slightly clavate at the monosorus apices. ........................................................................ 2. O. schlechtendalii. 1. Ultimate divisions broader, shorter, 0.6–1.4 mm wide, not distinct, cuneately joined in pairs, then again acutely joined, the segments thus in general 2–4 times flabellately parted or cleft, cuneate proximally. ..... 1. O. guatemalensis.

1. ODONTOSORIA GUATEMALENSIS

Specimens Examined. Chis (Purpus 6747, MO, PH, UC, US, 6748, UC, US, 6872, UC, US; Martı´nez S. 19320, MEXU; Matuda 2435, K, MEXU, US; Breedlove 31621, DS, MEXU, NY; Ventura & Lo´pez 1090, ENCB, XAL).

2. ODONTOSORIA SCHLECHTENDALII Odontosoria schlechtendalii (C. Presl) C. Chr., Index Filic. 209. 1905. Figs. 207A–C. Davallia divaricata Schltdl. & Cham., Linnaea 5: 617. 1830, hom. illeg., non Blume, 1828. Davallia schlechtendalii C. Presl, Suppl. Tent. Pterid. 129. 1836. Stenoloma schlechtendalii (C. Presl) Fe´e, Me´m. Foug. 5: 330. 1852. Microlepia schlechtendalii (C. Presl) Mett., Fil. Hort. Bot. Lips. 104. 1856. Odontoloma schlechtendalii (C. Presl) E. Fourn., Mexic. Pl. 1: 32. 1872. Odontosoria divaricata (Schltdl. & Cham.) J. Sm., Hist. Fil. 264. 1875. Lindsayopsis divaricata (Schltdl. & Cham.) Kuhn, Festschrift zu 50 Ja¨hr. Junil. D. Kgl. Realschule zu Berlin 27. 1882. Lindsaea schlechtendalii (C. Presl) Christ, Farnkr. Erde 296. 1897. Type. Mexico. [Veracruz:] Schiede & Deppe [803] (B!–2 sheets; isotype LE!).

Rhizomes short-creeping, 2–5 mm diam.; rhizome scales castaneous, very narrow, 2–3 ⫻ 0.5 mm; fronds 1.7–5 m long, arching, pendent on cliffs or scrambling or prostrate on ground, clumped; stipes ca. 1⁄4–1⁄2 the frond length, castaneous, glabrous; blades ovate-lanceolate to linear-lanceolate, 4–5 times pinnate, skeletonized; segments linear, mostly 3–5 ⫻ 0.3–0.5 mm, ultimate divisions subdichotomously forked, glabrous; sori terminal on the segments, ca. same width as the segments, appearing as obconical pockets at the segment apices.

Odontosoria guatemalensis Christ, Bull. Soc. Bot. Gene`ve 1: 229. 1909. Type. Guatemala. Cuesta Grande, “Hacienda de las Nubes,” Bernoulli & Cario 402 (presumably P, but not found). Figs. 207D–F. Differs from O. schlechtendalii by the characters given in the key. Odontosoria guatemalensis somewhat larger species than O. schlechtendalii, judging from the thickness (0.6–1.4 mm) of the stipes and rachises (mostly 2–4 mm diam.), and the ultimate divisions shorter, broader, and less divaricate.

Distribution. Wet clay roadside banks, clearings, and slumps in wet montane forests; 250–2200 m. Mexico; Guat, Bel, Hond. Selected Specimens Examined. Chis (Calzada et al. 9049, NY; Matuda 223, MEXU, NY, 18571, MEXU). Gro (Hinton 11189, NY). Oax (Maya J. 3262, NY; Mickel 7237, NY). Pue (Lorea 1102, FCME; Marquez R. 715, XAL). Ver (Harriman 14782, NY; Nee & Diggs 24879, XAL). Unverified, Doubtful, or Mistaken Reports. Mich (reported by Mickel & Beitel, 1988, but not verified).

Distribution. Scandent, along roadbanks; 1000–1700 m. Mexico; Guat, Hond.

In the sterile condition this species closely resembles Eriosorus flexuosus in frond size and dissection. The sori are quite different in the two, Odontosoria schlechtendalii having marginal cup-like

OLFERSIA

involucres and Eriosorus flexuosus having the sporangia naked along the segment veins. They can be distinguished even in the juvenile condition by Odontosoria schlechtendalii having the rhizomes clothed with thin scales (rather than hairs), and lacking hairs on the laminae.

8 6 . O L E A N DR A Oleandra Cav., Anales Hist. Nat. 1: 115. 1799. Type: Oleandra neriiformis Cav. Terrestrial, epipetric, or occasionally epiphytic; rhizomes slender, long-creeping or sometimes climbing or erect, dictyostelic, densely scaly, scales peltate, rhizomes bearing long stiff roots; fronds commonly monomorphic, articulate at prominent joint, leaving short stalks (phyllopodia), borne at wide intervals or in loose clusters; blades simple, entire, glabrous to pubescent, hairs often septate, midribs slightly to rather densely scaly; veins free, unbranched or forked near base, parallel, often fusing with cartilaginous margin; sori round, borne irregularly on veins, but mostly near midribs; indusia round-reniform to reniform, attached at deep sinuses; spores bilateral, perispore winged, coarsely ridged, and often spinulose between the ridges; x⫽41, 40? Oleandra is a pantropical genus with about eight species in the neotropics (Lellinger, unpubl.) and about 30 species in the Paleotropics from Africa to Polynesia; however, these numbers were considerably reduced by R. Tryon (1997, 2000), and reflect a much broader species concept. Oleandra is distinctive in its long-creeping, scandent to erect rhizomes, simple blades with stipes articulate above the rhizome (leaving short to often very long phyllopodia), and reniform indusia borne on parallel free veins. Its center of distribution in the neotropics is from Costa Rica to Colombia and Venezuela. The affinities of Oleandra are uncertain, but probably with Arthropteris, a small genus occurring from Africa to Asia and Polynesia, extending to the Juan Fernandez Islands. Historically, Oleandra has been included with genera of the Davalliaceae (e.g., by Moran in Davidse et at., 1995). Others have included the genus in a broadly circumscribed Dryopteridaceae (e.g., A. F. Tryon & R. M. Tryon, 1982; Smith in Steyermark et al., 1995). Still others place it in its own family, Oleandraceae, along with one or two other small genera (e.g., Kramer in Kubitzki, 1990). Its coarsely ridged spores and base chromosome number clearly point to an alliance with the dryopteroid ferns sensu lato. References Maxon, W. R. 1914. The American species of Oleandra. Contr. U.S. Natl. Herb. 17: 392–398; Tryon, R. M. 1997. Systematic notes on Oleandra. Rhodora 99: 335–343; Tryon, R. M. 2000. Systematic notes on the Old World fern genus Oleandra. Rhodora 102: 428–438.

1. OLEANDRA ARTICULATA Oleandra articulata (Sw.) C. Presl, Suppl. Tent. Pterid. 78, t. 2, f. 12. 1836. Figs. 208A–C.

427

Aspidium articulatum Sw., J. Bot. (Schrader) 1800(2): 30. 1801 [1802]. Lectotype (chosen by Maxon, 1914: 394). Plumier, Traite´ Foug. Ame´r., pl. 136. 1705, illustrating a plant from Martinique. For discussion, see Joncheere, Taxon 18: 538–541. 1969. Aspidium nodosum Willd., Sp. Pl., ed. 4, 5(1): 211. 1810, hom. illeg. superfl. Oleandra nodosa (Willd.) C. Presl, Suppl. Tent. Pterid. 78. 1836. Type. Based on same plate as Aspidium articulatum Sw. Name incorrectly applied to this species by Morton (Amer. Fern J. 58: 105–107. 1968).

Rhizomes long-creeping to scandent, brownish, not pruinose; rhizome scales orange- to red-brown, persistent, 4–6 mm long, spreading (giving the rhizomes a bristly appearance), linearlanceolate, attenuate, margin minutely glandular-ciliate; fronds distant, borne several cm apart; phyllopodia 5–10(–30) mm long, lacking scales; stipes 5–22 cm long; blades elliptic, 15–35 ⫻ 3–5 cm, glabrous, lustrous, firm-herbaceous, bases broadly to narrowly cuneate, apices acuminate to caudate; sori in several, irregular rows, often rather close to the midrib to about halfway to margin; indusia 0.5–1 mm diam., glabrous; 2n⫽82 (Jam, PR, Trin).

Distribution. Epiphytic on tree trunks, logs, or epipetric in crevices of sandstone bluffs, seasonal evergreen forests; 350– 900 m. Mexico; Guat, Bel, Hond, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Guy, Sur, Ec, Peru, Braz, Bol. Specimens Examined. Chis (Breedlove 23583, DS, NY). Ver (Wendt et al. 3846, CHAPA, MEXU, NY).

Oleandra lehmannii Maxon (which includes O. guatemalensis Maxon) occurs as far north as Guatemala, and should be looked for in Chiapas. It can be distinguished by its appressed, bicolorous rhizome scales and sori in one (rarely two) rows on either side and near the midrib. Oleandra bradei Christ of Costa Rica, which has creeping rhizomes with spreading scales and glabrous blades, has deciduous rhizome scales and pruinose rhizomes.

8 7 . O L F E R SI A Olfersia Raddi, Opusc. Sci. 3: 283. 1819. Type: Olfersia corcovadensis Raddi [⫽ Olfersia cervina (L.) Kunze].

428

ONOCLEOPSIS

Hemiepiphytic or terrestrial; rhizomes long-creeping, densely scaly; fronds holodimorphic; stipes scaly; sterile fronds 1-pinnate, terminal pinna conform, pinna pairs 4–12, ascending, coriaceous, axes grooved adaxially; veins parallel, oblique to costae, simple or usually forking at their base, free except for a marginal commissural vein on each pinna; fertile fronds much contracted, bipinnate, pinnules broadly adnate, sporangia in masses and essentially covering both surfaces of pinnules; indusia absent; spores bilateral with thick, folded perispore; x⫽41. Olfersia comprises two species, the widespread O. cervina and O. alata C. Sa´nchez & Garcı´a Caluff, endemic to eastern Cuba (Sa´nchez V. et al. 1991). The genus is closely related to, and often placed in, Polybotrya (which see) but differs from it in its conform terminal pinna, parallel veins uniting at the margin, and sporangia borne on both upper and lower surfaces of the fertile frond. Placement near Polybotrya, Cyclodium, and Maxonia, and within the Dryopteridaceae s.l. is also suggested by recent phylogenetic analyses (Cranfill, unpubl. data). References Moran, R. C. 1986. The neotropical fern genus Olfersia. Amer. Fern J. 76: 161– 178; Sa´nchez V., C., M. Garcı´a C. & C. Zavaro P. 1991. Nueva especie cubana del ge´nero Olfersia (Polypodiaceae–Dryopteridoideae). Fontqueria 31: 229–233.

1. OLFERSIA CERVINA Olfersia cervina (L.) Kunze, Flora 7: 312. 1824. Figs. 209A–C. Osmunda cervina L., Sp. Pl. 2: 1065. 1753. Acrostichum cervinum (L.) Sw., Syn. Fil. 14. 1806. Polybotrya cervina (L.) Kaulf., Enum. Filic. 55. 1824. Lectotype (chosen by Proctor, Fl. L. Antill. 2: 223. 1977). Plumier, Traite´ Foug. Ame´r., pl. 154. 1705, based on material from Morne Rouge, Martinique.

Rhizomes long-creeping, 1–2 cm diam.; rhizome scales golden to orange-brown, linear-lanceolate, 10–20 ⫻ 1 mm, apices twisted, hair-like, margins entire; fronds clumped; stipes stramineous, shorter than blades, scaly at bases; sterile blades narrowly oblong to ovate-lanceolate, 30–85 ⫻ 20–35 cm, glabrous except for rare, catenate hairs along costae abaxially; pinnae lanceolate to lanceolate-oblong, 15–25 ⫻ 3–9 cm, broadly and unequally cuneate at bases, often excavate basiscopically, apices longacuminate, entire; fertile blades skeletonized, pinnules 0.8–1 cm ⫻ 2–3 mm, entire; 2n⫽82 (Jam).

Distribution. Terrestrial, on fallen logs and occasional tree trunks in wet lowland and montane forests; 200–1050 m. Mexico; Guat, Bel, Hond, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol. Selected Specimens Examined. Chis (Martı´nez S. 19863, 22690, MEXU). Oax (Martı´nez C. 527, ENCB, XAL, NY; Mickel 4006, ENCB, NY, UC, 6384, NY, UC, 6399, ENCB, NY). Ver (Liebmann s.n. [Pl. Mex. 2691], US; Ventura A. 12428, CHAPA, ENCB, 17030, IEB; Wendt et al. 3846a, CHAPA, NY, UC).

With its distinctive dimorphic fronds, parallel veins uniting in a marginal commissure, conform terminal pinna, and thicktextured, entire lateral pinnae, O. cervina is unlikely to be confused with any other Mexican species.

8 8 . O N O C L E O P SI S Onocleopsis Ballard, Amer. Fern J. 35: 1. 1945. Type: Onocleopsis hintonii Ballard. Terrestrial; rhizomes erect, stout, often stoloniferous from bases; rhizome scales concolorous, brown, lustrous; fronds clumped, dimorphic; stipes light brown to stramineous, with scattered concolorous scales; sterile blades pinnate, reduced at blade bases, apices pinnatifid; rachises similar to stipes; sterile pinnae lanceolate, sessile, bases truncate or subcordate, apices acuminate, margins broadly serrate; blades glabrous adaxially, short-pilose on veins and costae abaxially, membranaceous to chartaceous; fertile blades 2–3-pinnate, with laminar tissue nearly absent, fertile pinnules rolled into 3–4 globular divisions, brown at maturity; veins on sterile pinnae netted and without included veinlets, on fertile pinnae free; sori enclosed by inrolled pinnae, ultimate segments with 2 sori; indusia essentially absent or present as thin, hyaline scale for each sorus; spores bilateral; x⫽?. Onocleopsis is a monotypic genus, occurring from central Mexico to Guatemala at high elevations in or adjacent to mountain streams. It is closely allied to Matteuccia and Onoclea of north temperate regions. The genus is distinct by the large, 1pinnate sterile fronds with netted veins and 2–3-pinnate fertile fronds.

OPHIOGLOSSUM References Ballard, F. 1945. A new fern genus from Mexico and Guatemala. Amer. Fern J. 35: 1–3; Ballard, F. 1948. Further notes on Onocleopsis. Amer. Fern J. 38: 125–132; Gastony, G. J. & M. C. Ungerer. 1997. Molecular systematics and a revised taxonomy of the onocleoid ferns (Dryopteridaceae: Onocleeae). Amer. J. Bot. 84: 840–849; Kato, M. & N. Sahashi. 1980. Affinities in the onocleoid ferns. Acta Phytotax. Geobot. 31: 127–138; Lloyd, R. M. 1971. Systematics of the onocleoid ferns. Univ. Calif. Publ. Bot. 61: 1–86.

1. ONOCLEOPSIS HINTONII Onocleopsis hintonii F. Ballard, Amer. Fern J. 35: 1. 1945. Type. Mexico. Me´xico: Dist. of Temascaltepec, wet barranca at Los Hornos, Hinton 3297 (K!). Figs. 210A–D. Rhizome scales to 15 ⫻ 4 mm, ovate to lanceolate, longattenuate, entire; sterile fronds 50–200 cm long, blades 25–35 cm wide; fertile fronds 56–120 cm long, blades 10–20 cm wide; stipes 20–30 cm long; pinna pairs 17–28; sterile pinnae to 50 ⫻ 3 cm; fertile pinnae to 20 ⫻ 3.5 cm, ultimate fertile segments 1.5–2.5 mm long.

429

26115, CAS, IEB, NY). Me´x (Hinton 7228, UC, US, 16314, NY, UC, US). Mich (Santos Martı´nez 2073, IEB, MEXU). Mor (Pringle s.n., 19 Sep 1896, US). Pue (Wolfgang 1215, MEXU). Oax (Anderson & Laskowski 4166, ENCB, MICH, US; Mickel 4162, ENCB, NY, UC).

8 9 . O P H I O GL O SSU M Ophioglossum L., Sp. Pl. 2: 1062. 1753. Lectotype (chosen by J. Smith, Hist. Fil. 367. 1875): Ophioglossum vulgatum L. Cheiroglossa C. Presl, Suppl. Tent. Pterid. 56. 1845. Type: Cheiroglossa palmata (L.) C. Presl [⬅ Ophioglossum palmatum L.].

Terrestrial (epiphytic in O. palmatum); stems short, upright, fleshy; roots fleshy, commonly giving rise to new plants from root buds; fronds generally small, simple, entire, 1–several per plant, glabrous, composed of (1) a sterile blade which is simple, entire, with intricate reticulate venation, without a true midvein (the sterile blade palmately lobed in O. palmatum), (2) an erect fertile spike arising from the base of the sterile blades, with two vertical rows of large sporangia sunken into the axis tissue and opening by a horizontal slit (O. palmatum with several fertile spikes at the blade bases and distal part of petioles), the fertile spikes terminated by an apiculum to 1.5 mm long, and (3) a common stalk (petiole); spores tetrahedral-globose, yellowish; gametophytes subterranean, whitish or brownish, mycorrhizal, cylindrical, simple to branched; x⫽30. Ophioglossum is a genus of mostly small, terrestrial plants of disturbed grassy habitats, so inconspicuous that they can elude the casual eye. This cosmopolitan genus is thought to comprise 20–25 species. There are few taxonomic characters, and the circumscription of species is difficult. Consequently, findings of great disjuncts are often interpreted as distinct species. Eight species are known in Mexico. References

Distribution. Terrestrial, in soil or sand in water along margins of streams, cloud forests, pine-oak and pine-oak-fir forests; 1500–2450 m. Mexico; Guat. Selected Specimens Examined. Gro (Lorea 2640, FCME, IEB, MEXU). Jal (Cochrane 10666, IBUG, IEB, MEXU, UC, WIS; McVaugh

Clausen, R. T. 1938. A monograph of the Ophioglossaceae. Mem. Torrey Bot. Club 19: 1–177; Wagner, W. H., Jr., C. M. Allen & G. P. Landry. 1984. Ophioglossum ellipticum Hook. & Grev. in Louisiana and the taxonomy of O. nudicaule L. f. Castanea 49: 99–110; Zech, J. C., P. R. Manning & W. H. Wagner Jr. 1998. A new adder’s tongue (Ophioglossum: Ophioglossaceae) for North America. Sida 18: 307–313.

Key to the Mexican Species of Ophioglossum 1. Plants epiphytic; sterile blades in mature plants palmately lobed, pendent; several fertile spikes produced adaxially on a single blade; roots dichotomously branched. ......................................................................................................................................................... 5. O. palmatum. 1. Plants terrestrial; sterile blades in mature plants undivided, erect; one fertile spike per blade; roots unbranched. 2. Stems conspicuously globose-bulbous (3–11 mm diam.); sterile blades 0.8–4 ⫻ 0.8–2 cm with cordate bases. ................ 2. O. crotalophoroides. 2. Stems cylindrical to subglobose (2–5 mm diam.); sterile blades 0.5–15 ⫻ 0.2–5 cm, if less than 4 cm long, bases cuneate to subtruncate. 3. Principal veins forming large primary areoles in which are included secondary areoles; sterile blades usually strongly apiculate or acuminate at tips, rarely rounded. 4. Ribbon-like old leaf bases not persistent. .................................................................................................................................. 4. O. nudicaule. 4. Ribbon-like old leaf bases persistent. 5. Persistent leaf bases 0–4; roots blackish, 0.7–1.1 mm diam.; blades mostly elliptic, (0.5–)2–3.5 cm wide; roots about 15 (6–24) per plant. ............................................................................................................................................................. 3. O. engelmannii. 5. Persistent leaf bases 1–6(–10); roots tan, 0.4–0.7 mm diam.; blades mostly linear-lanceolate, 1–1.5 cm wide; roots about 25 (15–30) per plant. ................................................................................................................................................ 6. O. polyphyllum.

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OPHIOGLOSSUM 3. Principal veins forming areoles with included free veinlets but not smaller secondary areoles; sterile blades mostly rounded to acute at tips. 6. Basal leaf sheaths persistent, leathery; plants mostly 8–15 cm tall, not including fertile spikes. 7. Blades ovate to nearly reniform, cordate or truncate at base; plants mainly of open disturbed areas. ................... 7. O. reticulatum. 7. Blades mostly ovate-lanceolate to elliptic, broadest and abruptly tapering at bases; plants mainly of shaded forest floor. .......................................................................................................................................................................................... 8. O. vulgatum. 6. Basal leaf sheaths absent, if present, delicately membranous; plants mostly 2–8 cm tall, not including fertile spikes. 8. Leaf blades more or less upright and folded longitudinally (conduplicate); blade outline linear-lanceolate attenuate at tips; fertile spikes 1–2.5 times the sterile blade length. .............................................................................................. 1. O. californicum. 8. Leaf blades flat, not folded longitudinally; blade outline lanceolate to ovate-lanceolate; fertile spikes 2–6 times the sterile blade length. ................................................................................................................................................................. 4. O. nudicaule.

1. OPHIOGLOSSUM CALIFORNICUM

2. OPHIOGLOSSUM CROTALOPHOROIDES

Ophioglossum californicum Prantl, Ber. Deutsch. Bot. Ges. 1: 355. 1883. Syntypes. U.S.A. California: San Diego, Cleveland 9232, 6 Apr 1982 (SD), and 9 Apr 1892, (GH, NY!). Fig. 212A.

Ophioglossum crotalophoroides Walter, Fl. Carol. 256. 1788. Type. U.S.A. “Carolina,” Walter s.n. (BM?). Figs. 212E, F.

Stems upright, cylindrical, 2–4 mm diam., commonly bearing 2 leaves per stem; roots unbranched; fronds 3–5 cm long, 1–2 per stem, old leaf bases absent or inconspicuous; sterile blades erect to spreading, commonly more or less folded when alive, green, dull, without pale central band when dried, 1–4.3 ⫻ 0.5–1 cm, herbaceous, narrowly lanceolate, gradually tapering to base, apices acuminate; venation reticulate, with numerous parallel narrow areoles, each with one to several included veinlets; fertile spikes 1–2.5 times length of sterile blades.

Stems globose-bulbous, 3–11 mm diam.; roots unbranched; fronds 1–3 per stem, 2–5 cm long; sterile blades 0.8–4 ⫻ 0.8–2 cm, ovate, bases cordate, abruptly contracted, apices pointed to rounded; venation reticulate, not easily visible because of thick texture of blade, usually without included veinlets; fertile spikes 1–13 cm long, 0.5–8 times the length of the sterile blades.

Distribution. Terrestrial in damp meadows, pastures and open pinelands; 1000–3350 m. Se USA; Mexico; Guat, Hond, Nic, CR; Hisp; Col, Ven, Ec, Peru, Bol, Chile, Arg, Uru.

Distribution. Terrestrial in open grassy areas; 50–300(–2760) m. USA (Calif); Mexico. Specimens Examined. BCN (Jones 3750, MICH, NY, UC, US; Pringle 2821, CAS, MO, NY, US).

Ophioglossum californicum is distinct by its narrow, commonly folded sterile blades. The material from Baja California is typical, but material cited or named as such from the states of Durango, Chihuahua, DF, and Me´xico do not have folded blades and have longer than average fertile spikes; we treat these as O. nudicaule. More work is needed, however, to develop a clearer understanding of this complex.

Selected Specimens Examined. Chis (Xolocotzi & Sharp X434, MEXU). Dgo (McVaugh 21682, MICH, NY). Gro (Lorea 3388, 4400, FCME). Hgo (Medina C. 1666, ENCB). Jal (McVaugh 20590, MICH). Me´x (Pringle 4244, MO, NY, UC, US). Mich (Arse`ne 7410, MEXU). Nay (McVaugh 13337, MICH, US). Oax (Molseed s.n., Jul 1963, NY). Qro (Gonza´lez 1126, IEB). Sin (Sanders 2188, UCR).

Ophioglossum crotalophoroides is distinct in its cordate blades, globose-bulbous rhizomes, and the presence of very few included veinlets.

3. OPHIOGLOSSUM ENGELMANNII Ophioglossum engelmannii Prantl, Ber. Deutsch. Bot. Ges. 1: 351. 1883. Syntypes. U.S.A. Texas: New Braunfels, Lindheimer 53;

OPHIOGLOSSUM PALMATUM

Missouri, Engelmann s.n. (B?) (see Clausen, 1938: 139). Figs. 211D, E. Stems cylindrical, 2–4 mm diam., bearing 1-several leaves per stem; roots unbranched, blackish, 0.7–1.1 mm diam., about 15 (6–24) per plant; fronds 5–22 cm long, old leaf bases generally persistent, 0–4; sterile blades elliptic, 1–12 ⫻ 0.5–3.5 cm, bases narrowly to broadly cuneate, apices usually apiculate, rarely rounded; veins forming a bireticulate pattern, with large primary areoles and included secondary areoles; fertile spikes 3.5–16 cm long, 1–3 times the length of the sterile blades.

Distribution. Terrestrial on wooded slopes, presumably in limestone regions; 1000–2650 m. S USA; Mexico; Hond, CR. Selected Specimens Examined. BCS (Brandegee 671, GH, UC). Chih (Pringle 1564, NY). Chis (Breedlove 12882, NY). Coah (Chiang et al. 9624, LL, MEXU, MO, NY). DF (Lyonnet 256, MO, NY). Dgo (Reeder et al. 3401, GH, UC, US). Gro (Anderson & Anderson 4911, MICH). Jal (Pringle 8649, MO, UC). Me´x (Hinton 4301, LL, MO, NY). Mich (Dı´az Barriga 6221, MEXU). Nay (Ramı´rez R. 533, MEXU). Sin (Gentry 6359A, MICH, MO). SLP (Reeder et al. 3300, US). Son (White 4335, MICH). Unverified, Doubtful, or Mistaken Reports. Ags (Rosales 2119, HUAA, and two other collections, cited by Siquieros-Delgado & Gonza´lez-Adame, 2004 [as O. aff. engelmannii], but not verified).

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Ophioglossum tenerum Mett. ex Prantl, Ber. Deutsch. Bot. Ges.1: 352. 1883. Ophioglossum nudicaule L.f. var. tenerum (Mett. ex Prantl) R. T. Clausen, Mem. Torrey Bot. Club 19: 146. 1938. Type. U.S.A. Georgia: s. coll. s.n. (B) (see R. T. Clausen, 1938: 147).

Stems cylindrical or swollen but not globose, 2–4 mm diam.; roots unbranched; fronds 1–2 per stem, 2–8 cm long, basal sheaths inconspicuous, fragile, not persistent; sterile blades 0.5–2.5 ⫻ 0.2– 0.8 cm, lanceolate or elliptic to ovate–lanceolate, plane, bases cuneate or attenuate, apices acuminate; venation reticulate in smaller leaves, bireticulate in larger leaves, often obscured because of thick blade texture; fertile spikes 1.5–15 cm long, 2–6 times the length of the sterile blades; 2n⫽240 (India).

Distribution. Terrestrial in meadows and pastures; 1200–3100 m. Se USA; Mexico; Guat, Bel, Hond, Nic, CR, Pan; Cuba, Hisp; Col, Ven, Guy, Sur, Fr Gui, Ec, Galapagos, Peru, Braz, Bol, Arg; Africa, trop Asia, Australia, Hawaii. Selected Specimens Examined. Ags (McVaugh 18385, MICH). BCS (Brandegee s.n., 13 Sep 1893, UC). DF (Pringle 7086, ARIZ, MICH, NY). Dgo (Breedlove 45865, CAS). Hgo (Sa´nchez Mejorada 517, MEXU). Jal (McVaugh 16023, NY). Me´x (Mickel 713, ISC, MICH). Nay (McVaugh 18693, NY). Oax (Cruz Cisneros 1887, ENCB). Son (Steinman 93372, MEXU, MO). Tlax (Gonza´lez-Medrana 9366, MEXU). Ver (Rzedowski 36778, ENCB). Unverified, Doubtful, or Mistaken Reports. Chih (Pringle 1563, cited by Knobloch & Correll, 1979, but not verified). Mich (Arse`ne s.n., cited as O. ellipticum by Conzatti, 1939, but not verified).

The large, elliptical, apiculate blades with doubly areolate venation and cylindrical rhizomes distinguish O. engelmannii from O. vulgatum (q.v.) (obtuse, ovate blades with simply areolate venation). A large form of O. nudicaule, which has traditionally been treated as O. ellipticum, can be confused with the present species (see Wagner et al., 1984, for details) Small narrow specimens can be confused with O. polyphyllum.

Wagner et al. (1984) showed that two forms of O. nudicaule intergrade, the small form called var. tenerum, the larger one previously known as O. ellipticum. Both are found in Mexico, the smaller with reticulate veins, the larger with bireticulate veins. A pale band often occurs in the middle of dried blades.

4. OPHIOGLOSSUM NUDICAULE

5. OPHIOGLOSSUM PALMATUM

Ophioglossum nudicaule L.f., Suppl. Pl. 443. 1782. Type. South Africa. Cape of Good Hope, Thunberg s.n. (UPS-Thunb. 25286, microfiche UC!). Figs. 212C, D. Ophioglossum ellipticum Hook. & Grev., Icon. Filic. 1: pl. 40A. 1831. Type: Guyana. Demerara, Parker s.n. (K).

Ophioglossum palmatum L., Sp. Pl. 2: 1062. 1753. Fig. 211H. Cheiroglossa palmata (L.) C. Presl, Suppl. Tent. Pterid. 57. 1845. Ophioderma palmata (L.) Nakai, Bot. Mag. (Tokyo) 39: 193. 1925. Type. Plumier, Traite´ Foug. Ame´r., pl. 163. 1705, illustrating a plant from Hispaniola.

432

OPHIOGLOSSUM

Stems erect, cylindrical, 1–1.5 cm diam., with one to several leaves per stem; roots dichotomously branched; fronds pendent or spreading, 12–100 cm long, common stalk 5–50 cm long; sterile blades 4–50 ⫻ 4–30 cm, deeply palmately lobed, bases cuneate, the lobes long, mostly 1–2.5 cm wide, acute to obtuse at tips; venation reticulate, with included free veinlets; fertile spikes (1–)2–10(–16) per frond, 1–9 cm long, on adaxial side of blade bases, on short stalks to 6 cm long; juvenile plants with simple, linear-oblong leaves, and, if fertile, one central fertile spike.

Distribution. USA (Ariz, Tex); Mexico; Canary Isl., Azores, Africa, India, China, Hawaii. Selected Specimens Examined. BCS (Keill & McGill 8236, ENCB, MICH). Chih (McGill et al. 9270, ASU, NY). DF (Pringle 8538, GH, LL, NY, UC; Tryon & Tryon 5111, GH). Hgo (Pringle 8734, GH, MO, NY, UC). Oax (Purpus 3152, NY). Pue (Purpus 3152, NY, UC, US). SLP (Reeder et al. 3300, GH; Rzedowski 33406, ENCB, MEXU). Zac (Rollins & Tryon 58253, GH, US).

Distribution. Epiphytic in dense wet forests at low to middle elevations; 550–900 m. USA (Fla); Mexico; Bel, Hond, Nic, CR, Pan; Gr & L Ant; Col, Ven, Guy, Sur, Ec, Peru, Braz, Bol. Specimens Examined. Chis (Mu¨nch 172, DS). Oax (Calzada et al. 16575, MEXU; Galeotti 6603, BR; Rzedowski 33796, ENCB; Wendt et al. 5116, CHAPA).

The large divided blades and epiphytic habit readily distinguish this species.

6. OPHIOGLOSSUM POLYPHYLLUM Ophioglossum polyphyllum A. Braun in Seubert, Fl. Azor. 17. 1844. Type. Azores. Ins. Terceira, St. Barbara, Hochstetter 165 (B). Fig. 212B. Similar to O. engelmannii except for having more roots per plant (around 25 (15–30) vs. about 15 (6–24)), roots tan, 0.4–0.7 mm diam. (vs. blackish, 0.7–1.1 mm diam.), more persistent leaf bases 1–6(–10) (vs. 0–4), and often more slender blades; 2n⫽240 (India).

Pringle s.n. (10 Apr 1882, NY) from Baja California resembles O. polyphyllum in having old leaf bases and more roots. Perhaps these characters are not diagnostic.

7. OPHIOGLOSSUM RETICULATUM Ophioglossum reticulatum L., Sp. Pl. 2: 1063. 1753. Lectotype (chosen by Proctor, Fl. L. Antill. 2: 43. 1977). Plumier, Traite´ Foug. Amer., pl. 164. 1705, illustrating a plant from Hispaniola. Fig. 211A. Stems erect, cylindrical, 3–5 mm diam.; roots unbranched; fronds usually single, 7–40 cm long, basal sheaths conspicuous, leathery, persistent; sterile blades 1.5–9 ⫻ 1.2–5 cm, sessile or short-stalked, broadly ovate to nearly reniform, bases cordate or truncate, apices obtuse to sometimes acute, with minute apiculum when young; venation reticulate, areoles with included free veinlets; fertile spikes 6–23 cm long, 1–4 times the length of the sterile blades; 2n⫽ca. 960 (Jam), 872, 960, ca. 1140, ca. 1260 (India).

OSMUNDA

Distribution. Terrestrial in grassy fields and open or light woods; 900–2200 m. Mexico; Guat, Hond, Nic, CR, Pan; Gr & L Ant; Col, Ven, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol, n Arg; trop Africa, Asia. Selected Specimens Examined. Chis (Matuda 1614, NY). Col (Molseed 451, NY, UC). DF (Rose 9468, NY). Gro (Lorea 2135, NY). Jal (Pringle 11270, MO, NY). Me´x (Mickel 714, ISC, MICH). Mich (Labat 924, MEXU). Mor (Espejo 2612, IEB). Nay (McVaugh 16494, MICH). Oax (Hallberg 1803, NY). Qro (Dı´az B. & Zamudio 4981, IEB). Rev (Levin 2114, MICH). Sin (Ownbey & Ownbey 1892, NY, UC). SLP (Pringle 3816, MO, NY, UC). Ver (Purpus 8902, MO, UC).

Ophioglossum reticulatum is the most common species of the genus in Mexico and is distinguished by the cordate blade bases and cylindrical rhizomes. All species of Ophioglossum are notoriously difficult to find but probably more widespread and common than collections indicate.

8. OPHIOGLOSSUM VULGATUM Ophioglossum vulgatum L., Sp. Pl. 2: 1062. 1753. Lectotype (chosen by Jonsell & Jarvis, 1993: 71). Herb. Clifford 472, Ophioglossum 1 (BM). Figs. 211B, C, F, G. Ophioglossum vulgatum L. var. pycnostichum Fernald, Rhodora 41: 494. 1939. Ophioglossum pycnostichum (Fernald) A˚. Lo¨ve & D. Lo¨ve, Taxon 26: 324. 1977. Type. U.S.A. Virginia: Dinwiddie Co., east of Burgess Station, Fernald & Long 9796 (GH!; isotype PH). Ophioglossum pringlei Underw. ex Conz., Fl. Tax. Mex. 1: 141. 1939, hom. illeg. sine descr. Lat. Type. Mexico. Hidalgo: Trinidad Iron Works, Pringle 8924 (isotypes GH!, NY!, UC!). Ophioglossum dudadae Mickel, Brittonia 44: 313. 1992. Type. Mexico. Oaxaca: Distrito Ixtla´n, 32 km N of Ixtla´n de Jua´rez on Hwy 175 at the Llano de las Flores, Mickel 9639 (NY!).

Stems cylindrical, 2–4 mm diam.; roots unbranched; fronds usually 1 per stem (rarely several), (4–)12–36 cm long, basal sheaths conspicuous, leathery, persistent; sterile blades (2–)5–10(– 15) ⫻ 1.5–6 cm, mostly ovate-lanceolate to elliptic, broadest and abruptly tapering at bases, obtuse or minutely apiculate to broadly rounded at apices; venation reticulate with included free veinlets; fertile spikes 4–27 cm long, 1–3 times the sterile blade length; 2n⫽480 (Europe and India), ca. 1320 (USA).

433

Distribution. Terrestrial in shaded, moist to wet woods; 2200– 2900 m. Canada, se USA; Mexico; temperate Europe and Asia. Selected Specimens Examined. Chis (Breedlove 12911, 15336, DS). DF (Pringle s.n., 19 Aug 1896, NY). Hgo (Pringle 8924, ENCB, GH, MO, NY, UC, 15705, GH, MICH, MO; Wagner 85067, MICH). Oax (Mickel 3837, NY p.p., 9639, NY). Pue (Barnes & Land 503, GH). Ver (Storer 145, MICH).

Ophioglossum dudadae Mickel was based primarily on its relatively small size. Revisiting the type locality in 1998, we found the plants to be much larger. Another locality about 40 km away, the Llano Verde, had been grazed for years when we first visited there in 1969, plants of Ophioglossum were small. In 1998, in the absence of grazing the previous ten years, the angiosperm flora was taller. All Ophioglossum plants were larger and seemed to be O. vulgatum. It seems unlikely that O. vulgatum has replaced O. dudadae completely. It is more likely that the plants are sometimes small because of grazing or other environmental conditions. Consequently, we now treat O. dudadae as an environmentally induced form of O. vulgatum. Small plants of O. vulgatum resemble O. petiolatum Hook., but are distinct in roots 1.2–1.6 mm diam. (vs 1–1.2 mm), blades elliptic rather than lanceolate, vein areoles mostly elongate rather than isodiametric, fertile stalks and spikes only slightly exceeding the sterile blades, and occurrence at high elevations. The report of O. petiolatum in Chiapas (Smith, 1981 at 2200 m) may refer to small plants of O. vulgatum. Doubtful Species Ophioglossum petiolatum Hook., Exot. Fl. 1: 56. 1823. Type. Based on cultivated specimens, origin Antilles (K). For additional synonymy, see Clausen, 1938. See discussion under O. vulgatum.

9 0 . O SM U N D A Osmunda L., Sp. Pl. 2: 1063. 1753; Gen. Pl., ed. 5, 484. 1754. Lectotype (first chosen by Le´man?, Dict. Sci. Nat. 37: 9. 1825): Osmunda regalis L.

434

OSMUNDA

Osmundastrum C. Presl, Gefa¨ssbu¨ndel Farrn 18. 1847. Type: Osmundastrum cinnamomeum (L.) C. Presl [⬅ Osmunda cinnamomea L.]. For details of typification of these names, see Pichi Sermolli, Webbia 26: 506. 1972.

Terrestrial, of moist habitats; rhizomes horizontal to usually erect, stout, surrounded by persistent, tough leaf bases and a wiry dense root mantle, caudices often subterranean or partly aerial, usually dichotomously branched or with a single arborescent trunk, apices lacking scales, but sometimes hairy; stipes not articulate, bases somewhat flared or expanded just above base to form sheathing, stipule-like wings; fronds crown-forming, dimorphic or hemidimorphic, sporophylls taller and more erect, croziers circinate, clothed with matted white to brownish hairs; sterile blades (or sterile portions) pinnate-pinnatifid to 2(–3)pinnate, rarely 1-pinnate, laminae glabrous but axes densely hairy, especially at pinna bases and in young fronds; veins of sterile fronds (or sterile portions) free, simple or several times forked; fertile blades (or fertile portions) usually 2-pinnate, lacking green lamina, sporangia borne in dense paniculate clusters on ultimate veinlets; sporangia large, globose, splitting longitudinally across the distal end, with short, many-rowed stalks; spores green, short-lived, tetrahedral-globose, surface densely rugose or echinate; gametophytes epigeal, green, flat, cordate to elongate, the center thickened; x⫽22. Osmunda is subcosmopolitan, with about 10 species in temperate and subtropical regions. It occurs mostly in marshes, swamps, and other wet areas. The two Mexican species are very widespread, but relatively uncommon, except locally in suitable habitats. They are occasionally segregated into two genera. The other two genera in the family, Todea (South Africa and Australasia) and Leptopteris (Australasia) have sporangia borne on relatively unmodified pinnae. Osmunda is distinct by its dimorphic or hemidimorphic fronds more than once-pinnately divided, free veins, rhizome indument of hairs, green, globose spores, and nearly unique (in the ferns) chromosome base number. The family Osmundaceae is isolated phylogenetically and has an extensive fossil record dating back to the Carboniferous. Recent molecular studies indicate that the family is basalmost within the leptosporangiate ferns and forms the sister group to all other extant familes in the clade (Pryer et al., 1995, 2001). The eusporangiate ferns (Marattiaceae and Ophioglossaceae), plus Psilotaceae (whisk ferns) and Equisetaceae (horsetails) insert below the leptosporangiate ferns in most recent analyses based on molecular data. References Benedict, R. C. 1909. Osmundaceae. N. Amer. Flora 16: 27–28; Bobrov, A. 1968. Species novae Osmundacearum. Novosti Sist. Vyssh. Rast. 1968: 5–8; Hewitson, W. 1962. Comparative morphology of the Osmundaceae. Ann. Missouri Bot. Gard. 49: 57–93; Serbet, R. & G. W. Rothwell. 1999. Osmunda cinnamomea (Osmundaceae) in the Upper Cretaceous of western North America: Additional evidence for exceptional species longevity among filicalean ferns. Int. J. Pl. Sci. 160: 425–433.

Key to the Mexican Species of Osmunda 1. Fronds entirely dimorphic, the fertile and sterile ones different; sterile blades pinnate-pinnatifid, fertile blades

totally lacking laminar tissue and soon drying and collapsing; mature pinnae with woolly tufts of hair at their base abaxially. ..................................................... 1. O. cinnamomea. 1. Fronds partly dimorphic (hemidimorphic), the sporangia confined to a few distal pinnae (distal 1⁄4–1⁄3 of blades), with basal portion of blades sterile, bipinnate; mature pinnae glabrous. ............................... 2. O. regalis var. spectabilis.

1. OSMUNDA CINNAMOMEA Osmunda cinnamomea L., Sp. Pl. 2: 1066. 1753. Type. U.S.A. “Marilandia,” Kalm s.n. (LINN 1244.12.). Figs. 213D, E. Osmundastrum cinnamomeum (L.) C. Presl, Abh. Ko¨nigl. Bo¨hm. Ges. Wiss., ser. 5, 5: 326. 1848.

Rhizomes stout, horizontal, with hairs and old leaf bases; fronds 0.5–1.5 m long, holodimorphic; stipes 1⁄3 the frond length, stramineous, darker at base, glabrous or with some soft, lax, cinnamon hairs; sterile blades pinnate-pinnatifid, 35–100 ⫻ 20 cm, elliptic to ovate-lanceolate, gradually narrowed into a pinnatifid apex, chartaceous to subcoriaceous; pinnae opposite to subopposite, glabrous except for tuft of short hairs at base of each pinna abaxially, sessile, to 13 ⫻ 1.5–4 cm, spreading to ascending; pinnules slightly ascending or falcate, margins entire, broadly acute at tip, short hairs scattered on veins and margin of pinnae at base; veins free, 1-forked; fertile blades bipinnate, narrow, ephemeral, axes densely covered with reddish brown hairs; 2n⫽44 (Jam, CR).

Distribution. Swampy areas, often in the open, or in light woods by springs, in swamp at edge of canyon; 900–1900 m. Canada, e USA; Mexico; Guat, Hond, Salv, CR; Berm, Gr Ant; Col, Ven, Ec, Peru, Braz, Bol, Parag; e Asia. Specimens Examined. Chis (Breedlove 22198, DS, MEXU, MO, NY; Ghiesbreght 300, K, NY). Oax (Mickel 902, NY, US; Rivera Reyes 242, MEXU). Pue (Miranda 3428, MEXU; Sharp 45349, US). Ver (Liebmann s.n., Feb 1842, GH, photo XAL!). Unverified, Doubtful, or Mistaken Reports. Tab (reported by Mickel & Beitel, 1988, and by Magan˜a, 1992, but not verified).

The wholly dimorphic fronds and pinnate-pinnatifid sterile blades distinguish this species from O. regalis. Serbet and Rothwell (1999) reported fossils of this species from the Upper Cre-

PECLUMA

taceous of western North America that resemble the extant plants so closely that they cannot be distinguished.

2. OSMUNDA REGALIS VAR. SPECTABILIS Osmunda regalis L. var. spectabilis (Willd.) A. Gray, Manual, ed. 2, 600. 1856. Figs. 213A–C. Osmunda spectabilis Willd., Sp. Pl., ed. 4, 5(1): 98. 1810. Type. U.S.A. Pennsylvania: Muhlenberg s.n. (B-Willd. 19504!). Osmunda mexicana Fe´e, Me´m. Foug. 9: 43. 1857. Type. Mexico. Oaxaca: Galeotti 6388 (P?; isotypes BR!–2 sheets). Osmunda palmeri A. E. Bobrov, Novosti Sist.Vyssh. Rast. 1968: 8. Type. Mexico. Jalisco: Rı´o Blanco, Palmer 660 (LE!; isotypes GH!, K!, UC!, US!).

Rhizomes massive, compact, horizontal to suberect, apices with long hairs; fronds 35–180 cm long, hemidimorphic, the fertile part occupying the distal 1⁄4–1⁄3 of the blades and greatly constricted (lacking green laminar tissue); stipes 1⁄3–1⁄2 the frond length, glabrous, stramineous to reddish brown, darker at base; blades bipinnate, 20–35 cm wide, elliptic, broadly lanceolate; pinnae with a conform terminal pinnule, chartaceous to subcoriaceous, glabrous at maturity; pinnules distant, very shortly petiolulate (less than 1 mm) to sessile at blade tip, narrowly elliptic, bases rounded to subcordate, entire to subcrenulate, apices obtuse to acute, glabrous at maturity; veins 2-forked; 2n⫽44 (Chis, CR).

Distribution. Swampy areas in the open or light woods, pineoak-Liquidambar forests, oak and Magnolia forests, near barranca; (250–)900–1800(–2700) m. Canada, e USA; Mexico; Guat, Hond, Salv, Nic, CR; Berm, Bah, Cuba, Jam, Hisp; Col, Ven, Ec, Peru, Braz, Bol, Parag, Arg, Uru. Var. regalis occurs in Europe, Africa, and Asia. Selected Specimens Examined. Chis (Breedlove 21253, DS, MEXU, NY). Gro (Lorea 1943, FCME). Hgo (Ramı´rez & Pe´rez G. 677, MEXU). Jal (Mexia 1822, CAS, DS, NY, UC, US). Me´x (Hinton 5394, DS, MEXU, NY, UC). Mich (Garcı´a & Dı´az B. 3875, IEB). Nay (Palmer 1947, NY, US). Oax (Mickel 4972, NY, UC). Pue (Sharp 45348, MEXU, US). Tab (Cowan 3100, CAS, MEXU, NY). Ver (Ventura A. 18385, CAS, IEB, MEXU). Unverified, Doubtful, or Mistaken Reports. Ags (Siqueiros 2828, and one other collection, HUAA, cited by Siqueiros-Delgado & Gonza´lez-Adame, 2004, but not verified). Qro (Ameneyro s.n.,

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ENCB, cited by Dı´az-Barriga & Palacios Rios, 1992, but not verified).

This variety seems to occur throughout the New World. Fronds and fertile panicles are more open, the stipes and rachises less stout, and the blades less coriaceous than var. regalis, which was recently typified by Jonsell and Jarvis (in Jarvis et al., 1993) on the basis of Herb. Burser vol. 20: 26 (UPS, microfiche UC!). New World material looks distinct enough to be treated as a different species from O. regalis sensu stricto (Old World), but more study of both herbarium and living material is needed to properly delimit these taxa. In Mexico, O. regalis is far more common and widespread than O. cinnamomea, but there are nearly as many collections from Chiapas (about 35) as from all remaining states.

91. PECLUMA Pecluma M. G. Price, Amer. Fern J. 73: 109. 1983. Type: Pecluma pectinata (L.) M. G. Price [⬅ Polypodium pectinatum L.]. Mostly epiphytic, less often epipetric or terrestrial; rhizomes long- to often short-creeping, unbranched, never glaucous, sometimes proliferous by root buds; rhizome scales concolorous to bicolorous, non-clathrate, basally attached, surfaces glabrous or hairy (comose), margins entire, toothed, or short-ciliate; fronds monomorphic, distant (ca. 1.5 cm apart) or clumped; stipes black to reddish brown, rarely lighter in color, terete (never channeled), articulate, phyllopodia 1–2 mm long; blades pinnatisect with numerous linear or narrowly deltate pinnae, bases attenuate to truncate, pinnae curling in response to drought, sometimes with scattered to numerous scales along the axes; indument adaxially on rachises, costae, and sometimes laminae between veins of mostly septate hairs, abaxially with similar sometimes longer hairs on rachises and costae, laminae between veins often with inconspicuous appressed, septate hairs 0.1–0.2 mm; veins free, rarely casually anastomosing, simple or 1–4-forked; sori round, exindusiate, terminal on first acroscopic veinlet, in single rows on each side of costae; soral paraphyses simple or branched, multicellular, sometimes clavate at the tips; sporangia glabrous or with 1–3-celled hairs (0–5 per sporangium) on the capsules; spores yellowish to whitish, bilateral (globose in an apogamous species, P. dispersa), surfaces verrucate to tuberculate with SEM magnification, seemingly smooth under the light microscope; x⫽37. Pecluma comprises about 28 species, all neotropical. Closest affinities appear to be with those species of Polypodium that lack scales on the blades, particularly the group of free-veined species related to P. hartwegianum, P. chiapense, and P. diplotrichum. The species of Pecluma (as the “Polypodium pectinatumplumula complex”) were revised by Evans (1968) and later segregated as a genus separate from Polypodium by Price (1983). Some species in this group have sometimes been included among the grammitid ferns, in Ctenopteris (e.g., by Copeland, 1955), but the similarity is only superficial. Pecluma differs from Polypodium

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PECLUMA

s.str. in having often short-creepingt, non-branching, nonglaucous rhizomes, proliferous roots, non-clathrate basally attached rhizome scales, terete stipes and rachises, and pectinate fronds. From the grammitid genera, Pecluma differs by having bilateral, non-green spores, two-rowed sporangial stalks, usually comose rhizome scales, and hyaline to yellowish (vs. maroon), obviously septate and often crispate hairs. A peculiar hair type, called “ctenoid” by Evans (1968), is found along the stipes and rachises of several species, most com-

monly in P. consimilis, P. divaricata, and P. ptilodon. These hairs consist of an arching or appressed axis of about four cells, each with a 1–2-celled lateral branch, all arising from one side of the axis, producing a comb-like trichome. References Copeland, E. B. 1955. Ctenopteris in America. Philipp. J. Sci. 84: 381–475; Evans, A. M. 1968 [1969]. Interspecific relationships in the Polypodium pectinatumplumula complex. Ann. Missouri Bot. Gard. 55: 193–293; Price, M. G. 1983. Pecluma, a new tropical American fern genus. Amer. Fern J. 73: 109–116.

Key to the Mexican Species of Pecluma 1. Rachises abaxially brownish, lacking scales. 2. Rhizome scales orangish to ferruginous; rachises and laminae abaxially covered with silvery, spreading hairs. ...................... 7. P. hygrometrica. 2. Rhizome scales dark orange- or red-brown to blackish; rachises and laminae abaxially glabrous or with spreading to appressed hairs, these often light brown or reddish. 3. Rhizome scales ovate; pinnae 1.5–5 mm wide; ctenoid (comb-like) hairs absent along stipes and rachises abaxially. 4. Sori medial or supramedial; receptacular paraphyses few to numerous, simple and straight. .............................................. 1. P. alfredii. 4. Sori marginal or submarginal; receptacular paraphyses numerous, twisted and with a subterminal clavate, multicellular head. .......................................................................................................................................................................................10. P. sursumcurrens. 3. Rhizome scales linear-lanceolate; pinnae mostly 4–10 mm wide; ctenoid hairs present along stipes and rachises abaxially. 5. Proximal pinnae scarcely reduced, or lowermost at least half as long as the longest pinnae; sporangia glabrous. ....... 5. P. divaricata. 5. Proximal pinnae less than 1⁄2 as long as the longest pinnae, reduced to 3 or more auricles; sporangia glabrous or bearing hairs. 6. Laminar surfaces abaxially glabrous or with evenly distributed appressed trichomidia 0.1 mm; sporangia bearing hairs. ....................................................................................................................................................................................... 3. P. consimilis. 6. Laminar surfaces abaxially with oblong patch of erect hairs around sori; sporangia glabrous. ........ 9. P. ptilodon var. bourgeauana. 1. Rachises abaxially red-brown to black, with persistent scales, these usually conspicuous. 7. Rhizome scales lanceolate with acuminate apices, comose. 8. Pinnae linear, the proximal ones neither deflexed nor reduced to mere auricles; veins mostly 1-forked. ............................... 8. P. plumula. 8. Pinnae lanceate, the proximal ones deflexed, sometimes reduced to auricles; veins 2-forked. 9. Adaxial laminar surfaces with scattered hairs 0.3–0.6 mm, especially submarginally; rhizome scales entire; spores reniform, 64 per sporangium. ............................................................................................................................................................. 2. P. atra. 9. Adaxial surfaces glabrous or with sparse hairs; rhizome scales denticulate; spores globose, 32 per sporangium. ............. 4. P. dispersa. 7. Rhizome scales ovate to cordate, rounded to acute at apices, non-comose. 10. Rhizome scales golden or orangish; sporangia with hairs on the capsules; lamina surfaces adaxially with scattered hairs 0.3– 0.5 mm. ................................................................................................................................................................................................ 6. P. ferruginea. 10. Rhizome scales brown, red-brown, or dark orange-brown; sporangia lacking hairs; lamina surfaces adaxially glabrous. 11. Rachis scales broadly ovate, cordate, numerous and conspicuous, often inrolled at the bases; proximal pinnae Ⳳ perpendicular to the rachises. ......................................................................................................................................................... 1. P. alfredii. 11. Rachis scales narrowly elliptic, few and sometimes inconspicuous, flat, or seemingly absent. 12. Pinna margins lacking hairs. ................................................................................................................................................... 1. P. alfredii. 12. Pinna margins hairy. .....................................................................................................................................................10. P. sursumcurrens.

1. PECLUMA ALFREDII Pecluma alfredii (Rosenst.) M. G. Price, Amer. Fern J. 73: 113. 1983. Polypodium alfredii Rosenst., Repert. Spec. Nov. Regni Veg. 22: 15. 1925. Type. Costa Rica. Turrialba, Brade & Brade 697 (S; isotype US!). Polypodium tablazianum Rosenst., Repert. Spec. Nov. Regni Veg. 22: 14. 1925. Lectotype (chosen by Lellinger, Proc. Biol. Soc. Wash. 89: 727. 1977). Costa Rica. Cartago: Carpintera, Brade & Brade 149 (S; isolectotypes UC!, US!).

Rhizomes short-creeping, 3–7 mm diam.; rhizome scales light to dark reddish brown, 1–2 ⫻ 0.8–1 mm, ovate, rounded to acute at tips, entire; fronds 10–60 cm long, clumped; stipes dark reddish

brown to brown, 1/6–2/5 the frond length, with acicular hairs 0.1–0.2(–0.5) mm; blades pectinate, narrowly ovate, 4–6.5 cm wide; rachises abaxially with many tan, bullate or cordate, entire, acute scales ca. 2 mm long; pinnae linear, 2–5 cm ⫻ 1.5–3 mm, greatly reduced and deflexed at blade bases, bases of pinnae equilateral or more surcurrent, apices obtuse to acute, costae of proximal pinnae decurrent onto the rachises; veins free, 1-forked; indument adaxially of scattered hairs 0.1–0.3 mm on costae, laminar surfaces glabrous, abaxially the costal surfaces between veins with appressed hairs 0.1–0.2 mm, margins with occasional hairs 0.3–0.5 mm, or marginal hairs absent; sori round, medial, with clavate paraphyses 0.5–0.8 mm; sporangia glabrous; spores reniform, 64 per sporangium. Distribution. Epiphytic or epipetric in humid oak woods and

PECLUMA ALFREDII

on dry exposed banks; 600–3050 m. Mexico; Guat, Hond, Salv, Nic, CR. Pecluma alfredii is unique in having the combination reddish brown, ovate rhizome scales, and ovate-cordate, often bullate scales along the rachises abaxially. There is great variability in stipe and rachis color (reddish brown to nearly blackish), hair density and location (pinna margins glabrous or hairy), and size of pinnae. Var. alfredii is restricted to Costa Rica and is distinguished by the pinnae completely lacking hairs along the margins and the glabrescent rachises abaxially. In Chiapas, this is known by the Tzotzil names “ne kotz’ tzib” and “xelajtik tzib,” concoctions of which are used to treat snake bite and infections of the urinary tract. Key to the Mexican Varieties of Pecluma alfredii 1. Rachis scales linear-lanceolate abaxially. ................. 1b. var. occidentalis. 1. Rachis scales ovate-cordate abaxially. ....................... 1a. var. cupreolepis.

1a. Pecluma alfredii var. cupreolepis (A. M. Evans) A. R. Sm., comb. nov. Figs. 214C, D, 215M. Polypodium cupreolepis A. M. Evans, Ann. Missouri Bot. Gard. 55: 224. 1969. Pecluma cupreolepis (A. M. Evans) M. G. Price, Amer. Fern J. 73: 114. 1983. Type. Mexico. Michoaca´n: Hills of Patzcuaro, Pringle 3353 (US!; isotypes B, CAS!, DS!, GH!, MICH, MO!, NY!, UC!, US!).

Differing from var. occidentalis in having ovate-cordate scales on the rachises abaxially; pinna margins usually with at least a few hairs.

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cited as P. alfredii by Arreguı´n et al., 2001, but not seen, probably P. sursumcurrens). Ver (cited by Palacios-Rios, 1992, on the basis of Ventura 8462, ENCB, MEXU, NY, but that is P. plumula).

A specimen cited from Edo. Puebla (Purpus 1828, GH, MO, US) by Evans (1968) is from Iztaccihuatl and is mostly likely from Edo. Me´xico. The Chiapas collection cited is from the southern Sierra Madre. The specimen cited from Tamaulipas is atypical in having numerous small, cordate scales along the costae abaxially. Var. cupreolepis is very closely related to var. alfredii, and differs primarily by the more numerous hairs on the rachises abaxially. In addition, most specimens (but not the type) differ by the presence of septate hairs on the pinna margins. Most specimens cited as P. alfredii by Evans (1968) have rachis scales identical to those of the type of Polypodium cupreolepis. Even two of the syntypes of P. tablazianum Rosenst. (Brade 149, 696, UC), treated by Evans as a synonym of P. alfredii, have rachis scales of typical P. cupreolepis. Polypodium alfredii and P. cupreolepis were treated as conspecific by Moran (in Davidse et al., 1995). Pecluma alfredii var. cupreolepis also resembles P. ferruginea, but differs by its glabrous sporangia, broader, more cordate rachis scales, glabrous laminar surfaces adaxially, and reddish brown rhizome scales. 1b. Pecluma alfredii var. occidentalis A. R. Sm., var. nov. Type. Mexico. Revillagigedo Islands: Socorro, Mason 1633 (holotype CAS!). Figs. 214A, B. A var. cupreolepis squamis lineari-lanceolatis secus rhachides abaxialiter differt. (L., occidentalis, western, referring to the distribution of this variety in westen Mexico.)

Differing from var. cupreolepis in the linear-lanceolate scales on the rachises abaxially; pinnae margins lacking hairs.

Distribution. Epiphytic or epipetric, pine-oak forests; 1250– 2600 m. Mexico; Guat, Hond (Kimnach 400a, UC), Salv, Nic, CR. Selected Specimens Examined. Chis (Breedlove 31941, DS). Col (Goldsmith 40, UC). DF (Purpus 1828, NY, US). Dgo (Sa´nchez 726, MEXU). Gro (Breedlove 65115, CAS). Jal (Wilbur & Wilbur 1927, DS, US). Me´x (Weller 596, ENCB, UC). Mich (Rzedowski & de la Sota 18336, ENCB, NY, US). Mor (Hinton 17208, ENCB, NY). Oax (Mickel 5013, 6206, NY, UC). Pue (Lyonnet 460900007, MEXU). Sin (Gentry 6337, DS, NY). Tam (Stanford et al. 2055, NY). Unverified, Doubtful, or Mistaken Reports. Hgo (Pringle 13475, US, cited by Evans, 1968, but not verified; this no. also cited as P. sursumcurrens, MICH, from Edo. Puebla, Honey Station, by Evans, 1968). Qro (Amaneyro s.n., ENCB, and Ferna´ndez 4070, 4270, ENCB,

Distribution. Terrestrial, epipetric, and epiphytic on trunks of trees in forests; 650–2150 m. Mexico. Selected Specimens Examined. Dgo (Reeder & Reeder 2487, US). Jal (Breedlove 45663, CAS, MEXU; Dı´az L. 5597, UC). Rev (Barkelew 236, UC p.p., US; Felger 15817, UC, US; Flores Palacios 756, UC; Moran 15817, 25502, UC; Villarreal 59b, ENCB). Sin (Breedlove 17110, 44961, CAS; Correll & Correll 28860, MEXU, US; Ownbey & Ownbey 1939, US).

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PECLUMA

This variety is apparently confined to the northwestern part of the range of P. alfredii, in the Sierra Madre Occidental. Collections from Isla del Socorro are consistently without marginal hairs on the pinnae. Those from Sinaloa and Durango have a few cilia and thus approach var. cupreolepis. Collections from both areas lack the large cordate scales of var. cupreolepis. Barkelew 236, UC p.p., was cited by Evans (1968) as P. ferruginea, but the sporangia of this collection lack hairs.

4723, IEB). Tab (Matuda 3372, MEXU, US). Tlax (Vela´zquez s.n., ENCB). Ver (Copeland herb. 124, BM, MEXU, MICH, NY, UC, US). Unverified, Doubtful, or Mistaken Reports. Camp (Ortega & Uca´n Ek’ 1309, XAL, cited by Palacios-Rios, 2002a, but not verified).

Among the Pecluma species with black stipes, P. atra is distinct in its scattered adaxial blade hairs and slender, entire rhizome scales. Its closest ally is P. dispersa, which is the apogamous triploid hybrid with P. plumula as the other parent.

2. PECLUMA ATRA 3. PECLUMA CONSIMILIS Pecluma atra (A. M. Evans) M. G. Price, Amer. Fern J. 73: 113. 1983. Figs. 214J, K. Polypodium atrum A. M. Evans, Ann. Missouri Bot. Gard. 55: 237, f. 18. 1969. Type. Belize. El Cayo: Mountain Pine Ridge, San Augustin, Rı´o Frio, Lundell 6639 (US!; isotypes GH!, MICH!).

Rhizomes short-creeping, 4–6 mm diam.; rhizome scales reddish brown, 3 ⫻ 0.5–0.8 mm, lanceolate, comose, entire; fronds 20–65 cm long, clumped; stipes black, 1/6–2/5 the frond length, with spreading hairs 0.1–0.2 mm; blades narrowly oblong, 6–12 cm wide, slightly reduced at bases, deflexed, sometimes to auricles; rachises abaxially with reddish brown, linear-deltate scales; pinnae linear-deltate, 3–6 cm ⫻ 3–7 mm, obtuse, surcurrent in proximal half of blade, costae of proximal pinnae at right angles to the rachises; veins free, 1–2-forked; indument adaxially of sparse to moderately dense hairs 0.1–0.2 mm on costae, laminar surfaces with scattered septate hairs 0.3–0.6 mm, abaxially the costae with scattered hairs 0.2–0.5 mm, surfaces between veins with numerous appressed hairs 0.1–0.2 mm, margins with scattered hairs 0.3–0.5 mm; sori round, medial, with clavate paraphyses 0.3 mm; sporangia glabrous or some with hairs ca. 0.1 mm; spores reniform, 64 per sporangium.

Pecluma consimilis (Mett.) M. G. Price, Amer. Fern J. 73: 113. 1983. Figs. 214L, M, 215L. Polypodium consimile Mett., Ann. Sci. Nat. Bot., se´r. 5, 2: 253. 1864. Type. Colombia. Norte de Santander: Ocan˜a, Schlim 633 (B; isotypes BR, G, L).

Differs form P. ptilodon by: laminar surfaces glabrous abaxially or with evenly dispersed appressed hairs 0.1 mm, lacking erect hairs in oblong patches around the sori; sporangia with weak hairs 0.1 mm on the capsules.

Distribution. Epiphytic in montane forests; 100–1650 m. Mexico; Guat, Nic, CR, Pan; Jam, Hisp; Col, Ven. Also reported from Honduras and Ecuador by Moran (in Davidse et al., 1995). Selected Specimens Examined. Chis (Breedlove 20964, 27886, DS, MEXU, 38195, DS; Martı´nez S. 5954, MEXU). Oax (Mickel 4809, 5831, 6413, ENCB, NY, UC). Ver (Lira 186, ENCB, MEXU; Ventura A. 14733, ENCB; Wendt & Ishiki 5881, MEXU, NY). Unverified, Doubtful, or Mistaken Reports. QR (Chater 40, MEXU, cited as P. consimilis by Sousa & Cabrera, 1993, is P. dispersa).

Distribution. Terrestrial or epipetric, often on limestone, in wet lowland and montane rain forests, seasonal evergreen forests; 0–1150(–2250) m. Mexico; Guat, Bel, Hond, Nic. Selected Specimens Examined. Chis (Breedlove 21926, DS, MEXU, NY, 38180, 38680, DS). Oax (Martı´nez-Caldero´n 693, ENCB; Mickel 6823, ENCB, UC, US; Orcutt 5211, DS). Qro (Carranza & Dı´az B.

Pecluma consimilis is distinct by its brown stipes, comose rhizome scales, reduced proximal pinnae, and evenly distributed abaxial trichomidia, without hairs in an oblong patch around the sori. Most Mexican specimens of P. consimilis differ from P. ptilodon in bearing minute hairs on the sporangial capsules; however, this character is not completely reliable, as a few specimens

PECLUMA DIVARICATA

of P. consimilis appear to lack hairs and a few specimens of P. ptilodon have hairs on the sporangia. Two varieties of P. consimilis were recognized by Evans (1968), but the non-typical one, var. pastazensis, has since been elevated to species rank [Pecluma pastazensis (Hieron.) R. C. Moran; Nordic J. Bot. 15: 58. 1995] and is confined to Colombia and Ecuador.

4. PECLUMA DISPERSA Pecluma dispersa (A. M. Evans) M. G. Price, Amer. Fern J. 73: 114. 1983. Figs. 215N, O. Polypodium dispersum A. M. Evans, Amer. Fern J. 58: 173, t. 27. 1968. Type. U.S.A. Florida: Citrus Co., Pineola Grottoes, Evans 2008 (MICH!; isotypes NY!, TENN, US!).

Rhizomes short-creeping, 4–8 mm diam.; rhizome scales reddish brown, lustrous, 2–4 ⫻ 0.3–1(–1.5) mm, lanceolate, acuminate, entire; fronds 27–63 cm long, clumped; stipes black, 1/7– 2/7 the frond length, adaxial surfaces densely pilose with acicular hairs 0.1 mm and scattered, appressed ctenoid hairs; blades narrowly deltate, 5–10 cm wide, reduced to lobes; pinnae somewhat deflexed proximally, the largest 2.5–5 cm ⫻ 2–5 mm, obtuse, equilateral at bases, surcurrent and decurrent, costae of proximal pinnae decurrent onto rachises; rachises abaxially with dark reddish brown, lustrous, ovate to narrowly deltate, acuminate scales; veins free, 1–2-forked; indument adaxially of numerous hairs 0.1– 0.3 mm on costae, laminar surfaces with scattered hairs to ca. 0.5 mm, abaxially the costae with scattered septate hairs to 0.5 mm, surfaces between veins with appressed hairs 0.1–0.2 mm, margins with scattered to numerous hairs 0.3–0.5 mm; sori round (to oblong), medial, with a few clavate paraphyses 0.5– 0.8 mm; sporangia glabrous or often with several inconspicuous hairs ca. 0.1 mm; spores globose, 32 per sporangium; 2n⫽111, apogamous (Fla, Jam).

Distribution. Epipetric or in rock crevices (or epiphytic or terrestrial), pine-oak forests; 0–1700 m. USA (Fla), Mexico; Guat, Bel, Hond, Nic (Grijalva & Burgos 1545, UC); Gr Ant; Col, Ven, Ec, Galapagos, Braz, Bol.

439

Selected Specimens Examined. Chis (Breedlove 26041, DS, MEXU, NY, 32314, DS, NY). Hgo (Cota & Gimate 7776, NY). Oax (Herna´ndez G. 755, CHAPA, MEXU, MO, NY; Mickel 4737, ENCB, NY, UC, US). QR (Bashor 497, NY; Te´llez 3560, MEXU). SLP (Blassingame 1316, US; Pringle 3999, GH, US; Reeves 6134, ASU). Tam (Palmer 562, NY, US). Ver (Woronow 3002, US).

Among the black-stiped species of Pecluma, P. dispersa is distinct in its densely puberulent stipes and rachises adaxially, lanceolate, denticulate rhizome scales, and 32 globose spores per sporangium. It is similar to P. atra in having numerous deltate to lanceolate scales on the rachises abaxially, and these scales sometimes extend to the proximal portions of the costae. There are scattered septate hairs on the laminae adaxially, but usually fewer and more concentrated toward the pinna margins compared to P. atra (but see Reeves 6134, which equals P. atra in hairiness adaxially). On morphological and cytological grounds, Evans (1968) postulated that P. dispersa was the hybrid between P. atra, a diploid, and P. plumula, a tetraploid. It reproduces successfully by an apogamous life cycle, well documented by Evans.

5. PECLUMA DIVARICATA Pecluma divaricata (E. Fourn.) Mickel & Beitel, Mem. New York Bot. Gard. 46: 269. 1988. Figs. 215G, H, J. Polypodium divaricatum E. Fourn., Mexic. Pl. 1: 80. 1872. Type. Mexico. Veracruz: Zacuapan, Galeotti 6287 (P!; isotype BR!), cited as Galeotti 6278, in error, by Mickel & Beitel, 1988. Polypodium bolivianum Rosenst., Repert. Spec. Nov. Regni Veg. 5: 236. 1908. Pecluma boliviana (Rosenst.) M. G. Price, Amer. Fern J. 73: 113. 1983. Type. Bolivia. South Yungas, Sirupaya near Yanacachi, Buchtien 481 (S; isotypes P!, US!), cited incorrectly as Buchtien 421 by Mickel & Beitel, 1988.

Rhizomes short-creeping, 6–10 mm diam.; rhizome scales dark reddish brown with blackish tips, 1.5–2 ⫻ 0.5 mm wide, narrowly deltate, comose, entire; fronds 35–135 cm long, distant to clumped; stipes reddish brown, 1/5–1⁄2 the frond length, with scattered to dense acicular hairs 0.1–0.3 mm, ctenoid hairs conspicuous; blades (10–)15–27 cm wide, narrowly deltate, proximal pinnae not or only slightly reduced; rachises abaxially with numerous ctenoid hairs ca. 0.1 mm long, sometimes also with scattered septate hairs to 0.5 mm; pinnae 4–15 cm ⫻ 4–9 mm, obtuse to acute, straight to occasionally slightly deflexed, equilateral at bases, costae of proximal pinnae at right angles to the rachises; veins free, 2–3 forked; indument adaxially of sparse to moderately dense hairs 0.1–0.3 mm on costae, laminar surfaces glabrous or nearly so, abaxially the costae bearing ctenoid hairs like those of rachises, sometimes also with sparse septate hairs to 0.5 mm, laminar surfaces between veins with appressed hairs 0.1–0.2 mm, margins with occasional scattered hairs 0.3–0.5 mm; sori round, supramedial with few, clavate paraphyses 0.1–0.3 mm; sporangia glabrous; spores reniform, 64 per sporangium; 2n⫽74 (CR).

440

PECLUMA

short hairs 0.1 mm at the bases of costae, or absent, laminar surfaces with scattered hairs 0.3–0.5 mm, abaxially the costae with scattered hairs 0.5–1 mm, abaxial surfaces between veins with scattered appressed hairs 0.1 mm, marginal hairs numerous, septate, 0.4–1 mm; sori round, medial, with clavate paraphyses 0.5–0.8 mm; sporangia each with 2–5 hairs 0.1–0.2 mm long from capsules; spores reniform, 64 per sporangium; 2n⫽74 (Nay).

Distribution. Epiphytic, occasionally epipetric or terrestrial at bases of trees; 600–1600 m. Mexico; Guat, Bel, Hond, Nic, CR, Pan; Col, Ven, Ec, Galapagos, Peru, Bol. Selected Specimens Examined. Chis (Breedlove 32899, ENCB, 56483, 56519, 56567, ENCB, CAS; Palacios-Rios 2783, UC). Ver (Purpus s.n., DS, UC; Ventura A. 4952, ENCB, MICH, 11619, CHAPA, ENCB, XAL).

Pecluma divaricata is distinct in its brown stipes and the proximally truncate blades lacking greatly reduced pinnae. The pubescence along the rachises abaxially is peculiar, consisting of ctenoid hairs with tiny “teeth” less than 0.1 mm. Evans (1968) did not mention Polypodium divaricatum in his monograph of this group, although he annotated an isotype at BR as “Polypodium bolivianum?” in 1965. This is the largest species in the genus in Mexico, and differs from P. ptilodon by the blades being truncate proximally and the absence of a patch of erect hairs around the sori. Pecluma divaricata is probably most closely related to the widespread P. eurybasis (C. Chr.) M. G. Price, from Costa Rica to Bolivia and the Greater Antilles, and these two seemingly intergrade in portions of their common range in South America.

6. PECLUMA FERRUGINEA Pecluma ferruginea (M. Martens & Galeotti) M. G. Price, Amer. Fern J. 73: 114. 1983. Figs. 214N, O. Polypodium ferrugineum M. Martens & Galeotti, Me´m. Foug. Mexique 36, pl. 7, f. 2. 1842. Type. Mexico. Oaxaca: Forests of Zacatepec and Juquila, Galeotti 6354 (BR!, frag. US!; isotype K!).

Rhizomes short-creeping, 2–5 mm diam.; rhizome scales golden or orangish, 1–2 ⫻ 0.8–1 mm, ovate, acute, entire; fronds 8–40 cm long, clumped; stipes reddish brown, 1/6–2/5 the frond length, pilose with acicular hairs 0.1–0.3 mm; blades narrowly ovate, 2–6.5 cm wide, bases reduced, strongly deflexed; rachises abaxially light red-brown, with scattered, inconspicuous, linear to lanceolate scales 1.5–2 mm long, also with scattered to moderately dense septate hairs 0.5–1 mm; pinnae linear to narrowly ovate, acute, 1–2.5 cm ⫻ 2–3 mm, surcurrent in proximal half of blade, costae dark reddish brown to black, decurrent onto the rachises; veins free, simple or 1-forked; indument adaxially of

Distribution. Epipetric (rarely epiphytic), wooded ravine, pine and oak forests; 750–2800 m. Mexico; Guat. Selected Specimens Examined. Chis (Ventura & Lo´pez 3741, ENCB, MEXU, MO, XAL). Col (Lott et al. 2924, MEXU, NY, UC). Dgo (Seiler 1359, MO). Gro (Hinton 9607, NY, UC). Jal (Dı´az L. 8936, UC). Me´x (Hinton 4161, NY). Mich (Dı´az L. 9326, UC). Mor (Copeland herb. 125, MEXU, NY, UC, US). Nay (Bartholomew 2624, UC). Oax (Mickel 5554, MEXU, NY, UC, US). Sin (Hoover 348, UC). Unverified, Doubtful, or Mistaken Reports. Hgo (Copeland herb. 16056, MICH, cited by Evans, 1968, but not verified, and no other specimens have been seen from this area).

This is primarily a species of western and central Mexico. Among the brown-stiped species of Pecluma, P. ferruginea is distinct in its golden brown, non-comose rhizome scales, small fronds, and conspicuous capsular setulae; it also has persistent rachis scales abaxially. Copeland (1955: 440) discussed it his treatment of Ctenopteris, a genus belonging to the Grammitidaceae. Evans (1968: 227) cited a specimen (Barkelew 236, US! p.p.) from the Revillagigedo Islands as this species, but this specimen and a duplicate in UC are referable to P. alfredii var. occidentalis. Pecluma alfredii has been collected many times on Isla del Socorro, but P. ferruginea is not known from there.

7. PECLUMA HYGROMETRICA Pecluma hygrometrica (Splitg.) M. G. Price, Amer. Fern J. 73: 114. 1983. Figs. 215A–D. Polypodium hygrometricum Splitg., Tijdschr. Natuurl. Gesch. Physiol. 7: 409. 1840. Type. Surinam. Para, Splitgerber 1069 (L, photos BM!, C!).

Rhizomes short-creeping, 3–5 mm diam.; rhizome scales orangish to ferruginous, narrowly lanceolate, acuminate, incon-

PECLUMA PLUMULA

spicuously comose, margins minutely papillate or entire; fronds 10–50 cm long, clumped; stipes dark red-brown, 1⁄4–1/15 the frond length, ca. 1–1.5 mm diam., pilose with acicular, silvery hairs, ctenoid hairs absent; blades narrowly elliptic to lanceolate, 8–40 ⫻ 2–6 cm, subtruncate to truncate at bases; rachises abaxially lacking scales, with dense silvery hairs 0.1–0.5 mm; pinnae 10– 40 ⫻ 2–5 mm, symmetrically dilated at bases, obtuse to acutish at apices, costae decurrent onto rachises; veins free, 1-forked; indument adaxially of dense, silvery, acicular hairs, sometimes septate hairs on costae, veins, and laminar surfaces between veins, abaxially with similar and even denser hairs, appressed clavate hairs absent on the lamina between veins, margins with dense acicular hairs mostly 0.1–0.2 mm; sori round, medial; sporangia with 1–5 hairs on capsules; spores reniform, 64 per sporangium; 2n⫽74 (CR).

441

Polypodium pulchrum M. Martens & Galeotti var. minus E. Fourn., Mexic. Pl. 1: 76. 1872, nom. nud.

Rhizomes short-creeping, 3–7 mm diam.; rhizome scales reddish brown, dark towards margins, lanceolate, acuminate, margins toothed, papillose; fronds 8–60 cm long, clumped; stipes black, 1⁄8–1⁄3 the frond length, glabrous; blades narrow to linear elliptic, 3–11 cm wide, reduced to lobes at bases, the reduced lobes spreading, not deflexed; rachises abaxially with orangish to red-brown, ovate to lanceolate, sometimes bullate scales, margins toothed, papillate or fimbriate, also with a few septate hairs to 0.5 mm; pinnae linear-obtuse, 1.5–3(–6) cm ⫻ 1.5–3.5 mm, surcurrent and decurrent, costae black, decurrent onto rachises; veins free, mostly 1-forked; indument adaxially of sparse hairs 0.1– 0.3 mm on costae, laminar surfaces glabrous or nearly so, abaxially the costae with sparse septate hairs 0.3–0.5 mm and scattered bullate scales 0.5 mm long, laminar surfaces between veins with scattered appressed hairs, margins with a few hairs 0.2–0.5 mm; sori round, medial, with clavate paraphyses to 0.5 mm; sporangia with hairs 0.1 mm on capsules; spores reniform, 64 per sporangium; 2n⫽148 (Fla, Jam).

Distribution. Epipetric or epiphytic in seasonal evergreen forests; 200–400 m. Mexico; Guat, Hond, Nic, CR, Pan; Col, Ven, Guy, Sur, Ec, Peru, Bol. Specimens Examined. Chis (Breedlove 22524, 28505, DS, NY, 28576, DS; Matuda 16717, MEXU, 18116, MEXU, US, 18392, DS, ENCB, MEXU, US; Purpus 7226, NY, UC, US).

This differs from congeners in Mexico by its ferrugineous rhizome scales, proximally truncate to subtruncate blades, absence of scales on the rachises abaxially, relatively small fronds less than 50 cm long, and evenly pilose blades (short silvery hairs on axes as well as between the veins on both sides of the blades). Its affinities may be with species treated in Polypodium rather than to other species of Pecluma.

8. PECLUMA PLUMULA Pecluma plumula (Humb. & Bonpl. ex Willd.) M. G. Price, Amer. Fern J. 73: 115. 1983. Figs. 214E, F. Polypodium plumula Humb. & Bonpl. ex Willd., Sp. Pl., ed. 4, 5(1): 178. 1810. Ctenopteris plumula (Humb. & Bonpl. ex Willd.) J. Sm., Hist. Fil. 185. 1875. Lectotype (chosen by Evans, 1968: 230). Venezuela. Caracas, Bredemeyer s.n. (B-Willd. 19655-1!, photo BM, frag. NY!; isotype P-Humb.!). Polypodium pulchrum M. Martens & Galeotti, Me´m. Foug. Mexique 41, pl. 8, f. 2. 1842. Type. Mexico. Veracruz: Jalapa, Galeotti 6332 (BR!–photos UC!, US!; isotypes BR!–2 sheets, K–photo US!; frag. NY!).

Distribution. Epiphytic to terrestrial or epipetric, open rocky limestone slopes, pine-oak forests, oak forests, tropical evergreen forests; 150–1500(–2100) m. USA (Fla); Mexico; Guat, Bel, Hond, Salv (Villacorta et al. 872, MO), Nic, CR, Pan; Bah, Gr & L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol. Selected Specimens Examined. Camp (Collins 50, US). Chis (Breedlove 40159, DS). Hgo (Moore 1970, UC). Oax (Mickel 6115, ENCB, MEXU, NY, UC, US). Pue (Rzedowski 23374, DS). Qro (Rubio 2626, IEB). QR (Sanders 9840, UC). SLP (Rzedowski 27692, NY). Tam (Stanford et al. 985, ARIZ, DS, MEXU, NY, UC, US). Ver (Copeland herb. 126, BM, GH, MEXU, MICH, UC). Unverified, Doubtful, or Mistaken Reports. Gro (Urbina 46, FCME, cited by Lorea-Herna´ndez & Vela´zquez M., 1998, but not verified). Gto (Rzedowski 40576, IEB, cited by Dı´az-Barriga & PalaciosRios, 1992, but not verified). Mich (Arse`ne 6696, MEXU, and Rzedowski 40576, 42127, IEB, cited by Dı´az-Barriga & Palacios-Rios, 1992, but not verified). Sin (reported by Mickel & Beitel, 1988, probably on the basis of Dunn et al. 21813, ENCB, NY, but the specimen is P. alfredii, probably var. occidentalis).

442

PECLUMA

Pecluma plumula is distinct from congeners in Mexico by the combination of black stipes, lanceolate, toothed rhizome scales, proximal pinnae a little shortened but not deflexed, persistent lanceolate rachis scales, and setulose sporangia. Herna´ndez G. 755 (MEXU), cited by Mickel and Beitel (1988) as this species, is P. dispersa.

9. PECLUMA PTILODON VAR. BOURGEAUANA Pecluma ptilodon (Kunze) M. G. Price var. bourgeauana (E. Fourn.) A. R. Sm., comb. nov. Figs. 214G, H. Polypodium consimile Mett. var. bourgeauanum E. Fourn., Mexic. Pl. 1: 76. 1872 [“bourgaeanum”]. Syntypes. Mexico. Veracruz: In valle Cordobensi, Bourgeau 1431 (P–photo XAL, isosyntypes K!, L–photo UC!, NY!; frag. US!); Bourgeau 1436 (P; isosyntypes K–photo XAL, US!); Colombia. “Guadalupa,” Lindig 45 (P; isosyntype US!). Polypodium pectinatum L. var. caespitosum Jenman, Bull. Bot. Dept. 4: 125. 1897. Polypodium ptilodon (Kunze) M. G. Price var. caespitosum (Jenman) A. M. Evans, Amer. Fern J. 58: 170. 1968. Pecluma ptilodon (Kunze) M. G. Price var. caespitosa (Jenman) Lellinger, Proc. Biol. Soc. Wash. 98: 387. 1985. Type. Jamaica. Old England, Jenman s.n. (NY!).

Rhizomes short-creeping, 5–12 mm diam.; rhizome scales reddish brown with blackish tips, lustrous, 2–3 ⫻ 0.5 mm, narrowly deltate, comose, entire; fronds 30–105 cm long, clumped; stipes reddish brown to black, 1/15–1/7 the frond length, with scattered to dense acicular hairs 1 mm, ctenoid hairs conspicuous; blades 8–18 cm wide, narrowly deltate to ovate-deltate, gradually reduced to auricles proximally; rachises abaxially lacking scales, with moderately dense septate hairs to 1 mm, also with tiny appressed ctenoid hairs; pinnae 4–9 cm ⫻ 6–9 mm, obtuse to acute, straight to subfalcate, equilateral at bases or surcurrent, costae of proximal pinnae at right angles to rachises; veins free, 1–3(–4)-forked; indument adaxially of scattered septate hairs 0.1– 0.3 mm on costae, laminar surfaces glabrous, abaxially the costae sparsely to moderately hairy with septate hairs to 1 mm long, laminar surfaces between veins with scattered, appressed hairs 0.1 mm, also with short dense erect acicular hairs in an oblong patch around sori, margins with scattered hairs to ca. 0.5 mm; sori round or oblong, medial, with clavate, branched paraphyses 0.3 mm; sporangia usually lacking capsular hairs (Mexican specimens); spores reniform, 64 per sporangium; 2n⫽148 (SLP, Ver, Fla, Jam).

Distribution. Terrestrial, on fallen logs, or epipetric, slopes and steep ravines in seasonal evergreen forests, montane rain forests, pine-oak forests; 200–1500 m. USA (Fla), Mexico; Hond; Cuba, Jam, Hisp; Trin. Reported from Guatemala by Stolze (1981) and from Nicaragua by Go´mez (1976), as Polypodium ptilodon var. caespitosum. Selected Specimens Examined. Chis (Breedlove 21870, 22181, DS, NY). Hgo (Sa´nchez S. 442, US). Oax (Mickel 4819, ENCB, NY, UC, US). Pue (Ventura A. 21737, IEB, MEXU). Qro (Rzedowski 48179, IEB). SLP (Pringle 3974, ARIZ, DS, GH, MEXU, MO, NY, UC, US). Tam (Palmer 306, US). Ver (Ventura A. 9512, ASU, CAS, XAL, 12234, CHAPA, ENCB, IEB, XAL). Unverified, Doubtful, or Mistaken Reports. Mich (Arse`ne 5253, MEXU, cited by Dı´az-Barriga & Palacios-Rios, 1992, but not verified). Tab (reported by Magan˜a, 1992, as var. caespitosa, but not verified).

Pecluma ptilodon is unique among Mexican species in its oblong patch of erect hairs around the sori. In Mexico, it is likely to be confused only with P. consimilis, which see for discussion. Other distinguishing characteristics are the gradually reduced proximal pinnae (to mere semicircular lobes), lack of scales and presence of ctenoid hairs on the rachises abaxially, and the broad blades and broad pinnae with 2–3-forked veins. Breedlove 20964 and 27886, from Chiapas, cited as this species by Smith (1981), are P. consimilis. Variety ptilodon, from Peru and Bolivia, differs by having slightly longer stipes, larger fronds, smaller spores, and 2n⫽74. Two other South American varieties recognized by Evans (1968) appear more distantly related and possibly not conspecific.

10. PECLUMA SURSUMCURRENS Pecluma sursumcurrens (Copel.) M. G. Price, Amer. Fern J. 73: 115. 1983. Figs. 215E, F, K. Polypodium sursumcurrens Copel., Univ. Calif. Publ. Bot. 19: 291. 1941. Type. Mexico. Veracruz: Head of Orizaba valley, Copeland herb. 128 (MICH, photo UC!; isotypes BM, GH!, UC!, US!). Polypodium cyathicola Copel., Univ. Calif. Publ. Bot. 19: 292, t. 43. 1941. Type. Mexico. Veracruz: Cuantlancillo (Orizaba), Copeland herb. 127 (MICH!, photo UC!; isotypes NY!, UC!, US!).

PELLAEA

Rhizomes short-creeping, 3–5 cm diam.; rhizome scales brown, 1.5–2.5 ⫻ 1 mm, ovate, acute, entire, often somewhat comose; fronds 20–55 cm long, clumped; stipes blackish with brown lateral stripes or stipes dark reddish brown, 1/15–1/5 the frond length, with scattered hairs 0.5–0.8 mm; blades narrowly oblong, 5–8 cm wide, proximal pinnae reduced, deflexed; rachises abaxially lacking scales or with sparse brown, linear-lanceolate scales 0.8–1 mm long, also with moderately dense hairs 0.5–0.8 mm; pinnae linear to narrowly lanceolate, acute, 2.5–4 cm ⫻ 2–5 mm, basiscopic edge of each pinna perpendicular to the rachis, surcurrent in proximal blade half, costae usually decurrent onto rachises, sometimes at right angles; veins free, 1-forked; indument adaxially of sparse to moderate hairs 0.1–0.3 mm on costae, laminar surfaces glabrous or nearly so, abaxially the costae with sparse to moderate hairs 0.3–0.5 mm, laminar surfaces between veins with scattered, appressed, clavate hairs 0.1–0.2 mm, margins with scattered hairs 0.5–0.8 mm; sori round, with clavate paraphyses 0.3–0.5 mm; sporangia lacking capsular hairs; spores reniform, 64 per sporangium; 2n⫽74 (Oax).

443

lowed suit, e.g., Smith (1981), Mickel and Beitel (1988), and Palacios-Rios (1992). However, we now believe that the type of P. cyathicola is conspecific with the type of P. sursumcurrens, the latter differing from the former primarily in having larger fronds, submarginal sori, and more numerous, twisted soral paraphyses that are enlarged subapically (abortive sporangia?). This last characteristic is found in varying degrees in many specimens here regarded as P. sursumcurrens. Pecluma sursumcurrens differs from P. alfredii in absence of scales on the rachises abaxially, or if scales are present, they are usually very sparse and linear-lanceolate (rather than ovatecordate). In addition, the rhizome scales are usually somewhat comose, more acute at the tips, and often darker brown, and the stipe bases have numerous long, twisted, multicellular hairs ca. 1 mm. The pinna margins are always hairy, in contrast to some specimens of P. alfredii (including the type) and its var. cupreolepis (including the type). The position of the sori varies from submarginal to medial, and there are often numerous twisted paraphyses with a multicellular, clavate head.

92. PELLAEA Pellaea Link, Fil. Spec. 59. 1841, nom. cons. Lectotype (first chosen by Christensen, Index Filic. XXXIX. 1906): Pellaea atropurpurea (L.) Link [⬅ Pteris atropurpurea L.]. (See Pic. Serm., Webbia 9: 428. 1953.)

Distribution. Epiphytic in lowland and montane forests, cloud forests; 100–1600(–2600) m. Mexico; Guat. Selected Specimens Examined. Chis (Breedlove 22975, MEXU, 31331, DS, NY). Hgo (Rzedowski 30216, MEXU). Oax (Croat & Hannon 65839, UC; Mickel 1613, ENCB, UC). Pue (Riba et al. 266, MEXU, NY). Qro (Servı´n O. 875, 1430, IEB). SLP (King 4354, NY). Tab (Bourgeau 3613, UC). Ver (Croat 39539, UC; Ventura A. 13711, CAS, MEXU).

Evans circumscribed Polypodium sursumcurrens in an extremely narrow sense, and cited only the type and two additional specimens as belonging to this species: Pue (Honey Station, Pringle 13475, MICH), and Ver (Sharp 44822, US). One of these collections, Pringle 13475 (US), was also cited by Evans (1968) as P. cupreolepis (treated herein as Pecluma alfredii var. cupreolepis), with provenance “Hidalgo, Trinidad.” Both the type of Pecluma sursumcurrens and Sharp 44822 have the hairs abaxially infected with a black pyrenomycete fungus. This same fungus is sometimes found on many other specimens of P. sursumcurrens in varying degrees (including type material of Polypodium cyathicola), as well as other species of Pecluma in Mexico. Evans (1968) placed Polypodium cyathicola in synonymy under P. alfredii, which see for further discussion. Others have fol-

Terrestrial; rhizomes compact to long-creeping; rhizome scales linear, usually toothed, concolorous or bicolorous; fronds medium-sized, clumped to distant; blades 1–3-pinnate, naked to pubescent; veins free (rarely anastomosing); sori marginal or rarely elongate along the veins, the blade margins recurved to protect the sporangia; spores tetrahedral-globose, yellow to brown, smooth or rugose; x⫽29. Pellaea is a genus of about 35 species of temperate and subtropical dry, rocky regions. There is a major problem with the generic circumscription, and there has been confusion in the past, with many things being put into Pellaea that belong elsewhere. Pellaea allosuroides (Mett.) Hieron. is a Cheilanthes, P. rigida (Sw.) Hook. a Cheiloplecton, and P. intramarginalis (Kaulf. ex Link) J. Sm. a Mildella. Pellaea skinneri Hook., P. lozanoi Maxon, and P. seemannii Hook. have usually been placed here, but these species are more closely related to Cheilanthes. It is difficult to circumscribe Pellaea precisely. In some respects Pellaea resembles Cheilanthes and Argyrochosma, but generally has pale frond axes, or if dark, then with glabrous blades. Molecular data (Gastony & Rollo, 1995, 1998) suggest that Pellaea is sister to Astrolepis and Argyrochosma). References Gastony, G. J. & L. D. Gottlieb. 1985. Genetic variation in the homosporous fern Pellaea andromedifolia. Amer. J. Bot. 72: 257–267; Gastony, G. J. & M. D. Windham. 1989. Species concepts in pteridophytes: the treatment and definition of agamosporous species. Amer. Fern J. 79: 65–77; Mendoza, A., M. Windham, B. Pe´rez-Garcı´a & G. Yatskievych. 2001. Una nueva especie de Pellaea (Pteri-

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PELLAEA

daceae) del estado de San Luis Potosı´, Me´xico. Acta Bot. Mex. 57: 15–21; Tryon, A. F. 1957. A revision of the fern genus Pellaea section Pellaea. Ann. Missouri Bot. Gard. 44: 125–193; Tryon, A. F. 1968. Comparisons of sexual and apoga-

mous races in the fern genus Pellaea. Rhodora 70: 1–24; Tryon, A. F. 1972. Spores, chromosomes, and relations of the fern Pellaea atropurpurea. Rhodora 74: 220–241.

Key to the Mexican Species of Pellaea 1. Blades 1-pinnate; rachises and laminae abaxially densely lanose. .................................................................................................................... 10. P. ribae. 1. Blades 2–3-pinnate, rarely 1-pinnate; rachises and laminae abaxially glabrous or with scattered lax hairs. 2. Stipes brown to black; spores tan (brown in P. wrightiana). 3. Rhizome scales concolorous, tortuous with occasional teeth; stipes and rachises essentially terete with many long (1–2 mm) and short (0.1 mm) hairs; proximal pinnae and their terminal pinnules on long (15 mm) dark stalks; segments obtuse to acute at apices, not mucronate. ...................................................................................................................................................... 2. P. atropurpurea. 3. Rhizome scales bicolorous, each with a thick sclerotic central stripe; stipes and rachises sulcate and glabrous; proximal pinnae and their pinnules on short colorless stalks; segments mucronate at apices, each with colorless tip 1 mm long. 4. Segments pinnate; costae of proximal pinnae to 7 cm long. 5. Blades 2-pinnate. .................................................................................................................................................................... 13. P. truncata. 5. Blades 3-pinnate, the larger and basal pinnules mostly 3-parted. ................................................................................. 5. P. mucronata. 4. Segments 3-parted, costae of proximal pinnae 0–2 cm long. 6. Costae present; pinnae pinnate. ....................................................................................................................................... 15. P. wrightiana. 6. Costae absent; pinnae ternate. 7. Blades partially 1-pinnate, ternate pinnae in proximal 1⁄4–3⁄4 of blades; rachises with lax hairs. ............................ 14. P. villosa. 7. Blades virtually all with ternate pinnae; rachises glabrous. ...................................................................................... 12. P. ternifolia. 2. Stipes stramineous or tan; spores brown. 8. Blade axes strongly to slightly flexuous; pinnule axes densely to sparsely hairy with short reddish hairs 0.1–0.3 mm long; rhizome scales bicolorous with dark center; pinnae subcoriaceous, veins not apparent; blades 1–3-pinnate; epidermal cells not whitened on recurved margins. 9. Blade axes strongly flexuous; costae densely hairy; blades 2–3-pinnate. ...................................................................................... 8. P. ovata. 9. Blade axes slightly flexuous; costae sparsely hairy; blades 1–2-pinnate. ................................................................................. 7. P. oaxacana. 8. Blade axes straight; pinnule axes glabrous except for scattered hairs near pinnule bases; rhizome scales concolorous, light brown; pinnae thin, veins evident; blades 1–2-pinnate; epidermal cells whitened on recurved margins. 10. Segments strongly hastate or palmately lobed. ............................................................................................................................ 9. P. pringlei. 10. Segments round or oblong, not hastate. 11. Rhizomes stout and compact, over 5 mm diam. 12. Blades once-pinnate. ........................................................................................................................................................ 6. P. notabilis. 12. Blades bipinnate to tripinnate. 13. Segments rotundate-cordate; stipes and rachises usually glabrous; rhizome scales 5–6 mm long, tortuous; recurved margins with occasional hairs; terminal pinnules with equilateral bases. ....................................... 3. P. cordifolia. 13. Segments narrowly deltate to sagittate; stipes and rachises with scattered short hairs near pinnule base; rhizome scales straight, 3–4 mm; recurved margins glabrous; terminal pinnules with inequilateral bases (1–1.5 mm difference). ............................................................................................................................................ 11. P. sagittata. 11. Rhizomes slender, long-creeping, under 5 mm diam., rhizome scales bicolorous. 14. Veins obscure; n. Mexico (Son to Tam and Zac). ................................................................................................... 4. P. intermedia. 14. Veins visible; BCN, Cedros Isl, NL. .................................................................................................................... 1. P. andromedifolia.

1. PELLAEA ANDROMEDIFOLIA Pellaea andromedifolia (Kaulf.) Fe´e, Me´m. Foug. 5: 129. 1852. Figs. 217G, H, J, K. Pteris andromedifolia Kaulf., Enum. Filic. 188. 1824. Type. U.S.A. California: Chamisso s.n. (P, photo GH, US). Pellaea andromedifolia (Kaulf.) Fe´e var. pubescens D. C. Eaton, Ferns N. Amer. 1: 205. 1878. Type. Mexico. [Nuevo Leo´n:] Monterrey, Nuttall s.n. (K).

Rhizomes long-creeping, 2–4 mm diam.; rhizome scales linearlanceolate, bicolorous, black with brown margins, irregularly

dentate; fronds 10–60 cm long, 0.5–1 cm distant; stipes ca. 1⁄2 the frond length, stramineous to gray, terete or flattened adaxially, dull, glabrous, hairy or scaly; blades elongate-deltate, 2–3pinnate, to 20 cm wide; pinnae ascending or perpendicular to rachises, costae straight; ultimate segments elliptic to ovate, truncate to subcordate at bases, rounded apices, 3–15 mm, herbaceous, veins visible, margins recurved on fertile segments; surfaces glabrous to sparsely pubescent; sporangia 64- or 32-spored; 2n⫽58 (USA).

PELLAEA CORDIFOLIA

445

Distribution. Rocky slopes and ledges, usually on igneous substrates; 0–500(–1100) m. USA (Calif, Ore); Mexico.

Distribution. Limestone cliffs; 1500–2500 m. Canada, USA; Mexico; Guat.

Selected Specimens Examined. BCN (Atwood 20667, NY; Carter et al. 1047, UC; Ceta & Salgado 7428, ENCB; Montgomery 83-20, 83-33, NY; Pray 16645, NY; Shreve 6834, MICH; Wiggins 5152, 9967, US). Cedros Island (Palmer 749 1⁄2, NY).

Selected Specimens Examined. Chih (Soderstrom 887, LL). Chis (Breedlove 37179, DS, NY). Coah (Stanford et al. 391, US). Jal (Iltis et al. 29515, WIS). NL (Mueller & Mueller 362, US). Oax (Mickel 6314, NY). Pue (Arse`ne 3548, GH). SLP (Burgess 5406, ARIZ). Son (Hartman 357, NY, US). Tam (Stanford et al. 943, US). Ver (Bourgeau 3068, NY). Zac (Johnston et al. 11565A, LL p.p.).

According to Windham (in FNA Ed. Comm., 1993), Pellaea andromedifolia comprises three cytotypes: a sexually reproducing diploid, an apogamous triploid, and an apogamous tetraploid. The apogamous tetraploid apparently resulted from hybridization between diploid and triploid individuals (Gastony & Gottlieb, 1985).

2. PELLAEA ATROPURPUREA Pellaea atropurpurea (L.) Link, Fil. Spec. 59. 1841. Figs. 218A–H, J. Pteris atropurpurea L., Sp. Pl. 2: 1076. 1753. Allosorus atropurpureus (L.) Kunze ex C. Presl, Suppl. Tent. Pterid. 153. 1836. Platyloma atropurpurea (L.) J. Sm., J. Bot. (Hooker) 4: 160. 1841. Notholaena atropurpurea (L.) Keyserl., Polyp. Herb. Bunge. 30. 1873. Type. U.S.A. Virginia: Kalm s.n. (LINN 1246.18). [A. Tryon (1957) cited Rappahanock River, Clayton 682 (BM) as holotype]. For additional synonymy, see A. Tryon (1957).

Rhizomes short, compact, to 8 mm diam.; rhizome scales 5–6 mm, linear, dark orange, concolorous, dull, subentire with occasional teeth, tortuous; fronds to 50 cm long, subdimorphic; stipes 1⁄4–1⁄2 the frond length, terete, atropurpureous, pubescent with scattered white hairs of two lengths,1–2 mm long and 0.1 mm long; blades elongate-deltate, to 25 ⫻ 18 cm, 2-pinnate or rarely 1-pinnate, gray-green; pinnae petiolate to 15 mm, articulate, pinnule stalks dark, stopping at bases of pinnules, apices acute to rounded, distal pinnae entire, proximal ones pinnate, slightly pubescent along abaxial costae with 1–2 mm long hairs, 1 to several pairs of pinnules, ultimate segments linear, entire, subcoriaceous, ultimate pinnules on proximal pinnae 30–40 ⫻ 4 mm; fertile margins recurved, thin at edges, indusia 0.1 mm with entire margins, sori elongate about 2 mm along ends; spores tan; n⫽2n⫽87 (Canada, USA, NL). ⫻ 0.3

Unverified, Doubtful, or Mistaken Reports. Ags (Gonza´lezAdame 77, HUAA, cited by Siqueiros-Delgado & Gonza´lez-Adame, 2004, but not verified).

Pellaea atropurpurea is distinct by its dark axes, pinnate pinnae, and concolorous rhizome scales. There is considerable variation within Pellaea atropurpurea, and perhaps some of the atypical specimens reflect hybridization with P. notabilis. Windham (pers. comm.) has suggested that Pellaea atropurpurea has crossed with P. notabilis, the hybrid appearing much like the former species, but distinct in having the rachises ciliate abaxially (Fig. 218H), whereas P. atropurpurea has the rachises glabrous abaxially (though ciliate adaxially) (Fig. 218C). Gastony and Windham (1989) reported a tetraploid apogamous plant, 2n⫽116, that is a probable hybrid between triploid P. atropurpurea and diploid P. notabilis.

3. PELLAEA CORDIFOLIA Pellaea cordifolia (Sesse´ & Moc¸.) A. R. Sm., Amer. Fern J. 70: 26. 1980. Fig. 216D. Adiantum cordifolium Sesse´ & Moc¸., Naturaleza (Mexico City), ser. 2, 1(Append.): 182. 1890. Type. Mexico. [Distrito Federal:] Cuyuaca´n [Coyoaca´n] and San Agustı´n near Mexico City (MA). Pteris cordata Cav., Descr. Pl. 267. 1801. Pellaea cordata (Cav.) J. Sm., Cult. Ferns 4. 1857, hom. illeg., non Fe´e, 1852. Pellaea cardiomorpha Weath., J. Arnold Arbor. 24: 309. 1943. Pellaea sagittata (Cav.) Link var. cordata (Cav.) A. F. Tryon, Ann. Missouri Bot. Gard. 44: 166. 1957. Type. Mexico. [Morelos?:] Chapultepec, Ne´e s.n. (MA; isotype F). For additional synonymy, see A. Tryon (1957).

Rhizomes short, stout, compact; rhizome scales 5–6 ⫻ 0.3–0.8 mm, dull tan to reddish brown, linear-lanceolate, irregularly denticulate, tortuous; fronds to 30 cm long, erect; stipes 1⁄4–1⁄2

446

PELLAEA

the frond length, nearly terete to plane adaxially, stramineous and glistening, glabrous or rarely sparsely pubescent; rachises and segment stalks essentially glabrous, with occasional short hairs on pinnule stalks; blades narrowly oblong to deltate, 2-pinnate, light green, yellow-green to gray-green; pinnae widely spreading to slightly ascending, segments round, ca. 12 ⫻ 9–11 mm, bases cordate, terminal pinnule on each pinnae with equilateral base, pinnae at broader angle to rachises than in P. sagittata, with only a few hairs abaxially on veins and costae near terminal segments; modified indusia 0.1–0.3 mm wide, entire, occasional hairs along recurved margin, cells along recurved margins whitened; spores brown; 2n⫽58 (DF).

pubescent; pinnae at right angles to rachises, segments 8–17 mm long, short-petiolate, ovate or elliptic, articulate, truncate or cordate or rounded at base, obtuse to mucronate at apices, costae straight to slightly flexuous; adaxial surfaces glabrous; abaxial surfaces glabrous or pubescent; sori not continuous along margins, interrupted at apices, margins reflexed, narrow, hyaline; sporangia 32-spored; n⫽2n⫽87, 116, apogamous (USA).

Distribution. Rocky slopes and ledges, on a variety of substrates, including limestone and igneous rocks; 1900–2850 m. USA (Ariz, N Mex, Tex); Mexico.

Distribution. Dry, rocky slopes or basaltic cliffs in oak forests; 1900–3050 m. Sw USA (Tex); Mexico. Selected Specimens Examined. Ags (Rzedowski & McVaugh 862, MICH). Chih (Pringle 448, NY). Chis (Purpus 7248, NY). Coah (Wynd & Mueller 590, NY). DF (Matuda 18828, US). Dgo (Reveal 3025, NY). Gro (Lorea 4378, FCME). Gto (Kenoyer 1787, US). Hgo (Rose & Painter 6741, NY). Jal (McVaugh 16775, MICH). Me´x (Rose & Painter 6825, US). Mich (Arse`ne 2970, US). Mor (Fisher 35417, NY). NL (Mueller & Mueller 970, MEXU). Oax (Mickel 4091, NY). Pue (Purpus 2712, NY). Qro (Rose & Rose 11196, NY). SLP (Parry & Palmer 980, NY). Tlax (Langman 2589, US). Ver (Chazaro B. et al. 6094, XAL). Zac (Taylor & Taylor 6022, NY).

Pellaea cordifolia is distinct from P. sagittata in its roundcordate segments.

Selected Specimens Examined. Chih (Pringle 461, NY, 930, LL, NY). Coah (Hinton 16700, ARIZ, US; Palmer “1426 & 1427,” NY). NL (Correll & Johnston 19922, NY; Hinton 18100, IEB). Son (White 3303, ARIZ). Tam (Breedlove 63463, CAS). Zac (Johnston et al. 11565A, LL p.p.; Lloyd & Kirkwood 22, GH). Unverified, Doubtful, or Mistaken Reports. Ags (reported by de la Cerda Lemus & Siqueiros Delgado, 1985, but not verified).

Plants of Pellaea intermedia in Mexico are mostly apogamous triploids and tetraploids; a sexual diploid has been found near Saltillo, Mexico (A. Tryon, 1968). Given the morphological similarity among the three cytotypes, the North American polyploids probably were derived from the Mexican diploid through autopolyploidy. From Pellaea andromedifolia it differs in obscure (vs. visible) veins, and its distribution in northern Mexico (Sonora to Tamaulipas and Zacatecas vs. Baja California Norte).

4. PELLAEA INTERMEDIA Pellaea intermedia Mett. ex Kuhn, Linnaea 36: 84. 1869. Type. Mexico. “Herb. Fournier” (B). Figs. 217L–O. Pellaea intermedia var. pubescens Mett. ex Kuhn, Linnaea 38: 85. 1869. Type. U.S.A. New Mexico: Wright 1190 (label no. 825) (B; isotypes GH!, MO).

Rhizomes long-creeping, 1.5–2 mm diam.; rhizome scales linear-lanceolate, bicolorous, black with light brown margins, finely serrulate, 2–4 ⫻ 0.3–0.8 mm; fronds to 50 cm long, 0.5–1 cm distant; stipes 1⁄2–3⁄5 the frond length, tan to gray to stramineous, adaxially flattened or terete, glabrous, dull; blades deltateovate to ovate-lanceolate, mostly 2-pinnate, occasionally 3pinnate, 4–20 cm wide, subcoriaceous; rachises straight and

5. PELLAEA MUCRONATA Pellaea mucronata (D. C. Eaton) D. C. Eaton in Emory, Rep. U.S. Mex. Bound. 2(1): 233. 1859. Figs. 218K, L. Allosorus mucronatus D. C. Eaton, Amer. J. Sci. Arts 22: 138. 1856. Type. U.S.A. California: near San Francisco Bay, A. B. Eaton s.n. (YU!, photo GH). Pellaea ornithopus Hook., Sp. Fil. 2: 142, t. 116A. 1858. Type. Mexico. Hartweg 2042 (K; isotypes GH, P). Pellaea longimucronata Hook., Sp. Fil. 2: 143. 1859, nom. illeg. since Hooker included the type of Allosorus mucronatus D. C. Eaton. Pellaea wrightiana Hook. var. longimucronata (Hook.) Davenp., Cat. Dav. Herb., Suppl., 46. 1883. Type. U.S.A. New Mexico: Wright 2131 (K!, photo GH; isotypes GH, US!, YU!).

PELLAEA OAXACANA

447

Rhizomes compact, ascending, 5–10 mm diam.; rhizome scales linear-subulate, 0.1–0.3 mm wide at bases, bicolorous, black, with brown, erose-dentate margins; fronds to 45 cm long, clumped; stipes 1⁄2–2⁄3 the frond length, dark brown, red brown, or purplish black, lustrous, glabrous, adaxially flattened to shallowly grooved; blades ovate-deltate to ovate-lanceolate, (2–) 3pinnate, 4–11(–18) cm wide, glabrous; pinnae perpendicular to rachises; ultimate segments narrowly oblong, often folded lengthwise in half, pinnules divided into 3’s, segments 2–12 mm long, bases rounded, apices mucronate, coriaceous, glabrous, margins recurved on fertile segments, covering more than half the abaxial surface; adaxial and abaxial surfaces glabrous; sori marginal; sporangia 64-spored, mixed with farina; 2n⫽58 (USA). Distribution. On limestone, bushy hillsides; 320–1100 m. Mexico. Specimens Examined. NL (Hinton 20008, MEXU; Knobloch 1963, GH; Sa´nchez 316, US, 613, 619, MEXU). Tam (Bartlett 10352, US; Kemp, in 1902, NY; Stanford et al. 2129 GH; Yatskievych 83-319, UC).

The long, slender, undivided pinnae and stramineous frond axes are distinct for this species.

7. PELLAEA OAXACANA

Distribution. Cliffs and rocky slopes, on a variety of acidic to mildly basic substrates; 0–2300 m. USA (Calif, Nev); Mexico. Selected Specimens Examined. BCN (Carter 1046, MEXU; Franklin & Chandler 5688, NY; Montgomery 83-36, NY; Taylor & Taylor 15606, NY; Wiggins 9821, US; Wiggins & Gillespie 4151, GH, MEXU, MICH, MO, US). Guad (Moran 6166, UC; Palmer 100, GH, MO, NY, YU). Son (Felger 87-42, ARIZ, MEXU, TEX; Palmer 554, US).

In Mexico the blades are tripinnate proximally and the pinnae usually Ⳳ perpendicular to the rachis and not overlapping (vs. bipinnate with the pinnae ascending and overlapping in subsp. californica (Lemmon) Windham of California).

6. PELLAEA NOTABILIS Pellaea notabilis Maxon, Contr. U.S. Natl. Herb. 10: 500. 1908. Type. Mexico. Tamaulipas: near Victoria, Palmer 234 (US!). Figs. 217A–C. Rhizomes compact; rhizome scales appressed, orange-tan, concolorous, serrulate, attenuate; fronds 12–43 cm, clumped, monomorphic; stipes ca. 1⁄3 the frond length, terete, stramineous, pubescent; rachises straight; blades 1-pinnate, 3.5–14 cm wide; pinnae 4–7 pairs, linear-lanceolate, entire, 3–7 ⫻ 0.7–1 cm, subcoriaceous, glabrous; veins obscure; spores 64 per sporangium, tan; 2n⫽58 (NL, Tam).

Pellaea oaxacana Mickel & Beitel, Mem. New York Bot. Gard. 46: 271. 1988. Type. Mexico. Oaxaca: Distrito Sola de Vega, 35–41 km S of Sola de Vega, 15–21 km N of bridge at Juchatengo, Mickel 6279 (NY!). Fig. 216B. Rhizomes creeping, slender; rhizome scales 2–3 ⫻ 0.5–0.8 mm, bicolorous, central portion dark reddish brown, margins wide, light brown, erose-denticulate; fronds to 45 cm long; stipes ca. 2/5 the frond length, stramineous, convex to plane adaxially, glabrous; rachises straight to slightly flexuous, costae and costules glabrous except for a few short hairs 0.1–0.3 mm on adaxial surfaces near base of segments; blades deltate, pinnate to bipinnate; pinnae slightly deflexed, segments ovate to sagittate, 10–20 ⫻ 10–15 mm, bases cordate, apices obtuse, subcoriaceous, glabrous, terminal pinnule with equilateral base; veins obscure or nearly so; margins recurved, nearly entire, Ⳳ differentiated, indusia 0.1 mm wide; spores brown.

448

PELLAEA

Distribution. Dry, rocky slopes; 800–1750 m. Mexico. Selected Specimens Examined. Gro (Lorea 794, FCME). Jal (Palmer 731, US). Me´x (Tejero-Dı´ez 2606, IZTA). Mich (Nelson 6546, US). Mor (Palacios Vargas 47, FCME). Oax (Mickel 3846, 4947, 6251, NY). Nay (Te´llez 9330, MEXU). Qro (Arse`ne 10649, US; Rose 11195, US).

Although intermediate in morphology between P. ovata and P. sagittata, P. oaxacana appears to be a fertile species. It is distinguished from P. ovata by the pinnae being only slightly flexuous and reflexed, pinnule tips somewhat acute, costae and costules only sparsely hairy, and fronds 1–2-pinnate. From P. sagittata, P. oaxacana differs by the bicolorous rhizome scales, slightly flexuous and retroflexed pinnae, terminal pinnules with equilateral bases, and papillate indusia.

8. PELLAEA OVATA Pellaea ovata (Desv.) Weath., Contr. Gray Herb. 114: 34. 1936. Fig. 216C. Pteris ovata Desv., Me´m. Soc. Linn. Paris 6: 301. 1827. Type. Peru. s. coll. s.n. (P-Desv., not found, photos GH, UC). Pteris flexuosa Kaulf. ex Schltdl. & Cham., Linnaea 5: 614. 1830. Allosorus flexuosus (Schltdl. & Cham.) Kunze, Linnaea 13: 136. 1839. Pellaea flexuosa (Schltdl. & Cham.) Link, Fil. Spec. 60. 1841. Platyloma flexuosa (Schltdl. & Cham.) J. Sm., Bot. Mag. 72(Compendium): 21. 1846. Type. Mexico. [Veracruz:] Near Jalapa, Schiede & Deppe s.n., Aug 1828 (B; isotype LE!).

Rhizomes short- to moderately creeping, slender, ca. 2 mm diam.; rhizome scales 2–3 ⫻ 0.5–0.8 mm, bicolorous, central portion lustrous black, margins thin, light brown, erose, tip hairlike; fronds to 60 cm long; stipes ca. 1⁄3 the frond length, stramineous, plane on top, glabrous to sparsely hairy with scattered black specks, rachises flexuous, costae and costules densely short-pubescent with reddish hairs 0.1–0.3 mm long; blades deltate, 2–3-pinnate, the pinnae retroflexed, segments elliptic to ovate, glabrous, articulate, bases cordate, apices obtuse to truncate, 8–12 ⫻ 5–10 mm, terminal pinnule of each pinnae with base equilateral; veins obscure, subcoriaceous; margins recurved with narrow modified entire indusia 0.1 mm wide; spores brown; 2n⫽58 (Tex, CR), n⫽2n⫽87, apogamous (SLP).

Distribution. Dry, rocky slopes and moist oak forests; 900– 2550 m. USA (Tex); Mexico; Guat, Hond, Nic, CR; Hisp; Col, Ven, Ec, Peru, Bol, Arg. Selected Specimens Examined. Ags (McVaugh & Koelz 123, MICH). BCS (Leo´n 3415, UC). Chih (Gentry 1538, UC). Chis (Ghiesbreght 204, US). Coah (Wynd & Mueller 318, US). DF (Lyonnet 861, NY). Dgo (Ortega 5313, US). Gro (Hinton 11305, US). Gto (Duges s.n., in 1880, NY). Hgo (Chase 7217, F, MO). Jal (McVaugh 17408, MICH). Me´x (Hinton 7206, MO). Mich (Arse`ne 3645, US). Mor (Rose et al. 10193, US). Nay (McVaugh 16337, MICH). NL (Mueller & Mueller 1130, GH). Oax (Pringle 5951, GH). Pue (Copeland herb. 108, UC). Qro (Tryon & Tryon 5154, MO). Sin (Breedlove 18048, NY). SLP (Rzedowski 24706, NY). Son (Rose et al. 13113, US). Tam (Bartlett 10313, US). Tlax (Acosta P. 2657, ENCB). Ver (Bourgeau 3067, NY). Zac (McVaugh 17783, MICH).

Pellaea ovata is distinct by its strongly flexuous blade axes and densely hairy costae. There are two forms of this species, an apogamous 32-spored form and a sexual 64-spored form.

9. PELLAEA PRINGLEI Pellaea pringlei Davenp., Gard. & Forest 4: 555. 1891. Type. Mexico. Jalisco: near Guadalajara, Pringle 2591 (GH!; isotypes F, GH!, MEXU!, NY!, P!, UC!, US!, YU!). Figs. 217D–F. Rhizomes short, stout, compact; rhizome scales 5–6 ⫻ 0.1–0.2 mm, dull tan to orange, linear-lanceolate, with hair-like tips, entire; fronds 14–38 cm long, erect; stipes 3⁄5–3⁄4 the frond length, plane adaxially, stramineous and lustrous, with scattered hairlike scales near bases; blades narrowly oblong to narrowly deltate, 1-pinnate, bluish green; pinnae strongly hastate, with 3–5 irregular lobes, 1.5–5.2 ⫻ 1.9–4.8 cm; stalks of the rachises and pinnae with sparse hair-like scales to glabrous; laminae glabrous; indusia ca. 1 mm wide, entire, glabrous; spores brown.

PELLAEA SAGITTATA

Distribution. Terrestrial in tropical deciduous forests or oakpine forests, sometimes in calcareous soils; 1300–2000 m. Mexico. Selected Specimens Examined. Col (McVaugh 16074, MICH). Gro (Hinton 9475, NY). Jal (Pringle 11793, MICH). Me´x (Hinton 1538, NY). Mich (Pe´rez & Zamudio 2612, IEB). Mor (Lyonnet 2563, MEXU; Pringle 7081, US, 7947, NY). Nay (Mexia 709, US). Sin (Gentry 6523, GH, NY).

Pellaea pringlei is distinct by its strongly hastate pinnae.

10. PELLAEA RIBAE Pellaea ribae Mendoza & Windham, Acta Bot. Mex. 57: 15. 2001. Type. Mexico. San Luis Potosı´: San Nicolas Tolentino, 3 km antes de Puerto Sta. Gertrudis, viniendo de Buenavista, Mendoza 250 (UAMIZ!; isotypes MO, NY!, UAMIZ!, UT!). Figs. 219A–C. Rhizomes compact, horizontal to ascending, 3–5 mm diam.; rhizome scales linear, concolorous orange-tan, somewhat flexuous at apices, entire, 1–1.3 ⫻ 0.1–0.2 mm; fronds monomorphic, 6– 20 cm long, clumped; stipes 1/6–1⁄4 the frond length, black, lustrous, terete, densely lanose with white hairs of two lengths, dense curled hairs 1–2 mm long and straight ca. 5–6 mm long; blades linear-oblong, 1-pinnate, 1.5–3 cm wide; rachises straight; pinnae 8–28 pairs, mostly deflexed, increasingly so proximally, distally perpendicular to rachises, linear-oblong, 6–16 mm long, sterile ca. 5 mm wide, fertile ca. 2 mm wide, incurved, bases subcordate, apices rounded, coriaceous; adaxial surfaces glabrous; abaxial surfaces densely lanose; pinna margins undifferentiated, curved, protecting sori; spores brown.

449

Distribution. Dry gypsum ledges, matoral xerofilo; 1260–1350 m. Mexico. Specimen Examined. SLP (Heil 5347, BRY).

This species is unique in the genus in having lanose axes and abaxial pinna surfaces.

11. PELLAEA SAGITTATA Pellaea sagittata (Cav.) Link, Fil. Spec. 60. 1841. Fig. 216A. Pteris sagittata Cav., Descr. Pl. 267. 1801. Allosorus sagittatus (Cav.) C. Presl, Suppl. Tent. Pterid. 153. 1836. Platyloma sagittata (Cav.) J. Sm., J. Bot. (Hooker) 4: 160. 1841. Type. Mexico. Cerro de Guadaloupe, Ne´e s.n. (MA).

Rhizomes short, stout, compact; rhizome scales 3–4 ⫻ 0.5–0.8 mm, straight, light brown, concolorous, dull, linear-lanceolate to lanceolate, irregularly denticulate; fronds to 80 cm long, erect; stipes 1⁄4–1⁄2 the frond length, nearly terete to plane on adaxially, stramineous, with narrow orange scales toward the base, otherwise glabrous or rarely sparsely puberulous, rachises straight; blades narrowly oblong to deltate, 1–2-pinnate, light green, yellow-green to gray-green; pinnae widely spreading to slightly ascending, entire to pinnate, segments short-stalked, 20–28 ⫻ 8–14 mm at base, narrowly deltate to sagittate, entire, cordate at bases, apices acute, subcoriaceous, glabrous to sparsely pubescent at pinna bases with scattered hairs and scales, terminal pinnule of each pinnae with strongly inequilateral base (1–1.5 mm difference); margins recurved, glabrous, with whitened cells on recurved portion, indusia well differentiated, 0.3–0.5 mm wide, papillate; spores brown; n⫽2n⫽87, apogamous (Mich).

450

PELLAEA

Windham (1993) reported that subsp. arizonica occurs in Arizona, Texas, Chihuahua, and Sonora, without mentioning the status of P. ternifolia in the rest of Mexico. Specimens from Chihuahua and Sonora do indeed match the larger pinna size and ungrooved rachises of subsp. arizonica. In the rest of the country, pinna dimensions generally, but not always, agree with those of subsp. ternifolia, but there is every transition in the rachises from having distinct grooves to being flattened or terete. There is an especially large specimen from Guerrero, but it is not known whether it is abnormal subsp. ternifolia, disjunct subsp. arizonica, or perhaps an unnamed polyploid taxon. Key to the Mexican Subspecies of Pellaea ternifolia

Distribution. Rocky slopes in pine-oak woods; 1350–3050 m. USA (Tex); Mexico, Guat, Pan; Col, Ec, Peru, Bol. Selected Specimens Examined. Chih (Knobloch 5940, US). Chis (Ghiesbreght 303, GH). DF (Pringle 10152, NY). Dgo (Mendoza R. 156, UAMIZ). Gto (Dı´az B. & Zamudio 3482, IEB). Hgo (Mexia 2760, UC). Jal (McVaugh 12974, MICH). Me´x (Bourgeau 1163, GH). Mich (Feddema 215, MICH). Mor (Hitchcock & Stanford 7089, UC). Oax (Pringle 4887, NY). Pue (Purpus 2712a, NY). Qro (Basile 3, NY). SLP (Schaffner 959, GH). Tlax (Clausen s.n., 23 Aug 1955, MEXU). Zac (Correa 173, MICH). Unverified, Doubtful, or Mistaken Reports. Ags (Siqueiros 4148, HUAA, and eight other collections, cited by Siqueiros-Delgado & Gonza´lez-Adame, 2004, but not verified). Gro (Lorea 848, FCME, cited by Lorea-Herna´ndez & Vela´zquez M., 1998, but not verified).

Pellaea sagittata is distinguished from P. cordifolia by the ovate-deltate to sagittate (vs. nearly round) segments. The specimen cited from Durango has extraordinarily large segments 25– 35 ⫻ 12–23 mm.

1. Pinnae short-petiolate, 1–2 mm; BCS. .............. 12c. subsp. brandegeei. 1. Pinnae sessile; non-peninsular Mexico. 2. Largest segments less than 18 mm long; stipes and rachises grooved adaxially. ............................... 12a. subsp. ternifolia. 2. Largest segments more than 18 mm long; stipes distally and rachises rounded or slightly flattened adaxially. ............................................................. 12b. subsp. arizonica.

12a. Pellaea ternifolia subsp. ternifolia

Figs. 219D–F.

Stipes distally and rachises grooved or flattened adaxially; rachises glabrous or with a few widely scattered hairs; pinnae sessile, completely glabrous; largest ultimate segments (excluding terminal pinnae) usually less than 18 mm; spores usually 39–45 ␮m diam.; 2n⫽58 (DF, Mor, Pue, SLP, USA), 116 (Dgo, Me´x, Pue, SLP).

12. PELLAEA TERNIFOLIA Pellaea ternifolia (Cav.) Link, Fil. Spec. 59. 1841. Pteris ternifolia Cav., Descr. Pl. 266. 1801. Lectotype (chosen by C. Chr., Dansk Bot. Ark. 9(3): 22. 1937). Peru. Guarimaya valley near Guamantanga, Ne´e s.n. (MA; isotype F). For additional synonymy, see A. Tryon (1957).

Rhizomes short, stout, compact; rhizome scales 5–6 ⫻ 0.3–0.5 mm, bicolorous, linear, central portion dark reddish brown to black, margins thin, whitish tan, erose; fronds to 50 cm long, erect, monomorphic; stipes 1⁄4–1⁄3 the frond length, atropurpureous to black, lustrous, glabrous, terete to plane or grooved adaxially; blades linear, bipinnate almost to tips; pinnae ternately divided, gray-green, segments linear to lanceolate, 16 ⫻ 1–2 mm, entire, mucronate, each with 1 mm colorless tip, glabrous, coriaceous; margins reflexed, scarious, indusia 0.1 mm wide with papillate margins; spores tan. Distribution. Terrestrial and epipetric on rocky slopes and ledges in dry forests; 200–4000 m. Sw USA; Mexico; Central America; w South America; Hawaii.

Distribution. Rocky slopes and ledges on igneous substrates, dry, rocky slopes of mountains, oak or pine-oak forests, or Bursera-Acacia or cacti associations; (700–)1700–3000 m. USA (Ariz, Tex); Mexico; Guat, Hond, Nic, CR; Hisp; Col, Ven, Ec, Peru, Braz, Bol, Chile, Parag, Arg; Hawaii. Selected Specimens Examined. Ags (McVaugh 18425, NY). BCN (Montgomery 83-30, NY). Chih (Correll & Gentry 22920, NY). Chis (Breedlove 40471, DS). Coah (Johnston et al. 11897, NY). DF (Orcutt 4245, NY). Dgo (Breedlove 44111, MEXU). Gto (Galvan 3221, ENCB). Hgo (Medina C. 430, XAL). Jal (Breedlove 61658, NY). Me´x (Bourgeau 457, NY). Mich (Hinton 15497, NY). Mor (Orcutt 3827, MEXU, US). NL (Rodrı´guez L. 156, UNL). Oax (Mickel 3849, NY). Pue (Ventura A. 1555, NY). Qro (Weller 631, UC). SLP (Parry & Palmer 982, NY). Son

PELLAEA TRUNCATA

451

(Phillips 274, MICH). Tlax (Herna´ndez X. 334, MEXU). Ver (Scoville s.n., in 1891, NY). Zac (Feddema 2256, MEXU). Unverified, Doubtful, or Mistaken Reports. Gro (Lorea 4706, ENCB, IEB, cited by Lorea-Herna´ndez & Vela´zquez M., 1998, but not verified).

Pellaea ternifolia is widespread and quite variable in Mexico, especially in regard to pinna size and rachis grooving. See subsp. arizonica and brandegeei, and P. villosa for discussion. 12b. Pellaea ternifolia subsp. arizonica Windham, Contr. Univ. Michigan Herb. 19: 42. 1993. Type. U.S.A. Arizona: Cochise Co., SW slopes of Dragoon Mts, ca. 1.59 km NNE of Granite Spring, Windham 246 (UT!; isotypes ASC!, ASU!, MO!, UC!, US!). Figs. 219G, H. Stipes terete or slightly flattened adaxially in distal portions; rachises glabrous or with a few widely scattered hairs; pinnae sessile, glabrous; largest ultimate segments (excluding terminal pinnae) usually more than 18 mm; spores averaging 46–53 ␮m diam. 2n⫽116 (USA).

Distribution. 700–2000 m. Mexico; Hawaii. Selected Specimens Examined. BCS (Brandegee 662, GH, UC; Carter 2409, UC, US, 2685, US, 3360, MEXU; Chambers 921, UC; Jones 24556, 27013, NY, UC; Moran 18911, UC, US; Purpus 559, US; Tenorio L. 10601, GH, NY).

Material of Pellaea ternifolia from Baja California (subsp. brandegeei) has the pinna dimensions and rachis grooving of subsp. ternifolia. Some Hawaiian specimens of P. ternifolia have stalked pinnae, suggesting that part of the population arrived from Baja California.

13. PELLAEA TRUNCATA Pellaea truncata Goodd., Muhlenbergia 8: 94. 1912. Type. U.S.A. Arizona: Cochise Co., Mule mountains, in dry rocky places, Goodding 977 (US!). Figs. 218M, N.

Distribution. Cliffs and rocky slopes, on acidic substrates; 1720–2400 m. USA (Ariz, Tex); Mexico. Selected Specimens Examined. Chih (Pringle 920, NY). Gro (Hinton 9223, NY). Gto (Duges s.n., in 1880, NY). Jal (McVaugh 17096, NY). Me´x (Hinton 4234, NY). Mor (Fisher 35414, NY). Nay (Te´llez 12468, MEXU). Oax (Mickel 4941, NY). Pue (Arse`ne 1837, US). Sin (Breedlove 35708, MEXU). SLP (Parry & Palmer 982, NY). Son (Reichenbacher 1127, ARIZ). Ver (Ventura A. 178, NY).

12c. Pellaea ternifolia subsp. brandegeei (C. C. Hall) Mickel, comb. et stat. nov. Figs. 219L–N. Pellaea brandegeei C. C. Hall, Amer. Fern J. 37: 111, photo 112. 1947. Type. Mexico. Baja California Sur: Cape region, Sierra de la Laguna, 1893, Brandegee s.n. (UC!; isotype US!).

Pellaea ternifolia subsp. brandegeei differs from subsp. ternifolia in the pinnae being distinctly short-petiolulate 1–2 mm. All specimens of subsp. brandegeei have grooved distal stipes and rachises. The distal 1–3 pinna pairs are undivided.

Rhizomes compact, ascending, 5–10 mm diam.; rhizome scales linear-subulate, bicolorous, black with brown, erose-dentate margins; fronds somewhat dimorphic, sterile shorter and less divided than the fertile, 10–40 cm long, clumped; stipes ca. 1⁄3 the frond length, castaneous to dark brown, lustrous, adaxially flattened or shallowly grooved, glabrous or nearly so; blades ovatedeltate, bipinnate, 5–18 cm wide; rachises straight, glabrous; pinnae 5–8 pairs of divided pinnae; ultimate segments narrowly oblong, 4–10 mm long, coriaceous, glabrous, apices mucronate; adaxial and abaxial surfaces glabrous; sporangia 64-spored, mixed with farina; spores brown; 2n⫽58 (USA).

452

PELLAEA

Distribution. Cliffs and rocky slopes, on various substrates but rarely observed on limestone; 500–1650 m. W USA; Mexico.

Distribution. Rocky slopes and ledges on igneous substrates; 1900–2250 m. USA (Tex); Mexico.

Selected Specimens Examined. BCN (Brandegee s.n., 20 Apr 1889, GH, NY; Montufar L. 74, ENCB; Moran 7996, UC; Wiggins 9826, 16846, DS). BCS (Moran 11671, DS, UC, 18727, UC, US). Son (McManus 737, MEXU; Phillips 338, 419, MICH).

Selected Specimens Examined. Chih (Pringle 446, NY). Coah (Wynd & Mueller 574, NY). Dgo (Maysilles 8288, NY). Gto (Galvan 2332D, ENCB). Hgo (Copeland herb. 109, NY). Me´x (Rzedowski 22894, NY). Mich (Rzedowski 49942, IEB). NL (Rodrı´guez L. 174, UNL). Pue (Purpus 4032, UC). SLP (Limon 51, MEXU). Son (White 4221, US).

Unverified, Doubtful, or Mistaken Reports. Chih (Correll 23306, cited by Knobloch & Correll, 1962, without herbarium, not verified).

Pellaea truncata crosses with P. wrightiana in areas of overlap, and the hybrids can be distinguished by their malformed spores.

The generally less divided blades and the laxly hairy rachises suggest that Pellaea villosa may have originated as a cross between P. ternifolia and P. ribae.

14. PELLAEA VILLOSA

15. PELLAEA WRIGHTIANA

Pellaea villosa (Windham) Windham & Yatsk., Novon 13: 361. 2003. Figs. 219J, K. Pellaea ternifolia (Cav.) Link subsp. villosa Windham, Contr. Univ. Michigan Herb. 19: 43. 1993. Type. Mexico. Hidalgo: Lena Station, rocky hills, Pringle 10025 (LL; isotypes ARIZ, BRY, CAS, CHAPA!, COLO, DAO, ENCB, F, GH, IND, KANU, LL, MEXU!, MSC, NY!, OKLA, SMU, TEX, UC, US, VT, WIS).

Blades 1-pinnate distally, pinnae ternate in proximal 1⁄4–3⁄4; stipes atropurpureous to black, distally terete or slightly grooved adaxially, rarely grooved in distal portion; rachises villous, especially in axils of pinnae; pinnae with hairs scattered along main veins abaxially; largest ultimate segments (excluding terminal pinnae) usually more than 18 mm; spores tan, usually 46–53 ␮m diam.; 2n⫽116 (Dgo, Pue, SLP).

Pellaea wrightiana Hook., Sp. Fil. 2: 142. 1858. Figs. 219O–Q. Pellaea ternifolia (Cav.) Link var. wrightiana (Hook.) A. F. Tryon, Ann. Missouri Bot. Gard. 44: 153. 1957. Type. U.S.A. New Mexico: Wright 2130 (K; isotypes GH!, NY!, US!, YU!).

Rhizomes compact, ascending, 5–10 mm diam.; rhizome scales linear-subulate, bicolorous, erose-denticulate, 0.1–0.3 mm wide at bases; fronds monomorphic, to 40 cm long, clumped; stipes 1 ⁄3–2⁄5 the frond length, dark brown, lustrous, adaxially flattened or shallowly grooved, glabrous; blades linear-oblong, 2-pinnate, 1.5–5 cm wide; rachises straight, glabrous; pinnae perpendicular to rachises or slightly ascending, with 3–9 ultimate segments; ultimate segments narrowly oblong, 5–20 mm long, each with mucronate apices, coriaceous, glabrous; adaxial and abaxial surfaces glabrous; sporangia 64-spored, mixed with sparse farina; spores brown; n⫽2n⫽87, 116 (USA).

PENTAGRAMMA

453 References

Alt, K. S. & V. Grant. 1960. Cytotaxonomic observations on the goldback fern. Brittonia 12: 153–170; Tryon, R. M. 1962. Taxonomic fern notes. II. Pityrogramma (including Trismeria) and Anogramma. Contr. Gray Herb. 189: 52–76; Yatskievych, G., M. D. Windham & E. Wollenweber. 1990. A reconsideration of the genus Pityrogramma (Adiantaceae) in western North America. Amer. Fern J. 80: 9–17.

1. PENTAGRAMMA TRIANGULARIS Pentagramma triangularis (Kaulf.) Yatsk., Windham & E. Wollenw., Amer. Fern J. 80: 15. 1990. Distribution. Cliffs and rocky slopes, in pine and oak woods; 1450–2200 m. Sw USA; Mexico. Selected Specimens Examined. BCN (Brandegee s.n., 14 May 1893, UC; Wiggins & Demaree 5024, DS, US). BCS (Gentry 4283, GH, MO; Gentry & Fox 11775, MEXU, US). Chih (Johnston 8090, GH; Pennell 19209, US; Wendt et al. 9971, MEXU, NY). Coah (Johnston & Muller 1309, GH, US). Son (Druery s.n., ca. 1895, US; White 4222, MICH).

According to Windham (in FNA Ed. Comm., 1993: 182), P. wrightiana, a tetraploid, hybridizes with Pellaea truncata and P. ternifolia subsp. arizonica to produce sterile triploids and tetraploids with intermediate morphology and malformed spores.

93 . PE N T A G R A M M A Pentagramma Yatsk., Windham & E. Wollenw., Amer. Fern J. 80: 13. 1990. Type: Pentagramma triangularis (Kaulf.) Yatsk., Windham & E. Wollenw. [⬅ Gymnogramma triangularis Kaulf.] Terrestrial or epipetric; rhizomes short-creeping, erect or ascending; rhizome scales bicolorous, each with black, sclerotic central stripe and tan margins, linear-lanceolate, entire; fronds monomorphic, clumped, 5–40 cm long; stipes castaneous to black, terete, glabrous, farinose or viscid-glandular; blades deltate-pentagonal, 1–2-pinnate-pinnatifid proximally, pinnatifid to pinnate-pinnatifid distally, herbaceous to coriaceous, adaxially glabrous or glandular, dull, abaxially farinose, farina white or yellow; ultimate segments sessile, adnate to rachises and costae their full width, deltate to oblong, margins not recurved, not differentiated; veins free, obscure; sporangia borne along veins, containing 64 spores, without indusia; spores tan to brown, tetrahedral; x⫽30. Pentagramma is a genus of only two species of western North America and Mexico (one species in northwestern Mexico with three subspecies). Traditionally, Pentagramma was included in Pityrogramma, based on the farinose fronds and gymnogrammoid sori, but evidence from morphology, cytology, and molecular studies have shown that the two genera are not closely related. Taxa within the genus have been treated variously, and further studies are needed.

Gymnogramma triangularis Kaulf., Enum. Filic. 73. 1824. Pityrogramma triangularis (Kaulf.) Maxon, Contr. U.S. Natl. Herb. 17: 173. 1913. Type. U.S.A. California: [near San Francisco fide Alt & Grant, 1960] 1816, Chamisso s.n. (B!).

Rhizomes compact, horizontal; rhizome scales castaneous to dark brown, lustrous, glabrous or sometimes viscid-glandular or rarely somewhat white-farinose proximally; blades broadly deltate-pentagonal, herbaceous to subcoriaceous; adaxial surfaces glabrous to glandular or viscid; abaxial surfaces farinose, farina white or yellow. Distribution. Terrestrial or epipetric in dry pine and oak woodlands; 30–1520 m. W Can and USA; Mexico. Key to the Mexican Subspecies of Pentagramma triangularis 1. Fronds viscid-glandular adaxially; the two pinna pairs above the basal pair irregularly pinnatifid to often entire; proximal basiscopic lobes of basal pinnae entire to undulate or crenate. ...................................................... 1c. subsp. viscosa. 1. Fronds glabrous or with scattered yellowish capitate glands adaxially, not viscid-glandular; the two pinna pairs above the basal pair regularly pinnatifid, proximal basiscopic lobes of basal pinnae pinnatifid, often deeply so. 2. Fronds with scattered yellowish, capitate, nonfarinose glands adaxially, white-farinose abaxially. ................................................................................ 1b. subsp. maxonii.

2. Fronds glabrous adaxially, yellow- or white-farinose abaxially. ............................................................ 1a. subsp. triangularis.

1a. Pentagramma triangularis subsp. triangularis Figs. 220A–C. Pentagramma triangularis (Kaulf.) Yatsk., Windham & E. Wollenw. subsp. semipallida (J. T. Howell) Yatsk., Windham & E. Wollenw., Amer. Fern J. 80: 16. 1990. Pityrogramma triangularis (Kaulf.) Maxon var. semipallida J. T. Howell, Leafl. W. Bot. 9: 223. 1962. Type. U.S.A. California: Butte Co., on metamorphic rocks in canyon of North Fork Feather River, 9 mi NE of Oroville, Howell 34696 (CAS!; isotypes G, US!).

Fronds 7–23 cm, clumped; stipes 3/5–4/5 the frond length, glabrous, not viscid-glandular; blades broadly deltatepentagonal, herbaceous to subcoriaceous, not viscid-glandular, proximal basiscopic lobes of basal pinnae pinnatifid, the two pinna pairs above the basal pair regularly pinnatifid; adaxial sur-

454

PENTAGRAMMA

faces glabrous; abaxial surfaces white- to bright yellow-farinose; 2n⫽60, ca. 90, 120 (USA).

Distribution. Pine and oak woodlands; 50–1520 m. USA (Ariz, Calif); Mexico. Distribution. Dry pine and oak woodlands; 30–1200 m. Can (BC), w USA (Calif, Idaho, Ore, Wash); Mexico. Selected Specimens Examined. BCN (Cleveland s.n., 23 Feb 1883, UC; Gallegos s.n., in 1923, MEXU; Orcutt s.n., 17 May 1886, US; Palmer 633, US; Raven et al. 12494, 12674, UC; Rovirosa s.n., 20 Feb 1923, MEXU). Guad (Franceschi 38, UC; Palmer 856, MEXU, UC; Wiggins & Ernst 82, UC).

Subspecies triangularis varies extensively in morphology, cytology, and chemistry. White-farinose specimens have sometimes been segregated as subsp. semipallida, but that seems to be just part of broader variation in farina color. Much work is needed to determine the significance of these variations. 1b. Pentagramma triangularis subsp. maxonii (Weath.) Yatsk., Windham & E. Wollenw., Amer. Fern J. 80: 16. 1990. Figs. 220D, E. Pityrogramma triangularis (Kaulf.) Maxon var. maxonii Weath., Rhodora 22: 119. 1920. Type. U.S.A. Arizona: Rincon Mtns, head of Rincon Valley, Blumer 3271 (US!; isotypes ARIZ!, DS!, GH!, MO!).

Fronds 6–22(–34) cm, clumped; stipes 1⁄2–3⁄4 the frond length, glabrous, not viscid-glandular, occasionally white-farinose proximally; blades broadly deltate-pentagonal, herbaceous (but not leathery), not viscid-glandular, proximal basiscopic lobes of basal pinnae pinnatifid, the two pinna pairs above the basal pair regularly pinnatifid; adaxial surfaces with scattered yellow capitate glands; abaxial surfaces white-farinose; 2n⫽120 (USA).

Selected Specimens Examined. BCN (Brandegee s.n., 20 Apr 1889, UC; Harvey 541, US; Moran 28107, SD). BCS (Brandegee s.n., Sep 1893, UC; Carter 2411, MEXU, UC; Gentry & Fox 11774, MEXU, UC, US; Jones s.n., 22 Sep 1930, UC). Guad (Drent s.n., in 1898, UC; Kuijt & Miller 1011, UC; Newcomb 179, UC, US)

Specimens intermediate between subsp. maxonii and both subsp. triangularis and subsp. viscosa occur. 1c. Pentagramma triangularis subsp. viscosa (Nutt. ex D. C. Eaton) Yatsk., Windham & E. Wollenw., Amer. Fern J. 80: 15. 1990. Figs. 220F, G. Gymnogramma viscosa Nutt. ex D. C. Eaton, Ferns N. Amer. 2: 16. 1879. Pityrogramma triangularis (Kaulf.) Maxon var. viscosa (Nutt. ex D. C. Eaton) Weath., Rhodora 22: 117. 1920. Pityrogramma viscosa (Nutt. ex D. C. Eaton) Maxon, Contr. U.S. Natl. Herb. 17: 173. 1913. Type. U.S.A. California: San Diego, Nuttall s.n. (PH!).

Fronds 9–37 cm, clumped; stipes 1⁄2–3⁄4 the frond length, sometimes viscid-glandular; blades broadly deltate-pentagonal, subcoriaceous, proximal basiscopic lobes of basal pinnae entire to undulate or crenate, the two pinna pairs above the basal pair irregularly pinnatifid to often entire; adaxial surfaces viscidglandular; abaxial surfaces white-farinose and viscid-glandular; 2n⫽60 (USA).

Distribution. Pine and oak woodlands; 50–1475 m. USA (Calif); Mexico.

PHANEROPHLEBIA Selected Specimens Examined. BCN (Brandegee s.n., 22 Feb 1883, UC; Gallegos s.n., 15 Feb 1923, MEXU; Johnston 3023, US; McKeever 5, 7, US; Raven et al. 12260, UC; Shreve 6415, MEXU; Thorne 61888, UC; Wiggins 4237, UC, 9743, UC, US; Wiggins & Gillespie 4045, MEXU).

Alt and Grant (1960) noted both diploid and tetraploid plants intermediate between subsp. viscosa and triangularis at some localities where these two co-occur. Intermediates also occur between subsp. viscosa and maxonii.

94 . P H A N E R O PH L E B I A Phanerophlebia C. Presl, Suppl. Tent. Pterid. 84. 1836. Type: Phanerophlebia nobilis (Schltdl. & Cham.) C. Presl [⬅ Aspidium nobile Schltdl. & Cham.]. Terrestrial (or among rocks); rhizomes compact, shortcreeping or ascending to erect, scaly; fronds small to mediumsized, less than 100(–135) cm, clumped; stipes scaly, especially at the bases, filiform-scaly to glabrescent distally; blades 1-pinnate, each with a conform terminal pinna; rachises usually with at least a few persistent filiform or lanceolate scales; pinnae coriaceous or chartaceous, spinulose-serrate or less commonly entire at margins, with tan appressed hairs 0.1–0.5 mm and hair-like scales to ca. 2 mm on the veins and tissue between the veins; veins pinnately branched, free to casually or sometimes regularly anastomosing, branches strongly upcurved and running toward the pinna margins; sori round, in mostly 2–4 series between costae and margins, indusiate or exindusiate in a few species, indusia firm, peltate, often caducous; spores bilateral, with cristate perispore; x⫽41. Phanerophlebia is a neotropical genus of 9 species, from southwestern United States (Texas to Arizona) to northern South America and Hispaniola; all but one species (P. haitiensis C. Chr., from Hispaniola and Honduras) occur in Mexico. Most

455

species grow in mesic forests at middle to higher elevations, often in calcareous rock crevices or calcareous soil. Phanerophlebia is clearly a member of the large dryopteroid clade, also including Polystichum, Dryopteris, Arachniodes, and probably Stigmatopteris and Ctenitis in Mexico; Adenoderris may also be a part of this clade. Phanerophlebia has traditionally been distinguished from Cyrtomium on the basis of free vs. netted veins, but some phanerophlebias have netted veins and some cyrtomiums have free veins. The distinction of Phanerophlebia (1-pinnate fronds with multiseriate sori) from Polystichum (many narrow pinnae) is tenuous, and the entire group (polystichoid ferns) needs more work to see if there are valid generic distinctions. Yatskievych et al. (1988) and Stein et al. (1989) provided evidence from chloroplast DNA restriction site differences that the morphological similarities between Phanerophlebia and Cyrtomium are the result of convergent evolution. Therefore, if lumping is to be done, Phanerophlebia should merged with Polystichum rather than with just Cyrtomium. Little and Barrington (2003) have evidence that Phanerophlebia is monophyletic and the sister of Polystichopsis. This clade forms the sister group of Polystichum, including Cyrtomium and Cyrtomiopsis, which nest within Polystichum. References Little, D. P. & D. S. Barrington. 2003. Major evolutionary events in the origin and diversification of the fern genus Polystichum (Dryopteridaceae). Amer. J. Bot. 90: 508–514; Maxon, W. R. 1912. Notes on the North American species of Phanerophlebia. Bull. Torrey Bot. Club 39: 23–28; Morton, C. V. 1957. Observations on cultivated ferns. II. The proper generic name of the holly fern. Amer. Fern J. 47: 52–55; Stein, D. B., G. Yatskievych & G. J. Gastony. 1989. Chloroplast DNA evolution and phylogeny of some polystichoid ferns. Biochem. Syst. Ecol. 17: 93–101; Underwood, L. M. 1899. American ferns–II. The genus Phanerophlebia. Bull. Torrey Bot. Club 26: 205–216; Yatskievych, G. 1992. Innovations in the fern genus Phanerophlebia. Novon 2: 445–446; Yatskievych, G. 1996. A revision of the fern genus Phanerophlebia (Dryopteridaceae). Ann. Missouri Bot. Gard. 83: 168–199; Yatskievych G. & R. Riba. 1999. Pteridofitas: Familia Dryopteridaceae. Phanerophlebia. Flora de Mexico 6(4): 26–38; Yatskievych, G., D. B. Stein & G. J. Gastony. 1988. Chloroplast DNA evolution and systematics of Phanerophlebia (Dryopteridaceae) and related fern genera. Proc. Natl. Acad. Sci. USA 85: 2589–2593.

Key to the Mexican Species of Phanerophlebia 1. Veins regularly or irregularly but copiously anastomosing. 2. Lateral pinna pairs 8–18, 1.5–3.5 cm wide, spinulose along entire length; indusia light brown, 0.8 mm diam. ........................ 7. P. remotispora. 2. Lateral pinna pairs 0–4 (rarely to 6), 2–5 cm wide, spinulose only in distal half; indusia absent. 3. Stipes scales not overlapping, 4–6 mm long; buds absent in axils of distal pinnae; spores 30–42 ␮m long. ............................ 2. P. gastonyi. 3. Stipe scales usually overlapping, 6–11 mm long; buds often present in axils of distal pinnae; spores 41–60 ␮m long. ... 3. P. juglandifolia. 1. Veins free or exceptionally and casually anastomosing at rare intervals. 4. Pinnae 0–4 pairs, to ca. 8(–12) cm long, rounded to cordate at bases. .................................................................................................. 6. P. pumila. 4. Pinnae (2–)6–17 pairs, (2–)10–25 cm long, broadly cuneate, truncate, or rounded at bases. 5. Stipes densely and persistently scaly at bases and sometime throughout with tan, ovate scales; rhizome scales tan, 10–15 ⫻ 5–7 mm; plants emitting a skunk-like odor when fresh. ................................................................................................. 4. P. macrosora. 5. Stipes sparsely scaly with dark brown scales; rhizome scales dark brown, 3–7.5 ⫻ 1–3 mm; plants without a skunk-like odor when fresh. 6. Pinnae mostly auriculate at acroscopic bases; northwest and north-central Mexico. .......................................................... 1. P. auriculata. 6. Pinnae lacking an auricle at acroscopic bases. 7. Indusia shriveled at maturity, convex, lacking a distinct central umbo; nearly throughout Mexico. ............................... 5. P. nobilis. 7. Indusia persistent at maturity, concave, with a distinct (in color and relief) raised central projection (umbo); northeast and north-central Mexico. ..................................................................................................................................... 8. P. umbonata.

456

PHANEROPHLEBIA

1. PHANEROPHLEBIA AURICULATA Phanerophlebia auriculata Underw., Bull. Torrey Bot. Club 26: 212. 1899. Figs. 221D–F. Cyrtomium auriculatum (Underw.) C. V. Morton, Amer. Fern J. 47: 54. 1957. Type. Mexico. Chihuahua: Mapula Mts., Pringle 831 (NY!; isotypes CAS!, GH!, K, NY!, PH, UC!, US!).

Rhizomes short-creeping to ascending, sometimes branched; rhizome scales concolorous, reddish brown to orangish, ovate, 3.5–7.5 ⫻ 1–2(–3) mm, margins ciliate; fronds 10–60(–75) cm long, very short fronds sometimes fertile; stipes 1⁄3–1⁄2 the frond length, stramineous, moderately scaly with tan, appressed often caducous scales similar to those of rhizomes; blades ovate to lanceolate, 13–50 cm long; rachises with sparse, tan, hair-like, twisted scales; buds absent from axils of distal pinnae; pinnae 2–8(–12) lateral pairs, ovate to lanceolate, usually falcate, 2–12 ⫻ (1–)2– 3(–3.5) cm, bases truncate to obliquely and broadly cuneate to nearly cordate, apices usually attenuate, lowermost pinnae inequilateral, excavate basiscopically, prominently auriculate acroscopically, margins spinulose-serrulate throughout; veins free or very casually anastomosing, 2–4-branched; sori in 2–4 series between costae and pinna margins, inner row 2–3 mm from costa, outer row often 1–3 mm from margins; indusia light brown, 0.6–0.9 mm diam., flat or concave, with a central umbo, shriveling at maturity; 2n⫽164 (Ariz, N Mex).

lis and P. umbonata, followed by chromosome doubling (allopolyploidy); this seems like a plausible scenario.

2. PHANEROPHLEBIA GASTONYI Phanerophlebia gastonyi Yatsk., Novon 2: 445. 1992. Type. Mexico. Chiapas: ca. 13 km NW of Berriozabal, 1000 m, Yatskievych et al. 85-182 (MO!; isotypes ARIZ, CHAPA, MEXU, IND, NY!, UAMIZ!, UC!). Figs. 222A–E, Q, R. Rhizomes short-creeping to ascending, not branched; rhizome scales bicolorous with broad, dark brown or sclerotic centers and narrow, lighter brown ciliate margins, ovate to ellipticlanceolate, 2.5–6 ⫻ 1.5–2 mm; fronds to 25–45(–55) cm long; stipes usually longer than blades, stramineous, with non-overlapping scales 4–6 mm long primarily at the base, caducous distally, reduced hair-like scales uncommon; blades ovate, 14–25 cm long; rachises glabrescent or sparingly scaly; buds absent from axils of distal pinnae; pinnae usually (0–)1–4 lateral pairs, ovate to lanceovate, sometimes somewhat falcate, 7–16 ⫻ 2–4 cm, bases obliquely cuneate to rounded, sometimes nearly equilateral, apices acute to attenuate, margins entire to slightly undulate proximally, spinulose-serrulate only in distal half, costae and veins abaxially with rare to sparse minute, hair-like scales 0.5–0.8 mm long on costae, 0.1–0.3 mm long on veins; veins 2–4-branched, with 1–3 series of Ⳳ regular to somewhat irregular marginal anastomoses; sori in (1–)2–4 series between costae and pinna margins, inner row 1–4 mm from costa, outer row 2–5 mm from margin; indusia absent; spores 30–42 mm long; 2n⫽82 (Chis, Oax, as P. juglandifolia, Ver).

Distribution. Terrestrial or in rock crevices, often granitic or quartzite, of canyons and ravines; 600–2150 m. USA (Ariz, N Mex, Tex); Mexico. Selected Specimens Examined. Chih (Correll & Gentry 23043, LL, UC, US, 23241, LL, NY; Palmer 450, NY, US; Reeves 4880, ASU; Spellenberg 6814, ASU, NY). Coah (Stewart 2198, LL). Son (Phillips 546, ARIZ, NY, US; Reeves 6374, ASU); Van Devender s.n., ARIZ, UC; Wiggins 7139, ARIZ, UC, US). Unverified, Doubtful, or Mistaken Reports. NL (reported by Aguirre-Clavera´n & Arreguı´n-Sa´nchez, 1988, but the specimen on which this was based is more likely P. umbonata. One collection from Sonora (Reeves 6374) is more like P. nobilis in the pinna length (to 12 cm) and large number of lateral pinna pairs (ca. 12), but the proximal pair of pinnae are auriculate, like P. auriculata, on one frond (but not four others). Yatskievych (1996) postulated that P. auriculata, a tetraploid species, may have arisen through hybridization between P. nobi-

Distribution. On limestone outcrops in wet montane and pineoak-Liquidambar forests; 900–2200 m. Mexico; Hond? (see comments under P. juglandifolia). Selected Specimens Examined. Chis (Breedlove 21652, DS, MEXU, NY, 30867, DS, MEXU, 31495, DS; Palacios-Rios 2901, UC). Oax (Mickel 3786B, NY, 6044, MO, NY, UC, 6182, MO, NY). Ver (Conant 726, GH, MEXU; Ventura A. 9265, MEXU, NY, 12342, IEB).

This species is exceedingly closely related to P. juglandifolia, differing primarily by the fewer, smaller stipe base scales with a blackened central area. In addition, P. gastonyi always lacks rachis

PHANEROPHLEBIA MACROSORA

buds (sometimes present in P. juglandifolia) and has veins that are not so highly anastomosing. The morphological differences between these two species seem of less significance to us than the differences between P. nobilis and P. remotispora, which Yatskievych treated as conspecific. Chromosome counts indicate that P. gastonyi is diploid, and one of the probably parents of the tetraploid species, P. juglandifolia (Yatskievych, 1996). A collection from Oaxaca (Geith 95-M007, NY) has simple blades ca. 10–12 ⫻ 2.5–3 cm, and is fertile. The veins are anastomosing, but the slender stipes, ca. 1 mm diam., are nearly devoid of scales and so the specimen is difficult to name. It appears to be a depauperate specimen of P. gastonyi. Judging by the rhizome scales and absence of rachis buds, a collection (Palma G. 67, XAL) determined by Yatskievych as P. juglandifolia from Veracruz is P. gastonyi. This same collection and Ramı´rez 527 (XAL), also P. gastonyi, were reported by Palacios-Rios (1992) as the only collections of P. juglandifolia from Veracruz. Herna´ndez M. 5916 (ENCB), from Hidalgo, cited as P. juglandifolia by Yatskievych (1996: 189), also seems to us to be P. gastonyi.

3. PHANEROPHLEBIA JUGLANDIFOLIA Phanerophlebia juglandifolia (Humb. & Bonpl. ex Willd.) J. Sm., J. Bot. (Hooker) 4: 187. 1841. Polypodium juglandifolium Humb. & Bonpl. ex Willd., Sp. Pl., ed. 4, 5(1): 195. 1810. Cyrtomium juglandifolium (Humb. & Bonpl. ex Willd.) T. Moore, Index Fil. lxxxiii. 1857. Type. Venezuela. Caripe, Humboldt & Bonpland s.n. (BWilld. 19688!, sheets 1a, 1b, frag. & drawings NY!; isotype P-Humb.!–2 sheets).

Rhizomes short, suberect, not branched; rhizome scales light brown, bicolorous with broad to narrow dark brown or sclerotic centers and tan ciliate margins, ovate-lanceolate, 3–10 ⫻ 1.5–5 mm; fronds mostly 60–90 cm long; stipes stramineous, equaling or longer than blades, with a few, brownish, ovate, usually overlapping scales 6–11 mm long similar to those of the rhizomes and scattered, twisted, hair-like scales 1–2 mm long; blades ovate, 20–45 cm long; rachises sparingly scaly to glabrescent; pinnae 2– 4(–6) lateral pairs, elliptic to ovate, subfalcate, 9–10(–20) ⫻ 2– 3(–5) cm, bases rounded to broadly cuneate, often inequilateral, apices acute to acuminate, margins entire to spinulose only in distal 1⁄3, costae and veins abaxially with rare to sparse minute, hair-like scales 0.5–0.8 mm long on costae, 0.1–0.3 mm long on veins; buds present or absent in axils of distal pinna, 0–1(–6) per blade; veins 3–5-branched, Ⳳ regularly anastomosing about midway between costae and margins; sori in 3–4(–6) series between costae and pinna margins, inner row 2–3 mm from costae, outer row 2–3 mm from margins; indusia absent; spores 41–60 ␮m long; 2n⫽164 (Chis, as P. cf. remotispora, CR).

457

Distribution. Terrestrial in wet forests; (700–)1350–2700 m. Mexico; Guat, Hond, Salv, Nic, CR, Pan; Col, Ven. The specimen cited by Yatskievych (in Davidse et al., 1995: 216) from Honduras lacks a bud and has a single pair of lateral pinnae; it appears more like P. gastonyi to us. Selected Specimens Examined. Chis (Breedlove 26840, DS, MEXU, NY, 34692, CHAPA, DS, NY, 41708, 42564, DS, MEXU, 46731, 48670, CAS; Dressler 1613, NY, US). Hgo (Herna´ndez M. 5916, ENCB, MO). Unverified, Doubtful, or Mistaken Reports. Gro (Lozano V. 215, FCME, cited by Lorea-Herna´ndez & Vela´zquez M., 1998, but not verified; this could also be P. gastonyi).

Phanerophlebia juglandifolia is distinct in its few pinna pairs, regularly anastomosing veins, and in the usual presence of a bud in the axil of a distal pinna. However, this last character is not always reliable in Mexico, as several specimens lack buds. Nor is it very reliable in Central America, where all 12 specimens (in UC) from Costa Rica lack buds, as do 4 of 6 Panamanian specimens seen. The species is very closely related to P. gastonyi (which see) and thought to be an allopolyploid derivative of that species (Yatskievych, 1992, 1996). Specimens cited as this species by Mickel and Beitel (1988) are P. gastonyi (Mickel 6044, 6182), as determined by Yatskievych.

4. PHANEROPHLEBIA MACROSORA Phanerophlebia macrosora (Baker) Underw., Bull. Torrey Bot. Club 26: 213. 1899. Figs. 222M–P. Aspidium juglandifolium (Humb. & Bonpl. ex Willd.) Kunze ex Klotzsch var. macrosorum Baker, J. Bot. 25: 25. 1887. Cyrtomium macrosorum (Baker) C. V. Morton, Amer. Fern J. 47: 55. 1957. Type. Costa Rica. Cooper s.n. (K!– frag. NY!; isotype US!). Phanerophlebia guatemalensis Underw., Bull. Torrey Bot. Club 26: 214. 1899. Cyrtomium guatemalense (Underw.) C. V. Morton, Amer. Fern J. 47: 55. 1957. Type. Guatemala. Quiche´: San Miguel Uspanta´n, Heyde & Lux s.n. [Donn. Sm. 3241] (NY!; isotypes GH!, P!, US!).

Rhizomes suberect to erect, unbranched; rhizome scales concolorous, tan, ovate-lanceolate, 10–12(–15) ⫻ 5–7 mm, margins ciliate-denticulate; fronds 70–135(–200⫹) cm long; stipes 1⁄3–1⁄2 the frond length, stramineous, densely scaly with mostly spread-

458

PHANEROPHLEBIA

ing tan, persistent scales similar to those of rhizomes; blades ovate to deltate, 40–90 cm long, smelling like a skunk when fresh; rachises with scattered to dense, pale, ovate, denticulate, caducous scales mostly 1–3 mm long, the narrowest ones twisted, hairlike; buds absent from axils of distal pinnae; pinnae 6–17 lateral pairs, linear-lanceolate, straight to slightly falcate, (8–)15–25 ⫻ 2–4.5 cm, bases inequilateral, broadly cuneate to rounded, the proximal ones basiscopically excavate, with auricles acroscopically, margins faintly crenulate and minutely spinulose; veins free, 3–4-branched; sori round, in 3–5 series between costae and pinna margins, inner row 2–3 mm from costa, outer row 1–10 mm from costa; indusia tan, caducous, 0.8–1 mm diam., flat or concave centrally, erose, not umbonate; 2n⫽82 (Oax, CR).

Rhizomes suberect to erect, usually unbranched; rhizome scales concolorous, brown, ovate to ovate-lanceolate, 3.5–10 ⫻ 1–3 (–4) mm, margins denticulate to ciliate; fronds (15–)40–100 cm long; stipes 1⁄3–1⁄2 the frond length, stramineous, densely scaly with appressed to spreading scales similar to those of rhizomes, scales brown to tan, often caducous or abraded; blades ovate to ovate-lanceolate, (10–)20–55 cm long; rachises with sparse, hairlike, twisted, caducous scales; buds absent from axils of distal pinnae; pinnae mostly 6–15 lateral pairs, linear-lanceolate, somewhat falcate, (3–)6–13(–20) ⫻ (1–)1.5–3 cm, bases rounded to broadly cuneate, apices attenuate, lowermost pinnae often inequilateral excavate basiscopically, usually without auricles acroscopically, margins spinulose-serrate throughout; veins free or only casually anastomosing near pinna margins, 1–3-branched; sori in 2–3(–4) series between costae and pinna margins, inner row 2–3 mm from costa, outer row 1–3 mm from margin; indusia tan, caducous, 0.6–1.3 mm diam., flat or concave, not umbonate; 2n⫽82 (Me´x, Oax).

Distribution. Terrestrial in cloud forests, pine forests, at middle to higher elevations; 1550–2750 m. Mexico; Guat, Salv, CR, Pan. Selected Specimens Examined. Chis (Breedlove 41713, DS; Matuda 5230, LL, MEXU, TEX). Gro (Lorea 4090, FCME; Martı´nez S. 5067, ARIZ, ENCB, MEXU, UAMIZ). Hgo (Croat & Hannon 65955, UC; Rebolledo V. 291, CHAPA, ENCB, NY). Me´x (Tejero-Die´z 2255, IZTA). Mich (Rzedowski 46201, UAMIZ). Oax (Yatskievych 83-467, UC). Ver (Narave F. 319, IEB, XAL, 370, XAL).

Phanerophlebia macrosora is distinct by its many pinna pairs, free veins, and rather densely scaly stipes. When fresh, plants produce a strong, unpleasant, skunk-like odor (Mickel & Beitel, 1988; Yatskievych, 1996), apparently the only species in the genus that has this characteristic. Putative hybrids with P. juglandifolia are known from Costa Rica (e.g., Yatskievych 86-31a, NY, UC). Fuentes 760 (CHAPA, NY), from Jalisco, determined as this species by Mickel (1992), is P. nobilis. Croat & Hannon 65955, from Hidalgo, was cited by Yatskievych (1996) as P. nobilis var. remotispora.

5. PHANEROPHLEBIA NOBILIS Phanerophlebia nobilis (Schltdl. & Cham.) C. Presl, Suppl. Tent. Pterid. 85, t. 2, f. 19. 1836. Figs. 221A–C. Aspidium nobile Schltdl. & Cham., Linnaea 5: 610. 1830. Cyrtomium nobilis (Schltdl. & Cham.) T. Moore, Index Fil. lxxxiii. 1857. Type. Mexico. Veracruz: “Hacienda de la Laguna,” Schiede [760] (B!).

Distribution. Terrestrial in montane forests; (1050–)1500– 2800(–3200) m. Mexico. Selected Specimens Examined. Chih (Gentry 7988, ARIZ, MEXU, UC, US). Chis (Matuda 4688, LL, MO, NY). DF (Rzedowski 27229, ENCB, LL, NY, TEX). Gro (Lorea 643, ENCB). Hgo (Gonza´lez Q. 2419, LL, NY). Jal (Mexia 1558, UC, US). Me´x (Hinton 403, NY). Mich (Rzedowski 46201, IEB, UAMIZ). Oax (Camp 2744, UC). Pue (Purpus 2454, UC). Qro (Ferna´ndez N. 3373, ENCB, IEB, NY; Ferna´ndez & Zamudio 992, NY). SLP (Schaffner s.n., in 1876, NY). Sin (Ownbey & Ownbey 1936, US). Son (Gentry 3660, ARIZ, UC). Tam (Johnston 12777, TEX). Tlax (Nicolas s.n., MEXU, UC, US). Ver (Copeland herb. 39, GH, MEXU, MICH, MO, UC, US). Unverified, Doubtful, or Mistaken Reports. Mor (Camacho G. 1058, ENCB, cited by Yatskievych, 1996, but not verified).

Yatskievych et al. (1988) treated P. remotispora as a variety of P. nobilis. He later (1992, 1996) cited apparently fertile hybrids between the two varieties, Arse`ne 5961 (GH!, MO, US), from Michoaca´n, and Fisher s.n. (MO!), from Veracruz. The former appears to us to be typical P. nobilis (var. nobilis of Yatskievych, 1996); the latter has well formed spores and appears to be more or less typical of P. remotispora (P. nobilis var. remotispora of Yatskievych (1996)). These hybrids were said to have intermediate morphology and casually anastomosing veins. However, for

PHANEROPHLEBIA REMOTISPORA

nearly the entire range of these two taxa, there is little or no difficulty in distinguishing them. In fact, the two species seem to be more easily distinguished than does P. gastonyi from P. juglandifolia, both of which were treated as species by Yatskievych. The free veins, higher elevation, generally narrower pinnae, and soral position all serve to distinguish P. remotispora from P. nobilis. Seldom, if ever, do they grow together, a fact acknowledged by Yatskievych (1996). Matuda 1365 (US), from Maltrata, Edo. Veracruz, is very atypical in having only 3 pairs of lateral pinnae and more cordate proximal pinnae. Maxon had earlier determined it as P. pumila. Yatskievych identified it as P. nobilis, and we tentatively do so also. Hinton 403, cited above as P. nobilis, was determined by Yatskievych (1996) as P. nobilis var. remotispora. Matuda 4688, from Chiapas, was determined by Smith (1981) as a very small, possibly precociously fertile specimen of P. macrosora, but Yatskievych believed it to be P. nobilis; we now concur.

6. PHANEROPHLEBIA PUMILA Phanerophlebia pumila (M. Martens & Galeotti) Fe´e, Me´m. Foug. 5: 282. 1852. Figs. 222F–H, J–L. Aspidium pumilum M. Martens & Galeotti, Me´m. Foug. Mexique 64, pl. 17, f. 1. 1842. Cyrtomium pumilum (M. Martens & Galeotti) C. V. Morton, Amer. Fern J. 47: 54. 1957. Type. Mexico. Oaxaca: “Llano Verde et del Carrizal,” Galeotti 6251 (BR!, photos LL!, US!; isotype P!). Phanerophlebia lindenii E. Fourn., Mexic. Pl. 1: 100, pl. 4. 1872. Type. Mexico. Chiapas: “In pineto pr. Ciudad Real” [San Cristo´bal Las Casas], Linden s.n. (P!; isotype G, photo US!). Linden 1551 (K!) is probably part of the same gathering.

Rhizomes short, erect, unbranched; rhizome scales concolorous, brown, ovate, 3–5 ⫻ 2–3 mm, margins ciliate-denticulate; fronds mostly 10–30(–45) cm long; stipes usually 1⁄2 or more of the frond length, stramineous to tan, with dense, persistent, lanceolate scales toward bases, scales becoming filiform distally, sometimes abraded, (3–)5–10 ⫻ 0.2–1.5 mm; blades ovate, subcoriaceous, (3.5–)6–20(–35) cm long; rachises with scattered to dense, filiform scales; buds absent from axils of distal pinnae; pinnae 0–3(–4) lateral pairs, ovate to lanceolate, straight or subfalcate, to 1.5–8(–12) ⫻ 1–2(–3) cm, the terminal usually the largest, bases rounded to truncate or subcordate, apices rounded to acute, lowermost pinnae inequilateral, excavate basiscopically, sometimes subauriculate acroscopically, margins spinulose throughout; veins free or only occasionally casually anastomosing near pinna margins, 2–3-branched; sori in 1–3 series between costae and margins, inner row (1–)2–5 mm from costa, outer row 2–4 mm from margin; indusia subpersistent, tan, 0.8–1.5 mm diam., flat or convex, not umbonate, erose at margins; 2n⫽164 (Chis, Oax).

459

Distribution. Epipetric and in rock crevices, limestone sinks, and on cliffs, in moist forests; 2000–2950 m. Mexico; Guat. Selected Specimens Examined. Chis (Breedlove 31816, DS, MEXU, TEX; 55521, DS). Gro (Nelson 2222, UC, US). Mich (Hinton 12428, LL, MEXU, US, 15935, ENCB, MEXU, NY, US). Oax (Breckon & Christman 622, IEB, UC; Gereau & Martin 2011, NY; Hallberg 848, MEXU; Yatskievych 85-139, ARIZ). Unverified, Doubtful, or Mistaken Reports. Qro (Ferna´ndez 4585, ENCB, Rzedowski 42551, IEB, and Rzedowski 46146, QMEX, cited by Arreguı´n et al., 2001, none seen, but Rzedowski 42551 is P. nobilis at ENCB, UAMIZ).

Phanerophlebia pumila is distinguished by having free veins, large persistent indusia, coriaceous texture, and few pairs of cordate-based pinnae. Rzedowski 42551 and 44614b (IEB), from Quere´tero, were cited by Dı´az-Barriga and Palacios-Rios (1992) as P. pumila, but are merely juvenile sterile or nearly sterile forms of P. nobilis, with 1–4 pairs of lateral pinnae; the pinnae are less coriaceous and not cordate, thus agreeing better with P. nobilis, which is common in Quere´tero.

7. PHANEROPHLEBIA REMOTISPORA Phanerophlebia remotispora E. Fourn., Mexic. Pl. 1: 100. 1872. Figs. 222S, T. Cyrtomium remotispora (E. Fourn.) C. V. Morton, Amer. Fern J. 47: 54. 1957. Phanerophlebia nobilis (Schltdl. & Cham.) C. Presl var. remotispora (E. Fourn.) Yatsk., Novon 2: 446. 1992. Type. Mexico. Veracruz: Ixhuatlancillo, Bourgeau 2349 (P!; isotypes BM!, GH!, K!, MICH, MO, LE!, NY!, P!, UC!, US!).

Rhizomes suberect to erect, usually unbranched; rhizome scales concolorous, brown, ovate to ovate-lanceolate, 3.5–10 ⫻ 1–3 (–4) mm, margins denticulate to ciliate; fronds mostly 60–125 cm long; stipes 1⁄3–1⁄2 the frond length, with dense brownish, ovate to lanceolate scales at bases; blades ovate to ovatelanceolate, 35–65 cm long; rachises glabrescent or with sparse, thin, curled, fibrillose scales 2–4 mm long, mostly at pinna bases; buds absent from axils of distal pinnae; pinnae 8–18 lateral pairs, linear-lanceolate, straight to often falcate, mostly 9–18 ⫻ 1.5–3.5 cm, bases rounded to broadly cuneate, apices attenuate, lowermost pinnae inequilateral, not auriculate, distal pinnae equilat-

460

PHLEBODIUM

eral, margins spinulose most of their length, decidedly so at pinna apices; veins anastomosing about 2⁄3 the way to pinna margins; sori in 2–4(–5) series between costae and pinna margins, inner row mostly (2–)3–5 mm from costa, outer row 1–3 mm from margin; indusia tan, caducous, 0.8 mm diam., flat or concave, not umbonate; 2n⫽82 (Chis, Hgo, Ver).

caducous scales similar to those of rhizomes; blades ovate to lanceolate, 13–50 cm long; rachises with sparse, tan, hair-like, twisted scales; buds absent from axils of distal pinnae; pinnae 7–18 pairs, ovate-oblong to usually linear-lanceolate, falcate, 4–15 ⫻ 1.5–2.6 cm, bases rounded to broadly cuneate to truncate, apices attenuate, lowermost pinnae inequilateral, excavate basiscopically, not auriculate acroscopically, margins spinulose-serrate throughout; veins free or only occasionally and sporadically anastomosing, 1– 3-branched; sori in 2–3 series between costae and pinna margins, inner row 2–3 mm from costa, outer row often submarginal; indusia firm, light brown, 0.6–1 mm diam., convex with a raised darker umbo centrally, persistent and not shriveling at maturity; 2n⫽82 (USA, NL, SLP).

Distribution. Terrestrial or epipetric in wet forests, often on calcareous rocks (limestone sinks); 250–2200 m. Mexico; Guat. Selected Specimens Examined. Chis (Breedlove 21363, LL, NY; Yatskievych 85-186, ARIZ, NY, UAMIZ, UC). Hgo (Yatskievych 83-128, ENCB, NY). Me´x (Guillemin s.n., NY). Oax (Mickel 7248, UC). Pue (Lorea 1095, FCME). Qro (Lorea 686, FCME, IEB). SLP (Rzedowski 10571, ENCB, NY). Ver (Copeland herb. 40, MEXU, UC; Duran E. & Burgos 739, XAL). Unverified, Doubtful, or Mistaken Reports. Mich (reported by Mickel & Beitel, 1988, but not verified, the specimen used as a basis is probably P. nobilis; reported by Tejero-Dı´ez & Arreguı´n-Sa´nchez, 2004, but not verified).

From P. nobilis, P. remotispora differs in having anastomosing veins, first series of sori usually more distant from the costae, broader pinnae, and occurrence at generally lower elevations. It appears to be absent from western Mexico, where P. nobilis is relatively common. This appears to be a classic case of ecogeographic speciation, with P. remotispora very common on the Caribbean slope and in southern Mexico, and P. nobilis very common on the Pacific slope; however, there is slight interdigitation of the ranges (see map by Yatskievych, 1996: 191). See discussion under P. nobilis for additional comments.

8. PHANEROPHLEBIA UMBONATA Phanerophlebia umbonata Underw., Bull. Torrey Bot. Club 26: 211. 1899. Figs. 221G, H, J, K. Cyrtomium umbonatum (Underw.) C. V. Morton, Amer. Fern J. 47: 54. 1957. Type. Mexico. Nuevo Leo´n: Near Monterry, Pringle 1982 (NY; isotypes GH!, K, PH, UC!, US!, YU).

Rhizomes short-creeping to ascending, often branched; rhizome scales concolorous, brown, ovate, 2.5–4.5 ⫻ 2–4 mm, margins ciliate; fronds (20–)40–90 cm long; stipes 1⁄3–1⁄2 the frond length, stramineous, moderately scaly with tan, appressed often

Distribution. Terrestrial or epipetric on limestone cliffs, in moist, mesic, sheltered canyons and ravines, pine-oak forests, often on calcareous rocks; 550–2000 m. USA (Tex); Mexico. Selected Specimens Examined. Coah (Wynd & Mueller 349, ARIZ, MEXU, NY; Lyonnet 421000001, CAS, MEXU; Stewart 1533, LL, US). NL (Pringle 13739, ARIZ, LL, TEX, UC, US). Qro (Rubio 1492, UAMIZ). SLP (Mickel 563, ENCB, NY; Pringle 3403, UAMIZ, UC). Tam (Bartlett 10095, NY, US; Stanford et al. 1069, ARIZ, NY, UC; Valde´s R. 1940, NY, TEX). Unverified, Doubtful, or Mistaken Reports. Chih (Jones s.n., CAS, cited by Yatskievych, 1996, but not verified).

This species is distinguished by having free veins, usually 10 or more pinna pairs, exauriculate pinnae, and firm, persistent, large indusia with a central umbo (nipple), hence the specific epithet umbonata. Phanerophlebia umbonata has, in general, more persistently scaly rachises than P. nobilis, a close relative. The range of P. umbonata is more or less confined to northeastern and north-central Mexico and southwestern Texas. Yatskievych (1996) cited Gentry 8119 (MICH, UC), from Sonora, as representing this species, but the UC specimen seems to us to be P. nobilis.

95. PHLEBODIUM Phlebodium (R. Br.) J. Sm., J. Bot. (Hooker) 4: 58. Jul 1841. Polypodium sect. Phlebodium R. Br. in Horsfield, Pl. Jav. Rar.

PHLEBODIUM AREOLATUM

4. 1838 (see Smith, 1981: 168, for discussion of problems in typification). Chrysopteris Link, Fil. Spec. 120. Sep 1841, nom. superfl. for Phlebodium.

Epiphytic or epipetric; rhizomes creeping, branched, glaucous; rhizome scales concolorous, brown to golden, nonclathrate, peltately attached, surfaces glabrous, margins ciliate; fronds monomorphic, articulate; stipes brown to stramineous, channelled adaxially; blades deeply pinnatifid to pinnatisect with rounded sinuses, broadly oblong to subdeltate; pinnae broad, entire, chartaceous to coriaceous, usually glabrous, often glaucous; veins netted, row of costal areoles without free included veins, then one to several rows of additional areoles, these with or without free included veins; sori round, exindusiate, within areole at the union of merging, included veins, in 1 to 7 rows between costa and segment margin, paraphyses absent; sporangia glabrous; spores bilateral, coarsely tuberculate; x⫽37. Phlebodium, as commonly construed, is a segregate of Polypodium s.l. It occurs in the New World tropics and subtropics and comprises about four species. The double included veins running to the sorus distinguish this small genus from species of Polypodium, which have free veins or a single included vein, if the veins are netted. Key to the Mexican Species of Phlebodium 1. Sori in one row on each side of the costae; pinnae 10–30 mm wide; (170–)550–2500 m. ........................................ 1. P. areolatum. 1. Sori in several rows on each side of the costae; pinnae 35– 45 mm wide; 0–500 m elevation. ............................... 2. P. decumanum.

1. PHLEBODIUM AREOLATUM Phlebodium areolatum (Humb. & Bonpl. ex Willd.) J. Sm., J. Bot. (Hooker) 4: 59. 1841. Figs. 223A–D. Polypodium areolatum Humb. & Bonpl. ex Willd., Sp. Pl., ed. 4, 5(1): 172. 1810. Goniophlebium areolatum (Humb. & Bonpl. ex Willd.) C. Presl, Suppl. Tent. Pterid. 186. 1836. Polypodium aureum L. var. areolatum (Humb. & Bonpl. ex Willd.) Baker in Hooker & Baker, Syn. Fil. 347. 1867. Type. Venezuela. Cumana´, Humboldt & Bonpland s.n. (B-Willd. 19645!). Polypodium araneosum M. Martens & Galeotti, Me´m. Foug. Mexique 33. 1842. Phlebodium aureum (L.) J. Sm. var. araneosum (M. Martens & Galeotti) L. D. Go´mez, Brenesia 18: 163. 1981. Phlebodium araneosum (M. Martens & Galeotti) Mickel & Beitel, Mem. New York Bot. Gard. 46: 277. 1988. Type. Mexico. Oaxaca: Villa Alta, Galeotti 6460 (BR!, photo NY!; isotypes BR!, US!; US! photo ex BR). Polypodium fulvum M. Martens & Galeotti, Me´m. Foug. Mexique 33. 1842. Type. Mexico. Oaxaca: “Environs de Villa-Alta,” “Monte de Tanga,” Galeotti 6463 (BR!, US! photo ex BR). Polypodium glaucinum M. Martens & Galeotti, Me´m. Foug. Mexique 32. 1842. Type. Mexico. “El Sabino pre`s d’Izmiquilpan,” Galeotti 6430 (BR!; US! photo ex BR).

Rhizomes creeping, 4–10 mm diam., pruinose; rhizome scales 6–10 mm long, reddish brown, lanceolate, each with enlarged, round, peltate base, margins denticulate throughout, especially on scale tips, darker at point of attachment; fronds (20–)28–90

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cm long; stipes 1⁄3–1⁄2 the frond length, stramineous to brown, glabrous; blades deeply pinnatifid, broadly oblong to ovatedeltate, bases not much narrowed, 20–40 cm wide; pinnae 4–13 pairs, 10–30 mm wide, linear-oblong to linear deltate, obtuse to acuminate, glabrous, often glaucous, margins entire, occasionally with low teeth or crenate; veins netted, with double included veinlets or infrequently single; sori round, medial or submedial (served by two excurrent veins), one row on each side of the costae; 2n⫽74 (Oax).

Distribution. Epiphytic, epipetric or terrestrial; lower montane rain forests, montane rain forests, pine-oak-Liquidambar forests, seasonal evergreen forests, tropical deciduous forests, pineoak forests; (170–)550–2500 m. USA (Fla); Mexico; Guat, Hond, Salv, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol, n Arg. Selected Specimens Examined. Ags (Garcı´a R. 2793, IEB). Chih (Bye 2660, MEXU). Chis (Breedlove 31497, DS, MEXU). Col (Lott et al. 2959, CAS, MEXU). DF (Arse`ne 10664, US). Dgo (Bravo 1103, IEB, MEXU). Gro (Acosta & Lo´pez 48, MEXU). Gto (Galva´n & Galva´n 2862, ENCB, IEB, MEXU). Hgo (Bartolomew 4039, CAS). Jal ( Wilbur & Wilbur 1441, MEXU). Me´x (Hinton 1529, MEXU). Mich (Pringle 3383, MEXU). Mor (Sa´nchez-Mejorada 162, MEXU). Nay (Calzada et al. 18820, MEXU). NL (Rodrı´guez A. herb. no. 010205, UNL). Oax (Martı´nez-C. 475, 898, MEXU, NY). Pue (Rzedowski 32421, ENCB, MEXU). Qro (Rubio 1301 IEB, MEXU). Sin (Vega & Palazuelos 827, MEXU). SLP (King 4283, US). Tam (Martı´nez 1460, IEB). Ver (Ventura A. 16807, XAL). Unverified, Doubtful, or Mistaken Reports. Camp (Uca´n Ek’ et al. 5917, UADY, cited by Palacios-Rios, 2002a, but not verified). Tab (reported by Magan˜a, 1992, but not verified).

This species has often been included in P. aureum (L.) J. Sm., e.g., by Smith (1981). The plants from Mexico and Central America have a single row of sori on each side of the costa. Proctor (1985) reported that specimens with sori in single rows predominate in the Greater Antilles, whereas Polypodium (Phlebodium) aureum L. (Sp. Pl. 2: 1087. 1753. Type. America, LINN 1251.10), with 2 rows of sori on each side of the costae, is common in Florida, Bahamas, Puerto Rico, the Lesser Antilles, and South America. Proctor and Evans (pers. comm.) reported that in Puerto Rico, where P. aureum, P. areolatum, and P. decumanum co-occur, both P. areolatum and P. decumanum appear to be diploids. Phle-

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PILULARIA

bodium aureum is their fertile, allotetraploid hybrid, and at least one sterile, triploid backcross hybrid was reported. Among the alleged synonyms of Phlebodium aureum, Polypodium leucatomas Poir. in Lamarck (Encycl. 5: 516. 1804. Type. French Guiana, LeBlond s.n. (P!-Lam., microfiche NY!) and Polypodium sporadocarpum Willd. (Sp. Pl., ed. 4, 5(1): 171. 1810. Type. B-Willd. 19644) both have more than one row of sori and therefore do not represent P. areolatum. Reports of Phlebodium aureum (L.) J. Sm. in Mexico generally apply to P. areolatum. Phlebodium araneosum has sometimes been treated as a distinct species (e.g., Mickel & Beitel, 1988), but the only distinguishing character is the presence of blade scales, and there is every transition between it and P. areolatum. Polypodium pseudoaureum Cav. (Descr. Pl. 2: 247. 1801) may possibly be an earlier name for this taxon, but since the type is sterile, there was no locality given for Ne´e’s specimen (collected on a trip around the world), and the pinnae are shorter than those of P. areolatum, use of this name should wait until the type can be studied in detail. The identity of P. pseudoaureum could be resolved by the study of stomate length. Phlebodium aureum would have longer stomates as it is a tetraploid.

2. PHLEBODIUM DECUMANUM Phlebodium decumanum (Willd.) J. Sm., J. Bot. (Hooker) 4: 59. 1841. Figs. 223E, F. Polypodium decumanum Willd., Sp. Pl., ed. 4, 5(1): 170. 1810. Type. Brazil. Hoffmannsegg s.n. (B-Willd. 19640!, frag. NY!).

Similar to P. areolatum except: sori in 4–7 rows on both sides of costae; rhizome scales larger (10–15 mm) and more twisted; clavate hairs in the abaxial vein axils minute (less than 0.1 mm long); pinnae wider (35–45 mm); 2n⫽74 (Trin).

Selected Specimens Examined. Chis (Breedlove 47197, CAS; Rovirosa 54, US). Gro (Ochoa F. 58, FCME). Oax (Herna´ndez M. & Orozco 1639, MEXU; Martı´nez C. 718, MICH, US). QR (Calzada et al. 6888, XAL). SLP (Carranza & Dı´az B. 4730, IEB, MEXU). Tab (Gilly & Herna´ndez X. 68, US). Ver (Sa´nchez-Mejorada 633, MEXU, US; Valdivia Q. 2031, XAL). Unverified, Doubtful, or Mistaken Reports. Camp (Zamora C. & Me´ndez D. 4995, UACAM, XAL, cited by Palacios-Rios, 2002a, but not verified). Yuc (Lundell 4835, MO, cited by Moran, in Davidse et al., 1995, but not verified; Castillo R. 106, UADY, cited by PalaciosRios, 2002b, but not verified).

96. PILULARIA Pilularia L., Sp. Pl. 2: 1100. 1753; Gen. Pl., ed. 5, 486. 1754. Type: Pilularia globulifera L. Aquatic or in shallow seasonally drying pools, rooted in soil; rhizomes long-creeping, slender, bearing hairs, roots arising at nodes; leaves filiform, circinate, terete, lacking expanded blades, with a single vascular strand, glabrous; sporocarps produced only on emergent plants, globose, indurate, borne singly at or just above ground level on unbranched stalks arising at bases of petioles, attached laterally to stalk apex (attached portion called raphe), lacking teeth, densely to sparsely hairy, hairs abraded with age, dehiscing into 4 valves; sori 2–4 per sporocarp; sporangia of two kinds: ovoid megasporangia, containing single megaspores, and fusiform microsporangia, containing numerous microspores; indusia thin, enclosing the sporangia; megaspores ovoid, filling the megasporangia, with a prominent papilla-like laesura, surface rugulose; microspores tetrahedral-globose, surface densely and coarsely rugose; x⫽10. Pilularia comprises about five very similar species, distributed in North America, South America, Europe, Australia, and New Zealand. The discreteness and interrelationships of species within Pilularia need re-investigation using modern methods. The genus is closely related to two other genera of heterosporous ferns, Marsilea (which see, for discussion of relationships within the family) and Regnellidium, the latter monotypic and restricted to southeastern Brazil and northeastern Argentina. Together, these three genera constitute the family Marsileaceae, which is now believed to be allied to the other two heterosporous fern genera, Salvinia and Azolla (Pryer et al., 1995; Pryer, 1999). References Dennis, W. M. & D. H. Webb. 1981. The distribution of Pilularia americana A. Braun (Marsileaceae) in North America, north of Mexico. Sida 9: 19–24; Johnson, D. M. 1993. Pilularia. p. 335 in FNA Ed. Comm., 1993; Pryer, K. M. 1999. Phylogeny of marsileaceous ferns and relationships of the fossil Hydropteris pinnata reconsidered. Int. J. Pl. Sci.160: 931–954.

Distribution. Epiphytic, often on palms, mostly on Gulf coastal plain; 0–500 m. USA (Fla); Mexico; Guat, Bel, Hond, CR; Gr & L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, s Braz, Bol, Parag, Arg, Uru.

1. PILULARIA AMERICANA Pilularia americana A. Braun, Monatsber. Ko¨nigl. Preuss. Akad. Wiss. Berlin 1863: 435. 1864. Type. U.S.A. Arkansas: Fort

PITYROGRAMMA

Smith, Nuttall s.n. (herbarium uncertain, but Nuttall collections known to be at BM, GH, LIV, and PH, among others; isotype NY!). Figs. 224A–C. Plants forming dense clones; rhizomes ca. 1 mm diam.; leaves 1.5–10 cm, sparsely pubescent to glabrescent; sporocarp stalks 1– 3 mm long; sporocarps globose, 1.6–2.7 mm diam., covered with matted hairs until mature; raphes (attached portion of sporocarp stalk) very short, lacking teeth; sori 4 per sporocarp; 2n⫽20 (Arkansas).

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9 7 . P I T Y R O GR A M M A Pityrogramma Link, Handbuch 3: 19. 1833. Type: Pityrogramma chrysophylla (Sw.) Link [⬅ Acrostichum chrysophyllum Sw.]. Trismeria Fe´e, Me´m. foug. 5: 164, pl. 14, figs. 1—2. 1852. Lectotype (chosen by Maxon, Pteridophyt. Porto Rico. 436. 1926): T. aurea Fee [⫽ Pityrogramma trifoliata (L.) R. Tryon]. For additional synonymy, see Lellinger, Mem. New York Bot. Gard. 23: 7. 1972.

Terrestrial; rhizomes erect, woody; rhizome scales concolorous; fronds medium-sized, clumped; stipes brown to atropurpureous, mostly glabrous; blades 1–3-pinnate, chartaceous to coriaceous, usually with white or yellow farina abaxially, infrequently with hairs (not ours); veins free; sporangia along veins, veins obscured by farina so sporangia appear in indistinct lines or scattered black dots; indusia and paraphyses absent; spores tetrahedral-globose, dark; x⫽30, 58, 60.

Distribution. In shallow ponds and seasonal pools, also in moist soils in semidisturbed areas and pastures; 300–2000 m. USA (scattered but widespread); Mexico; Col, Ven, s Braz, Bol, Chile, Arg. Specimens Examined. BCN (Thorne & Charlton 60050, ENCB; Wiggins 10087, DS, ENCB). Dgo (Taylor & Montgomery s.n., 9 Nov 1997, NY).

Most likely, this species is more common and widespread in Mexico than the relatively few collections suggest, at least in northwestern Mexico. Because of its inconspicuous and unfernlike appearance, probably it has been overlooked by collectors.

Pityrogramma is a genus of about 17 species, mostly of the American tropics; there are 12 species in America and 4 in Mexico. They are common plants of roadsides and other disturbed soil banks, of low to high elevations. One species of western North America, Pityrogramma triangularis, is now placed in Pentagramma, a genus differing from Pityrogramma in size, habit, habitat, chemistry, and spore morphology. Trismeria is often segregated from Pityrogramma because of its distinct habit, blade dissection, and spores. R. Tryon (1962) reported hybridization between Trismeria and species of Pityrogramma. Pityrogramma is related to Anogramma, less so to Eriosorus and Jamesonia, but is distinct by the combination of farinose abaxial blade surfaces, pinnately divided blades, gymnogrammoid sori, and erect woody rhizome. Reference Tryon, R. M. 1962. Taxonomic fern notes. II. Pityrogramma (including Trismeria) and Anogramma. Contr. Gray Herb. 189: 52–76.

Key to the Mexican Species of Pityrogramma 1. Distal pinnae entire; proximal pinnae entire to trifoliate, the segments linear-oblong. .......................................................................... 4. P. trifoliata. 1. Distal and proximal pinnae with numerous, pinnately arranged lobes or segments. 2. Stipe bases densely clothed with golden brown scales and rare, short, clavate hairs 0.1 mm; rhizome scales 8–12 ⫻ 0.5–1 mm, golden brown, becoming black with age; pinnae inequilateral, basiscopically exaggerated; pinnules on both sides of pinna at nearly right angles to costae; spores brown; blades pinnate-pinnatifid (to bipinnate-pinnatifid), broadest at bases; middle to high elevation (900–2000 m). ....................................................................................................................................................................... 3. P. ebenea. 2. Stipe bases with sparse golden brown scales or glabrous; rhizome scales 2–5 ⫻ 0.1–0.8 mm, golden brown or light brown; pinnae equilateral; pinnules ascending; spores reddish brown; blades bipinnate-pinnatifid to tripinnate, lanceolate; low to midelevation (50–1150 m). 3. Ultimate segments lanceolate; fronds medium-sized to large, to 120 cm long; rhizome scales 3–5 ⫻ 0.5–0.8 mm, dark golden brown with tooth-like undulations. .................................................................................................................................... 1. P. calomelanos. 3. Ultimate segments ovate; fronds medium-sized, less than 40 cm long; rhizome scales 2–3 ⫻ 0.1–0.3 mm, light brown, margins entire, tips hair-like, uniseriate, often forked. ..................................................................................................................... 2. P. dealbata.

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PITYROGRAMMA

1. PITYROGRAMMA CALOMELANOS Pityrogramma calomelanos (L.) Link, Handbuch 3: 20. 1833. Figs. 225A, B. Acrostichum calomelanos L., Sp. Pl. 2: 1072. 1753. Type. LINN sheet 1245.19, photos A, GH. Ceropteris serrata Fe´e, Me´m. Foug. 8: 81. 1857. Type. Mexico. [Veracruz:] Orizaba, Schaffner 162 (P, not found). For additional synonymy, see R. M. Tryon (1962).

Rhizomes erect; rhizome scales 3–5 ⫻ 0.5–0.8 mm, concolorous, dark golden brown, margins with acute, tooth-like undulations, not prolonged into hair-like uniseriate tips; fronds to 120 ⫻ 20 cm, clumped; stipes 1⁄3–1⁄2 the frond length, dark reddish brown to black, glabrous except for sparse golden brown scales at bases; blades lanceolate, bipinnate-pinnatifid to tripinnate, pinnae equilateral; ultimate segments ascending, lanceolate, texture thin, wrinkled, glabrous adaxially, white-farinose abaxially, margins toothed, slightly recurved; veins free, prominent; sori elongate along the veins, appearing as black specks scattered in the white farina; spores reddish brown; 2n⫽240 (Jam, Old World).

Distribution. Terrestrial, in open areas, open rocky slopes, roadbanks, sandy soils near streams, among rocks along streams, rarely in forests, pine-oak forests, tropical deciduous forests, riparian vegetation; 0–1200(–1600) m. USA (Fla); Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Bah, Gr & L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Galapagos, Peru, Braz, Bol, Parag, Arg; introduced in the Old World. Selected Specimens Examined. Chis (Breedlove 19930, DS, NY). Col (Palmer 1218, NY, US). Gro (Palmer 273, CAS, NY, US). Hgo (Copeland herb. 103, UC, US). Jal (Palafox 5394, GUADA, IEB, UC). Me´x (Hinton 7326, NY, US). Mich (Hinton 16295, DS, NY, UC). Nay (Gentry 19511, NY, US). Oax (Mickel 6668, NY, UC). Qro (Rzedowski 42828, ENCB, IEB). QR (Sanders et al. 9993, UC, UCR). Sin (Correll & Correll 28846, US). SLP (Storer 151, US). Tab (Bautista & Cowan 2335, CAS, NY). Tam (Haynes 4302, NY). Ver (Smith 2103, MEXU, MO, NY, UC).

Pityrogramma calomelanos is distinguished from P. ebenea by its finer blade dissection, ascending segments, and occurrence at lower elevations.

2. PITYROGRAMMA DEALBATA ˇ eske´ Akad. Pityrogramma dealbata (C. Presl) Domin, Rozpr. C Veˇd., Trˇ. 2, Veˇdy Mat. Prˇir. 51(15): 7. 1942 [1941]. Figs. 226A, B. Gymnogramma dealbata C. Presl, Reliq. Haenk. 1(1): 18, t. 3, f. 1. 1825. Lectotype (chosen by R. M. Tryon, 1962: 67). Panama, Haenke s.n. (PR–sheet 24358a, photo GH). Ceropteris schaffneri Fe´e, Me´m. Foug. 8: 80. 1857. Pityrogramma schaffneri (Fe´e) Weath., Contr. Gray Herb. 114: 25. 1936. Syntypes. Mexico. Veracruz: “pre`s d’Orizaba, dans les lieux secs et expose´s au soleil,” Schaffner 165a, 165b (P, not found).

Rhizomes erect; rhizome scales 2–3 ⫻ 0.1–0.3 mm, concolorous, light brown, entire, prolonged into long, hair-like, uniseriate, often divided tips; fronds less than 40 ⫻ 12 cm, clumped; stipes 1⁄3–1⁄2 the frond length, dark reddish brown, glabrous except for rare golden brown scales at extreme bases; blades lanceolate, bipinnate-pinnatifid to tripinnate, pinnae equilateral; ultimate segments ascending, cuneate, ovate, coriaceous, glabrous, white-farinose abaxially, rarely yellow, margins toothed, slightly recurved; veins free, depressed adaxially; sori elongate along the veins, appearing as black specks scattered in white farina; spores reddish brown.

Distribution. Roadside banks and wooded slopes of the Pacific slope, pine-oak forests, riparian situations with seasonal evergreen forests; 100–1300 m. Mexico; Guat, Hond, Salv, Nic, CR, Pan. Selected Specimens Examined. Chis (Breedlove 20648, DS, NY; Marcks & Marcks 884, ENCB, WIS). Col (Jones 515, CAS, DS, NY, UC, US). Gro (Lorea 4600, FCME). Jal (Pringle 4377, MEXU, NY, UC, US). Mich (Nelson 6937, US). Nay (Mexia 510, CAS, US). Oax (Mickel 6276, NY, UC; Roe et al. 544, ENCB, US). Ver (Pringle 6078, MEXU, MO, NY, UC, US).

Pityrogramma dealbata resembles P. calomelanos in having bipinnate-pinnatifid to tripinnate blades and occurring at low elevations, but is distinct in its cuneate, ovate segments, smaller fronds, and smaller rhizome scales.

PITYROGRAMMA TRIFOLIATA

3. PITYROGRAMMA EBENEA Pityrogramma ebenea (L.) Proctor, Brit. Fern Gaz. 9: 219. 1965. Figs. 226C–E. Acrostichum ebeneum L., Sp. Pl. 2: 1071. 1753. Lectotype (chosen by Proctor, Brit. Fern Gaz. 9: 219. 1965). Jamaica. s. coll. s.n. (LINN-1245.14). Acrostichum tartareum Cav., Descr. Pl. 242. 1801. Pityrogramma tartarea (Cav.) Maxon, Contr. U.S. Natl. Herb. 17: 173. 1913. Type. Peru. Near Guamantanga, Ne´e s.n. (MA; isotype P, not found, photo US). Pityrogramma tripinnata Domin, Spisy Prˇir. Fak. Karlovy Univ. 88: 7. 1928. Type. Mexico. San Luis Potosı´: Parry & Palmer 1004 (K!, photo GH, NY!, UC!; isotype NY!, US!). For additional synonymy, see R. Tryon (1962).

Rhizomes erect; rhizome scales 8–12 ⫻ 0.5–1 mm, concolorous, golden brown, black with age, entire, prolonged into hair-like, uniseriate tips; fronds to 100 cm long, clumped; stipes 1⁄3–1⁄2 the frond length, dark reddish brown, glabrous except bases with dense golden brown scales varying to hair-like scales 2 mm long and sparse, minute clavate hairs less than 0.1 mm long; blades deltate to narrowly triangular, to 40 cm wide, broadest at bases, 1–2-pinnate-pinnatifid; pinnae inequilateral, exaggerated basiscopically, segments perpendicular to costae, ovate, glabrous adaxially, coriaceous, white-farinose abaxially, margins strongly recurved; veins free, depressed adaxially; sori elongate along veins, appearing as black specks scattered in the white farina; spores brown; 2n⫽240 (Jam).

Distribution. Terrestrial on exposed banks, montane rain forests or cloud forests, pine or pine-oak forests; 900–2600 m. Mexico; Guat, Hond, Salv, Nic, CR, Pan; Gr Ant; Col, Ven, Ec, Galapagos, Peru, Braz, Bol. Selected Specimens Examined. Chis (Breedlove 23327, DS, NY). Col (Orcutt 4699, NY). DF (Schaffner 124, in 1875, NY). Dgo (Breedlove 18909, CAS, NY). Gro (Hinton 14190, NY, US). Gto (Rzedowski 51253, IEB). Hgo (Frye 2545, US). Jal (Mexia 1827, CAS, DS, UC, US). Me´x (Hinton 7434, MEXU, NY, US). Mich (Pringle 13642, CAS, UC). Mor (Lyonnet 1325, MEXU). Nay (Jones 23480, CAS, NY, UC). Oax (Mickel 5020, NY, UC). Pue (Fro¨derstro¨m & Hulte´n 747, NY). Qro (Rzedowski 48440, IEB). Rev (Dı´az-Luna 18058, GUADA, NY). SLP (Parry & Palmer 1040, NY, US). Sin (Keil & Canne 8847, NY). Ver (Ventura A. 230, ENCB).

Although P. ebenea generally occurs at higher elevations than P. calomelanos, the two species occasionally grow together in

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Mexico. Pityrogramma ebenea is distinguished by the pinnules being broadest at the base and held nearly at right angles to the costae.

4. PITYROGRAMMA TRIFOLIATA Pityrogramma trifoliata (L.) R. M. Tryon, Contr. Gray Herb. 189: 68. 1962. Figs. 226F–H, J, K. Acrostichum trifoliatum L., Sp. Pl. 2: 1070. 1753. Trismeria trifoliata (L.) Diels in Engl. & Prantl, Nat. Pflanzenfam. 1(4): 165. 1899. Lectotype (chosen by Proctor 1985: 203). Sloane, Voy. Jamaica 1: t. 45, f. 2. 1707, based on a plant from Jamaica.

Rhizomes erect; rhizome scales bicolorous, centers dark reddish brown, thickened, margins narrow, light brown, 3–4 ⫻ 0.5–0.8 mm; fronds 0.5–2.4 m long, clumped; stipes 1/5–1⁄3 the frond length, with scales at base numerous, castaneous, concolorous, light brown, to 5 ⫻ 1–1.5 mm, and scattered hairs 1 mm long; blades pinnate at tip to usually bipinnate at bases, to 15 cm wide, pinnae usually divided toward the frond base into (2–)3 pinnules oriented out of the plane of the frond, the segments linearoblong, irregularly serrulate, thin with prominent veins, glabrous adaxially, glabrous to white-farinose abaxially, non-articulate, sterile ones to 90 ⫻ 8–12 mm, fertile ones to 90 ⫻ 3–5 mm, margins recurved, unmodified; sori irregularly elongate along the veins from midribs to margins; spores reddish brown; 2n⫽120 (Jam).

Distribution. Wet open banks or meadows or seepage areas, tropical deciduous forests, pine-oak forests, coffee plantations; 100–1500 m. USA (Fla); Mexico; Guat, Hond, Salv, Nic, CR, Pan; Bah, Gr Ant; Col, Ven, Ec, Peru, Braz, Bol, Chile, Parag, n Arg, Uru. Selected Specimens Examined. BCS (Brandegee 655, NY, UC). Chis (Breedlove 14908, DS, ENCB, NY, US). Gro (Tenorio L. et al. 479, MEXU). Hgo (Lautner L01/44, UC). Me´x (Hinton 8464, NY, US). Mich (Torres 13646, IEB, XAL). Oax (Camp 2459, ENCB, NY, UC; Mickel 6662, NY). Pue (Lyonnet 510900009, MEXU). Ver (Ventura A. 4352, ENCB, MEXU, NY, XAL).

Pityrogramma trifoliata is often placed in a separate genus Trismeria. It crosses with P. calomelanos in Costa Rica, Panama, and Argentina.

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PLAGIOGYRIA

98 . PL A G I O G YR I A

resemblance to certain dimorphic species of Blechnum, under which name specimens often are mistakenly identified.

Plagiogyria (Kunze) Mett., Abh. Senckenberg. Naturf. Ges. 2: 265. 1858. Lomaria sect. Plagiogyria Kunze, Farrnkra¨uter 2: 61. 1850. Type: Plagiogyria euphlebia (Kunze) Mett. [⬅ Lomaria euphlebia Kunze].

References

Terrestrial; rhizomes rather stout, suberect, dictyostelic, lacking scales and hairs, sometimes stoloniferous; roots wiry; fronds dimorphic, the fertile ones more erect, with longer stipes, and with narrower pinnae, the sterile ones pinnatisect to barely pinnate and the pinnae adnate, crown-forming, erect to arching, glabrous at maturity, young fronds covered with a gelatinous secretion; stipes with 3 vascular bundles at base, fusing distally to a V- or U-shape, stipe bases flared or slightly expanded, blackish, persistent, subtriangular in cross section, covering the rhizomes, bearing 2 rows of obscure to prominent gland-like protuberances (aerophores) at their bases; blades chartaceous to subcoriaceous, pinna margins remotely serrulate to biserrate, especially at pinna tips; veins free, simple or 1-forked, occasionally 2-forked; fertile fronds with contracted pinnae, sporangia at maturity seemingly covering abaxial surfaces of fertile blades, actually in a line along both sides of costa, without true indusia, margins scarious and recurved, protecting the sporangia when young; sporangia sometimes with hair-like paraphyses, sporangial stalks of 4–6 rows of cells; annuli oblique, uninterrupted by stalks; spores tetrahedralglobose, with surface smooth or irregularly tuberculate, sometimes with scattered globules; gametophytes epigeal, chlorophyllous, cordate or somewhat elongate; x⫽66. Plagiogyria is a genus of generally high elevations, and the species often grow in cloud forests or in boggy areas with poor soils. According to most accounts, Plagiogyria comprises about 40–50 species, mostly of southern and eastern Asia (Japan to Malesia, China, and the Himalayas), but most authors agree that this number is inflated (e.g., see Kramer in Kubitzki, 1990). A more realistic estimate is 15–20 species worldwide, and Zhang and Nooteboom (1998) recognized only 11. In the New World there are at most six species (Lellinger, 1971), but only one is widespread. We feel that even six is too high, that a better estimate is three; Zhang and Nooteboom (1998) recognized only a single American species, but their treatment stressed more the Asian taxa. The characters of vein branching and toothing used to separate the American species by Lellinger seem tenuous and insufficient. More work, especially population field studies, is needed to evaluate better the American species of this genus. The genus has often been thought to be related to the Osmundaceae, but recent molecular data support the conclusion that it is more closely related to the tree ferns, Dicksoniaceae and Cyatheaceae (Pryer et al., 1995; Wolf et al., 1999). Plagiogyria is, however, a rather isolated genus, phylogenetically. It is is distinguished by having dimorphic fronds, lack of indusia, flared stipe bases, and sporangia with rather thick (ca. 6-seriate) sporangial stalks and slightly oblique annuli. There is a superficial

Copeland, E. B. 1929. The fern genus Plagiogyria. Philipp. J. Sci. 38: 377–415; Lellinger, D. B. 1971. The American species of Plagiogyria sect. Carinatae. Amer. Fern J. 61: 110–118; Riba, R., B. Pe´rez-Garcia & A. Mendoza. 2002. Pteridofitas. Familia Plagiogyriaceae. Flora de Me´xico 6(6): 1–7; Zhang, X.-C. & H. P. Nooteboom. 1998. A taxonomic revision of Plagiogyriaceae (Pteridophyta). Blumea 43: 401–469.

1. PLAGIOGYRIA PECTINATA Plagiogyria pectinata (Liebm.) Lellinger, Amer. Fern J. 61: 115. 1971. Figs. 227A–E. Lomaria pectinata Liebm., Mexic. Bregn. 233 (reprint. 81). 1849. Type. Mexico. Oaxaca: Cerro de Sempoaltepec, Liebmann s.n. [Pl. Mex. 2477, Fl. Mex. 68] (C!, frag. US!). Lomaridium semicordatum C. Presl, Epimel. Bot. 155. 1849 [1851]. Plagiogyria semicordata (C. Presl) Christ, Farnkr. Erde 176. 1897. Type. Colombia. s. coll. s.n. (PRC). Lomaria arguta Fe´e, Me´m. Foug. 8: 70. 1857. Plagiogyria arguta (Fe´e) Copel., Philipp. J. Sci. 38: 407. 1929. Type. Mexico. Veracruz: Valle´e d’Orizaba, Cerro del Agua, Schaffner 98 (P). Plagiogyria biserrata Mett., Abh. Senckenberg. Naturf. Ges. 2: 272, t. 15, f. 1–18. 1858. Lectotype (chosen by Copeland, 1929: 409). Venezuela. [Aragua:] Colonia Tovar, Moritz 400 (B, photo US; isolectotypes HBG. One of the other syntypes (Linden 556, B, isosyntype UC!) of this species is P. costaricensis Mett. ex Kuhn, and a third syntype is from Mexico (Karwinski s.n., B). Plagiogyria aequidentata E. Fourn., Mexic. Pl. 1: 133. 1872. Type. Mexico. Veracruz: Orizaba, Mu¨ller 723 (P?, Herb. Lenormand, not found). Plagiogyria truncata Mickel & Beitel, Mem. New York Bot. Gard. 46: 283. 1988. Type. Mexico. Oaxaca: Distrito Ixtla´n, trail from San Pedro Nolasco N to the Llano Verde, Mickel 3815 (NY!).

Rhizomes, including old stipe bases, 1.5–4 cm diam.; fronds 30–140 cm tall, the fertile ones more erect than the sterile; stipes of sterile fronds short, 1⁄4–1/5 the frond length, stramineous, dark at base, glabrous; blades deeply pinnatisect or pinnate at base, 6–20 cm wide, truncate to greatly reduced proximally, the shortest pinnae 1⁄2–1⁄4(–1/10) length of longest pinnae, as little as 1 cm long, blade apices acute to abruptly acuminate; pinnae spreading, the largest 3–10 cm ⫻ 4–12 mm, linear, curved to straight, dilate at the base, frequently more surcurrent than decurrent, acute, margins subentire to serrulate to biserrate; veins simple or onceforked at the costae, or one or both veins of a fork sometimes forked again midway toward margin; fertile fronds pinnate, sessile or very short-stalked ⬍ 1 mm, to 140 ⫻ 4–12 cm; stipes 1⁄3–1⁄2 the frond length; pinnae contracted, 3–7 cm ⫻ 2–4 mm; 2n⫽132 (Jam).

PLEOPELTIS

Distribution. Terrestrial in evergreen cloud forests, wet cloud forests of high elevations; 1750–3250 m. Mexico; Guat, CR, Pan; Cuba, Jam, Hisp; Col, Ven, Ec, Peru, Bol. Selected Specimens Examined. Chih (Correll & Gentry 23104, MO, UC). Chis (Breedlove 21995, DS, MEXU, NY). DF (Lyonnet 471200003, MEXU). Dgo (Palmer 124, NY). Gro (Lorea 3700, FCME). Hgo (Moore 3207, UC, US). Jal (Rose & Painter 7598, BM, NY, US). Me´x (Hinton 2808, BM, K, NY). Mich (Rzedowski 42670, IEB). Mor (Lyonnet 1419, BM, K, MEXU, MO, UAMIZ, UC). Oax (Pringle 4999, B, BM, BR, GH, LE, MO, NY, UC, US). Pue (Pringle 15703, CAS, MO, UC, US). Qro (Dı´az Barriga & Carranza 7378, IEB). Son (Van Devender 97-831, ARIZ). Ver (Ventura A. 9014, ENCB, NY).

We adopt a broader concept for this taxon in Mexico than has been applied in most recent floristic and revisionary accounts (an approach that has been taken to an even greater extent by Tryon and Stolze (1989a: 101) and also by Zhang and Nooteboom (1998). We now suspect that much of the variation in neotropical Plagiogyria is contained within a single species, the oldest name for which is the one adopted here. Thus, we would also include within the circumscription of P. pectinata several Antillean species recognized by Lellinger (1971), including P. maxonii Copel. in Jamaica and P. serrulata (Willd.) Lellinger in Hispaniola. Taxa that seem slightly more distinct in the New World include P. costaricensis Mett. ex Kuhn, from Costa Rica, Venezuela, and Ecuador to Bolivia, and P. fialhoi (Fe´e & Glaziou) Copel. in southeastern Brazil. Characters used by Lellinger (1971), Smith (1981), Mickel and Beitel (1988), and Lellinger (in Davidse et al., 1995) to distinguish the Mexican taxa include the degree and pattern of serrations on the margins of the sterile pinnae and the extent of reduction, if any, of the proximal pinnae. The former character we believe is largely a function of the number of vein forks (one or two), and hence a function of pinna width, which varies considerably in Plagiogyria; similarly, reduction of the proximal pinnae is extremely variable, even within populations and on a single plant. For example, specimens collected from around Lagunas de Zempoala, on the border between Edo. Morelos and Edo. Me´xico, show all possible transitional states, from virtually no reduction of the proximal pinnae to extreme reduction. We have observed that there is also great variation in size of plants in this and other populations, as well as size of fronds on a single plant. Possibly this is related to moisture and other environmental fluctuations,

467

from year to year. Thus, we are unable to apply the characters and character states discussed by previous authors to delineate Mexican specimens into two or three species. Furthermore, we find no geographical correlation to the morphological differences that do exist, other than that plants become increasingly smaller as one goes northwestward in Mexico, into habitats that are often drier, colder, and less equable. Evidence suggesting hybridity between species in Mexican specimens is suspect. Re-examination of spores of Mickel 6779 (NY, UC), cited by Mickel and Beitel (1988) as a putative hybrid between P. pectinata and P. truncata, shows that the spores are well formed, not particularly irregular as originally stated. We now regard this particular collection as simply one of the many variants in a variable species. Plagiogyria pectinata is based on a Liebmann collection from Oaxaca. The collection comprises three sterile fronds, which are narrow (8–9 cm) and taper strongly to proximal pinnae that are 1 ⁄3–1⁄4 the length of the longest pinnae.

99. PLEOPELTIS Pleopeltis Humb. & Bonpl. ex Willd., Sp. Pl., ed. 4, 5(1): 211. 1810. Type: Pleopeltis angusta Humb. & Bonpl. ex Willd. ⫻Pleopodium Schelpe & Anthony, Bothalia 15: 557. 1985. Type: ⫻Pleopodium simianum Schelpe & Anthony [⫽ Pleopeltis macrocarpa ⫻ Polypodium polypodioides subsp. ecklonii (Kunze) Schelpe].

Epiphytic, less commonly epipetric; rhizomes long-creeping, branched, rhizome scales concolorous to bicolorous, nonclathrate to rarely clathrate, peltate at bases, surfaces hairy (comose) or glabrous, margins entire to fimbriate or erose; fronds simple to pinnatifid, rarely pinnate-pinnatifid, distant to clumped, monomorphic to subdimorphic, stipitate, articulate; blades firm to coriaceous, sparsely to densely scaly with peltate, round to ovate-lanceolate scales; veins netted, rarely free, areoles with 1–3 free included veins; sori round to oblong, within costal areoles in one row on each side of midribs, at junction of several included veins (compital), exindusiate, with round, peltate scales, at least in immature sori; sporangia glabrous; spores yellowish, bilateral, with perispore shallowly to prominently verrucate; x⫽34, 35, 37? Pleopeltis is a segregate of Polypodium and as treated herein comprises 15–20 species (excluding Lepisorus of Asia). The distribution of the genus thus construed is primarily New World, with a few species in Africa to India and Sri Lanka. Pleopeltis is usually distinguished by the blades and sori with peltate, round scales, the areoles with free, included veins, and sori at the junction of several included veins. The generic limits of Pleopeltis have long been in need of careful scrutiny (Windham, 1993). Morphologically, a few species resemble species of Microgramma, but other species are more similar to scaly species of Polypodium. On the basis of molecular evidence, it is now believed that all scaly-bladed species treated herein as Polypodium are more appropriately placed in Pleopeltis, and that the circumscription of Polypodium should be restricted to those species that have essen-

468

PLEOPELTIS

tially scaleless blades (Smith et al., ms. in prep.; Schneider et al., unpubl. data). The implications of that for the Mexican fern flora are that the following taxa will ultimately need transfer to Pleopeltis: Polypodium alansmithii R. C. Moran; P. collinsii Maxon; P. cryptocarpon Fe´e; P. fallacissimum Maxon; P. furfuraceum Schltdl. & Cham.; P. guttatum Maxon; P. lepidotrichum (Fe´e) Maxon; P. lindenianum Kunze; P. madrense J. Sm.; P. platylepis Mett. ex Kuhn; P. plebeium Schltdl. & Cham.; P. polypodioides (L.) Watt var. knoblochianum Mickel; P. pyrrholepis (Fe´e) Maxon; P. remotum Desv.; P. rosei Maxon; P. rzedowskianum Mickel; P. sanctae-rosae (Maxon) C. Chr.; and P. villagranii Copel. Combinations in Pleopeltis already exist for Polypodium muenchii, P. polypodioides (L.) Watt var. aciculare and var. michauxianum, P. riograndense, and P. thyssanolepis, but in this work we treat them in the genus Polypodium, pending future publications. Polypodium fallax is treated here in Pleopeltis on the basis of the caducous, peltate, circular paraphyses on the blades. Polypodium percussum Cav., which is often placed in Pleopeltis (e.g, by Tryon & Tryon, 1982; Tryon and Stolze, 1993), is here treated in Microgramma (which see for further discussion). Several species of Pleopeltis appear to hybridize with species of Polypodium (as treated in this flora) to produce abortivespored or fertile species in the hybrid genus ⫻Pleopodium, which we treat here as a synonym of Pleopeltis, pending further study of generic limits of Pleopeltis and Polypodium. One such putative hybrid, ⫻Pleopodium fallacissimum (Maxon) Mickel & Beitel was thought to be the putative hybrid between Pleopeltis polylepis

var. erythrolepis and Polypodium guttatum. However, we now consider this taxon to be a good species, and treat it under Polypodium, even though its proper position we anticipate will eventually be in Pleopeltis. The distribution and density of the peltate scales on the blades are important characters for distinguishing species in Pleopeltis. In the key and species descriptions, the terms used to describe the scales are defined as follows: sparse ⫽ scales rarely touching and often space between scales for two or more similar scales; scattered ⫽ some scales touching and often space between scales for a single similar scale; dense ⫽ most scales overlapping other scales and with little or no space for any additional scales. References Aguirre-Clavero´n, R. & M. L. Arreguı´n. 1988. Claves de familias, ge´neros, especies y variedades de pterido´fitas del estado de Nuevo Leo´n, Me´xico. Anales Esc. Nac. Ci. Biol. 32: 9–61; Anthony, N. C. & E. A. Schelpe. 1985. ⫻Pleopodium–a putative intergeneric fern hybrid from Africa. Bothalia 15: 555–559; Hinton, J. & G. S. Hinton. 1995. Checklist of Hinton’s collections of the flora of south-central Nuevo Leo´n and adjacent Coahuila. Acta Bot. Mex. 30: 41–112; Hooper, E. A. 1994. Biosystematic analysis of the Pleopeltis macrocarpa complex in the Neotropics. Ph.D. Thesis, Univ. Kansas, Lawrence; Hooper, E. A. 1995. New combinations in the Pleopeltis macrocarpa group (Polypodiaceae: Polypodieae). Amer. Fern J. 85: 75–82; Mickel, J. T. & J. M. Beitel. 1987. Notes on ⫻Pleopodium and Pleopeltis in tropical America. Amer. Fern J. 77: 16–27; Rodrı´guez-Lo´pez, R. & M. L. Arreguı´n-Sa´nchez. 1992. Adiciones a la pteridoflora del estado de Nuevo Leo´n, Me´xico. II. Anales Esc. Nac. Ci. Biol. 37: 27– 34; Weatherby, C. A. 1922. The group of Polypodium lanceolatum in North America. Contr. Gray Herb. 65: 3–14; Wendt, T. 1980. Notes on some Pleopeltis and Polypodium species of the Chihuahuan Desert region. Amer. Fern J. 70: 5–11.

Key to the Mexican Species of Pleopeltis (hybrids not keyed) 1. Blades pinnatifid or pinnate-pinnatifid. 2. Blades pinnate-pinnatifid, 2–4 cm long; rhizomes thread-like, 0.5–0.8 mm diam. ................................................................................ 5. P. fallax. 2. Blades pinnatifid or pinnatisect, 5–20 cm long; rhizomes 1.5 mm or more in diam. 3. Rhizome scales bicolorous with brown-black central portion, 1.5–2 ⫻ 0.5–0.8 mm, comose; pinnae 1.5–7 mm wide; adaxial surfaces of blades with scattered fimbriate scales. ................................................................................................................................ 1. P. angusta. 3. Rhizome scales concolorous, lustrous brown, 4–6 ⫻ 1.5–2 mm, glabrous; pinnae 7–14 mm wide; adaxial surfaces of blades glabrous. ........................................................................................................................................................................................ Polypodium muenchii. 1. Blades simple. 4. Stipes absent to short (less than 1/10 of frond length), flattened; sori elongate; rhizome scales 0.3–0.5 mm long, comose with brown hairs 1–1.5 mm long. ........................................................................................................................................................................ 2. P. astrolepis. 4. Stipes present, short to long (1/10–1⁄2 of frond length), terete; sori round to slightly oblong; rhizome scales 0.8–3 mm long, glabrous or comose with long hairs. 5. Laminar scales lanceolate, 1 ⫻ 0.3–0.5 mm, scattered to dense, with fimbriate margins; rachises with scattered, lanceolate, black-centered scales 1–2 mm long, especially near base; soral scales largely lanceolate, 1 mm long, a few orbicular (0.5 mm long), all with fimbriate margins. ......................................................................................................................................................... 3. P. conzattii. 5. Laminar scales mostly orbicular, 0.3–1 mm wide, with some sparse to dense lanceolate scales, with fimbriate to subentire margins; rachises lacking black-centered lanceolate scales; soral scales largely orbicular, 0.3–1 mm wide, fimbriate to subentire. 6. Soral scales with brown centers and pale brown margins, 0.3–0.5 mm wide; laminar scales sparse, brown-centered, 0.3– 0.5 mm wide; fronds lanceolate, dimorphic, fertile 8–20(–25) mm wide, sterile 22–32(–40) mm wide; rhizome scales denticulate, dimorphic, on rhizomes 0.8–1 mm long, at base of stipes 2–3 mm long, densely comose. ........................... 6. P. mexicana. 6. Soral scales with brown-black centers and pale brown to whitish border, 0.3–1 mm wide; laminar scales sparse to dense, brown to dark reddish brown-centered, 0.3–1 mm wide; fronds narrowly elliptic to linear-elliptic, monomorphic, 6–22 mm wide; rhizome scales denticulate to erose, dimorphic to monomorphic, rarely to sparsely comose. 7. Laminar scales sparse, never overlapping or even contiguous, 0.5 mm wide; at least some of the veins abaxially, especially those nearest the midribs and base of blades, darkened and thickened; rhizome scales dimorphic,

PLEOPELTIS ANGUSTA

469

denticulate, along rhizomes strongly bicolorous, 1–1.5 mm long, at base of stipes and often at rhizome apices essentially concolorous, light brown, 1.5–2 mm long; 450–1900. ................................................................................. 4. P. crassinervata. 7. Laminar scales scattered to dense, sometimes overlapping, 0.1–0.3 mm wide; veins abaxially (except midribs) not visible; rhizome scales monomorphic, bicolorous with black centers, erose, 1.5–3 mm long; (2050–)2450– 3100 m. .........................................................................................................................................................................................7. P. polylepis.

1. PLEOPELTIS ANGUSTA Pleopeltis angusta Humb. & Bonpl. ex Willd., Sp. Pl., ed. 4, 5(1): 211. 1810. Polypodium angustum (Humb. & Bonpl. ex Willd.) Liebm., Mexic. Bregn. 186 (reprint 34). 1849. Type. Mexico. Near Ario, Humboldt & Bonpland s.n. (BWilld. 19729!; isotype P-Bonpl.!, photo US!).

Rhizomes long-creeping, 1.5–2 mm diam.; rhizome scales black with very narrow pale margins, 1.5–2 ⫻ 0.5–0.8 mm, comose, margins fimbriate; fronds distant, slightly dimorphic, fertile fronds narrower; stipes castaneous, 1⁄3–3⁄5 the frond length, glabrous, winged almost to base; blades deeply pinnatisect, 4–15 (–20) cm wide, with 1–5 pairs of lobes, these opposite to subopposite, linear, ascending, 4–12 cm ⫻ 1.5–7 mm; midribs (of rachises and costae) abaxially castaneous to black, prominent usually to blade and pinna apices; veins not readily visible; abaxial blade scales round, 0.3 mm wide, to ovate-lanceolate, 0.8–1 mm long, centers brown, margins denticulate, darker and more dissected around the sori; sori oblong, usually broader than the laminar space, with peltate soral scales 0.5–0.8 mm wide, centers brown-black, margins pale, fimbriate; 2n⫽68 (Guat), 74 (Oax, Guat), the latter number probably erroneous. Brazilian material referred to P. angusta belongs to the related P. pleopeltifolia (Raddi) Alston. Pleopeltis angusta, the type of the genus, differs from most Mexican species by the pinnatifid blades. In this character, it more closely resembles scaly species here treated in Polypodium. From P. fallax, which has pinnate-pinnatifid blades, P. angusta differs by the much larger fronds, thicker rhizomes, and blackened midribs. Key to the Mexican Varieties of Pleopeltis angusta 1. Pinna pairs 1–2; pinnae 4–7 mm wide; Pacific slope. ........................................................................................... 1a. var. angusta.

1. Pinna pairs 3–4(–5); pinnae 1.5–5 mm wide; Atlantic slope. .............................................................................. 1b. var. stenoloma.

1a. Pleopeltis angusta. var. angusta

Fig. 229N.

Differing from var. stenoloma by having 1–2 lateral pinna pairs per blade, pinnae 4–7 mm wide.

Distribution. Epiphytic on Pacific slope; 400–2800 m. Mexico; Guat, Salv, Nic. Lorea-Herna´ndez (in Davidse et al. 1995: 347) also cited this species from Honduras. Selected Specimens Examined. Chis (Matuda 5423, LL, US). Col (Bautista 16882, GUADA, UC). Dgo (Sa´nchez S. 724, US). Gro (Hinton 11067, LL, US). Jal (Breedlove 45740, CAS, ENCB). Me´x (Tejero-Dı´ez 2547, IZTA). Mich (Rzedowski 39682, ENCB). Nay (Tenorio L. 16802, IEB, MEXU). Oax (Mickel 6172, ENCB, UC; Rzedowski 21318, DS, ENCB, TEX).

1b. Pleopeltis angusta var. stenoloma (Fe´e) Farw., Amer. Midl. Naturalist 12: 297. 1931. Figs. 229O–R. Drynaria stenoloma Fe´e, Me´m. Foug. 5: 272. 1852. Type. Mexico. [Oaxaca: Distr. Ixtla´n:] Talea, Galeotti 6532 (P!). Drynaria tortulosa Fe´e, Me´m. Foug. 5: 271. 1852. Type. “Cuba.” Linden s.n. (P?). Smith (1981) considered the type locality in error; probably it is Mexico, Chiapas. Pteropsis angustifolia E. Fourn., Mexic. Pl. 1: 86. 1872. Syntypes. Mexico. Leibold s.n.; “in regione Orizabensi,” Bourgeau 2777 (P!), Mu¨ller 46, Bot., t. 53; “Orizaba, Huatusco, Cordoba,” Schaffner 226; “in regione Oajacense,” Salle s.n. (P!); [Oaxaca:] Talea, Galeotti 6532 (P!), 6308 (P!); Ario, Bonpland s.n. (P!).

Differing from var. angusta by pinna pairs 3–4(–5) per blade, pinnae 1.5–5 mm wide.

470

PLEOPELTIS

0.5–1 mm wide, centers pale to dark brown, margins pale brown, fimbriate, scales on midribs scattered, 0.8–1 mm long, lanceolate, cells black, sclerotic; sori oblong, 2–5 mm long, soral scales peltate, 0.1–0.3 mm diam., centers dark brown-black, margins pale brown, fimbriate; 2n⫽70 (CR), 136 (140?) (Mex, PR, Hisp; Hooper, 1994); 148 (PR), probably erroneous.

Distribution. Epiphytic on Atlantic slope; 300–1700(–2700) m. Mexico; Guat. Selected Specimens Examined. Chis (Breedlove 56561, CAS, ENCB; Hammel et al. 15681, MO, UC; Martı´nez S. 18687, TEX). Oax (Mickel 4724, 6377, 7265, ENCB, NY, UC; Wendt et al. 5029, CAS). Ver (Calzada 10305, UC; Purpus 8690, UC, US; Ventura A. 16640, CAS, IEB).

Intermediate specimens between the two varieties with narrow pinnae but few pairs (2–3) occur in Oaxaca: Tehuantepec (Hallberg 1694, 1714, 1739, all NY); along the Tuxtepec-Teotitla´n border (Hallberg 1348, NY).

2. PLEOPELTIS ASTROLEPIS Pleopeltis astrolepis (Liebm.) E. Fourn., Mexic. Pl. 1: 87. 1872. Figs. 228B–F. Polypodium astrolepis Liebm., Mexic. Bregn. 185 (reprint 33). 1849. Phlebodium astrolepis (Liebm.) Conzatti, Fl. Tax. Mex. 1: 100. 1939. Syntypes. Mexico. Oaxaca: Trapiche de la Concepcio´n, Liebmann s.n.; Veracruz: Mirador, Aug 1841 [erroneously 1842 in Hooper, 1994], Liebmann s.n. [Fl. Me´x. 87]. Both specimens are mounted on one sheet (C!). Smith (1981: 176) chose the lowermost specimen as lectotype. Grammitis lanceolata Schkuhr, Deutschl. Krypt. Gew. 1: 9, t. 7. 1804, non Polypodium lanceolatum L., 1753, nec Pleopeltis lanceolata (L.) Kaulf., 1824. Grammitis elongata Sw., Syn. Fil. 22, 213. 1806, non Pleopeltis elongata Kaulf., 1824. Polypodium elongatum (Sw.) Mett., Abh. Senckenberg. Naturf. Ges. 2: 88, t. 2, f. 8, 9. 1857, hom. illeg., non Schrader, 1818, nec Aiton, 1879. Gymnogramma elongata (Sw.) Hook., Sp. Fil. 5: 157. 1864. Polypodium lanceolatum L. var. elongatum (Sw.) Krug, Bot. Jahrb. Syst. 24: 131. 1897. Type. Jamaica. Swartz s.n. (S). Grammitis revoluta Spreng. ex Willd., Sp. Pl., ed. 4, 5(1): 139. 1810. Pleopeltis revoluta (Spreng. ex Willd.) A. R. Sm., Proc. Calif. Acad. Sci., ser. 4, 40: 230. 1975, non Pleopeltis revoluta Alderw., 1909. Type. Not stated, received from Sprengel (B-Herb. Willd. 19584, microfiche UC!).

Rhizomes long-creeping, 0.8–1.5 mm diam.; rhizome scales monomorphic, 0.3–0.5 mm long, ovate to ovate-lanceolate, comose with brown hairs 1–1.5 mm long, black-centered, margins narrow, pale, lacerate-denticulate; fronds (4–)8–11(–15) cm long; stipes usually less than 1/10 the frond length, 0–18 mm long, black, strongly flattened; blades simple, linear-lanceolate to narrow-elliptic to oblanceolate, margins thin; midribs abaxially black in proximal 1⁄2–3⁄4 of blade length; veins not readily visible; abaxial blade scales scattered to rare, round to ovate-lanceolate,

Distribution. Epiphytic, or epipetric at lower elevations; 0– 1850 m. USA (Fla); Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Sur, Fr Gui, Peru, Braz, Bol. Also cited from the Guianas and Ecuador by LoreaHerna´ndez (in Davidse et al., 1995). Selected Specimens Examined. Camp (Lundell 1161, US). Chis (Conrad & Conrad 2994, MO, UC). Col (Sanders 11715, UC). Gro (Lorea 4608, FCME). Jal (Mexia 1495, CAS, DS, IEB, UC). Nay (Breedlove 44322, CAS). Oax (Martı´nez C. 375, DS, UC). Tab (Rovirosa 618, US). Ver (Copeland herb. 148, UC). Unverified, Doubtful, or Mistaken Reports. Hgo (reported by Mickel & Beitel, 1988, but not verified). Mich (reported by Mickel & Beitel, 1988, but not verified). Qro (Gonza´lez P. 224, IEB, QMEX, cited by Dı´az-Barriga & Palacios-Rios, 1992, and Arreguı´n et al., 2001, but not verified). SLP (reported by Mickel & Beitel, 1988, but not verified). Yuc (Lundell 1161, F, cited by Lorea-Herna´ndez in Davidse et al., 1995, but this collection is from Campeche).

Among its simple-bladed congeners, P. astrolepis is distinct in its elongate sori, very short stipes, and small, comose rhizome scales. Specimens from the Pacific slope tend to be slightly longer (mostly 11–15 cm) than those on the Atlantic slope (where fronds are 8–10 cm), but the differences are not taxonomically significant. Pleopeltis ⫻sordidula (Maxon & Weath.) Mickel & Beitel (Amer. Fern J. 77: 21. 1987), based on Polypodium sordidulum Maxon & Weath. in Weatherby (Amer. Fern J. 17: 92. 1927. Type. Mexico. Veracruz: near Cordoba, Spence 114, GH!; isotype US!) is the putative hybrid between P. astrolepis and P. fallax, and is intermediate in several morphological characters between the putative parents (Fig. 228N; Mickel & Beitel, 1987). The filiform, rather sparsely scaly rhizomes bearing small, densely comose scales point to P. fallax as one parent. This hybrid is known only from Veracruz, as an epiphyte on coffee trees in montane forests, 900–1400 m: Ver (Copeland herb. 149, MEXU, MICH, NY, UC, US; Fisher s.n., DS; Fisher 37, US).

PLEOPELTIS CRASSINERVATA

Spence 111 (US), from the same locality as the type, is probably another Pleopeltis hybrid, of uncertain parentage, but between a simple-bladed and a deeply pinnatifid species. The blades are lobed, with about 12 pairs of segments, the midribs darkened, and the laminar scales strongly fimbriate.

3. PLEOPELTIS CONZATTII Pleopeltis conzattii (Weath.) R. M. Tryon & A. F. Tryon, Rhodora 84: 129. 1982. Figs. 228G, H, J, K. Polypodium conzattii Weath., Contr. Gray Herb. 65: 11. 1922. Type. Mexico. Oaxaca: Cerro San Felipe, Conzatti 678 (GH!).

Rhizomes creeping, 1.5–2 mm diam.; rhizome scales dimorphic, along rhizomes 0.5–1 mm long, linear-lanceolate, centers black with occluded cells, margins pale, narrow, fimbriate-serrulate, comose, at stipe bases 2–2.5 mm long, slender with long fimbriate hair-like teeth, centers dark; fronds monomorphic, distant; stipes 1⁄8–1/6 the frond length, atropurpureous, with blackcentered, linear-lanceolate scales with sharply serrulate margins; blades simple, linear-lanceolate to lanceolate, (12–)14–30(–35) ⫻ 0.7–2.5 cm; midribs abaxially black at blade bases or nearly throughout blade length, also with scattered, bicolorous, blackcentered scales 1–2 mm long, these especially dense proximally; veins not readily visible; abaxial blade scales dense, hyaline with centers red-orange, fimbriate-ciliate, mostly ovate-lanceolate, 1 ⫻ 0.3–0.5 mm; sori slightly oblong, soral scales mostly ovatelanceolate, similar to laminar scales, 0.5–1 mm long, centers brown to pale brown, margins fimbriate; 2n⫽68 (Oax; Hooper, 1994).

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4. PLEOPELTIS CRASSINERVATA Pleopeltis crassinervata (Fe´e) T. Moore, Index Fil. 345. 1862. Fig. 228A. Drynaria crassinervata Fe´e, Me´m. Foug. 8: 97. 1857. Polypodium crassinervatum (Fe´e) Kiaersk., Beretn. Bot. Haves Virks. 1871–73: 49. 1874. Polypodium lanceolatum L. var. crassinervatum (Fe´e) Weath., Contr. Gray Herb. 65: 8. 1922. Syntypes. Mexico. Veracruz: “Valle´e d’Orizaba,” Schaffner 173, 489; Co´rdoba, Schaffner 174a (not found in P; Schaffner 489, 174a in RB).

Rhizomes creeping, 1–1.5 mm diam.; rhizome scales dimorphic, 1–1.5 mm long along rhizomes, ovate-lanceolate, black, margins generally broad, pale, denticulate, rarely comose, at stipe bases 1.5–2 mm long, lanceolate, generally concolorous with only dark center spot, denticulate; fronds monomorphic; stipes 1/10–1⁄3 the frond length, terete; blades simple, linear-elliptic, (8–)10–30 ⫻ 0.7–1(–2) cm, thin; midribs abaxially blackish only at blade bases or nearly throughout the blade length; veins faintly to readily evident at blade bases near the midribs, dark; abaxial blade scales sparse, round, 0.5 mm diam., brown-centered, margins pale brown to whitish, fimbriate; sori oblong, soral scales peltate, 0.5– 0.8 mm wide, centers dark brown-black, margins pale brown, fimbriate; 2n⫽70 (Oax, SLP).

Distribution. Epiphytic in wet montane forests; (450–)700– 1900 m. Mexico; Guat, Hond, Nic, CR. Selected Specimens Examined. Chis (Breedlove 58423, CAS; Dressler 1596, MEXU, US). Hgo (Croat & Hannon 65892, UC). Oax (Mickel 7143, ENCB, UC). Pue (Rzedowski 29968, ENCB). Qro (Rzedowski 42787, IEB). SLP (King 4271, LL, TEX, UC). Tam (Sharp et al. 52088, US). Ver (Valdivia Q. 2143, UC; Ventura 9505, ASU, ENCB, IEB, XAL).

Distribution. Epiphytic in pine-oak woods; 1800–3000 m. Mexico. Selected Specimens Examined. Gro (Lorea 1267, IEB, 1517, FCME, 3145, 3296, IEB). Oax (Conrad & Conrad 3131, CAS, MEXU; Frame 331, UC; Lorence 3240, MEXU; Rzedowski 21029, DS, TEX).

Pleopeltis conzattii is distinguished by its lanceolate, fimbriatemargined blade scales and black-centered scales along the midribs. Its closest relative is P. mexicana, and a few specimens from Guerrero seem to show intermediacy.

Citation of this species from the states of Chihuahua, Durango, Sinaloa, Me´xico, and Morelos by Mickel and Beitel (1988) is based on misidentifications. Pleopeltis crassinervata is aptly named for its distinctive, conspicuous, dark veins near the rachises (midribs) in the proximal half of the blades. Specimens from Tamaulipas and some from San Luis Potosı´ lack the characteristic darkened veins at the base of the blades, and so could be mistaken for P. mexicana, but the rhizome scales are smaller and less comose, and the fertile and sterile blades are similar in size and shape (not dimorphic). The

472

PLEOPELTIS

blade margins of P. crassinervata tend to be more revolute than in either P. mexicana or P. polylepis. Pleopeltis ⫻melanoneuron Mickel & Beitel (Fig. 228L; Amer. Fern J. 77: 25. 1987) was described on the basis of a specimen from Mexico (Veracruz: Zacuapan, Purpus 5798, US, not found; isotype UC!). The type was earlier cited as a paratype of Polypodium sordidulum by Maxon and Weatherby, but we agree with Mickel and Beitel (1987) that this is not the same hybrid as Pleopeltis ⫻sordidula. The parentage suggested by Mickel and Beitel (P. crassinervata ⫻ fallax) seems not quite so obvious. The blackened lateral veins at the base of the blades are much darker than one finds in either of their suspected parents and suggests that P. angusta might possibly be involved in the parentage. Two additional collections of this hybrid have been made, the first also from Veracruz (Foster 14, US), and the second, lacking sori, from Hidalgo (30 km NNE of Tulancingo on Mex. 130, ca. 2000 m, Marcks & Marcks 718, WIS).

5. PLEOPELTIS FALLAX Pleopeltis fallax (Schltdl. & Cham.) Mickel & Beitel, Amer. Fern J. 77: 21. 1987. Figs. 228O–Q. Polypodium fallax Schltdl. & Cham., Linnaea 5: 609. 1830. Type. Mexico. [Veracruz:] Misantla, Schiede & Deppe 758 (isotype B!–photo BM!, LE!). Polypodium tuerckheimii Christ, Bull. Herb. Boissier, se´r. 2, 5: 254. 1905. Type. Guatemala. Alta Verapaz: Cubilquitz, Tu¨rckheim s.n. (P, photo BM).

Rhizomes long-creeping, 0.6–0.8 mm diam., green; rhizome scales monomorphic, black, clathrate, 0.5 mm long, ovate, thickwalled, with cluster of long brown hairs (ca. 1 mm long) at central point of attachment, rhizomes thus appearing densely pubescent; fronds monomorphic, distant; stipes ca. 1⁄3 the frond length, light brown with very narrow decurrent wing, with very sparse, minute, 0.3–0.5 mm, dark, appressed scales; blades oblong to narrowly deltate, 2–4 ⫻ 1–2 cm, pinnate-pinnatifid, pinnae 4–7 pairs, opposite to alternate, adaxially glabrous; midribs abaxially greenish to yellowish or light brown primarily at blade bases, costae not visible; veins not readily visible; abaxial blade scales sparse, reddish brown, lanceolate, serrulate, 0.5–0.8 mm long; sori round, at maturity exceeding the pinnae in width, peltate soral scales ovate-lanceolate, 0.1–0.3 mm long, centers dark brown-black, margins pale, fimbriate.

Distribution. Epiphytic and on rock ledges in mesic woods; 50–1500 m. Mexico; Guat, Bel, Hond, Nic, CR. Selected Specimens Examined. Chis (Palacios-Rios 2804, UC). Hgo (Gimate L. 838, ARIZ, LL). Oax (Mexia 9189, ARIZ, LL, UC, US). Pue (Rzedowski 29975, 32437, ENCB; Sharp 441201, US). Qro (Dı´az B. & Carranza 6521, IEB). SLP (Alcorn 1802, TEX). Ver (Faden et al. 76/101, UC; Ventura A. 5071, LL).

This species appears superficially like a Polypodium in its pinnate to pinnate-pinnatifid fronds, but we place it in Pleopeltis on the basis of its caducous peltate paraphyses. The species is distinct in its divided blades and thread-like rhizomes. Blatchley s.n. (US), from Mt. Orizaba (Veracruz), is a mixed collection of P. fallax and a possible hybrid involving that species. The putative hybrid has lobed blades, somewhat similar to those of P. ⫻sordidula (see under P. astrolepis), but the abaxial surfaces have more numerous, darker scales.

6. PLEOPELTIS MEXICANA Pleopeltis mexicana (Fe´e) Mickel & Beitel, Amer. Fern J. 77: 21. 1987. Figs. 228R–V. Drynaria mexicana Fe´e, Me´m. Foug. 8: 97. 1857. Syntypes. Mexico. Veracruz: Galeotti 6321 (P; not 6321, BR, which is the type of Polypodium galeottii M. Martens ⫽ Megalastrum subincisum); Puebla, Schaffner 179 (RB); Popocate´petl, Schaffner 292 (RB). Polypodium lanceolatum L. var. trichophorum Weath., Contr. Gray Herb. 65: 8. 1922. Pleopeltis macrocarpa (Bory ex Willd.) Kaulf. var. trichophora (Weath.) Pic. Serm., Webbia 23: 189. 1968. Type. Mexico. Me´xico: Lava fields near Eslaba, Pringle 11797 (GH!; isotypes CAS!, US!).

Rhizomes creeping, 1.5–2 mm diam.; rhizome scales dimorphic, 0.8–1 mm long along rhizomes, ovate-lanceolate, densely comose, centers black, margins narrow, pale, denticulate, 1.5–2.5 mm long at stipe bases, narrowly lanceolate, centers black to rarely concolorous, margins narrow, pale, denticulate; fronds dimorphic, sterile ones 2–3 times the width and 1⁄2–3⁄4 the length of the fertile, distant; stipes 1⁄4–1⁄3 the frond length, terete, 1–8 cm long; blades coriaceous, dimorphic, simple, the sterile ones lanceolate, 10–20(–25) ⫻ 2.2–3.2(–4) cm, the fertile ones lanceolate to linear-lanceolate, (12–)15–25(–30) ⫻ 0.8–2(–2.5) cm; midribs abaxially black nearly to blade apices; veins not readily visible; abaxial blade scales sparse, round, 0.3–0.5 mm diam.,

PLEOPELTIS POLYLEPIS

brown-centered with margins pale brown to white, fimbriate; sori oblong, soral scales peltate, 0.3–0.5 mm diam., similar to laminar scales; 2n⫽68 (Oax; Andrews, 1994).

Distribution. Epiphytic to epipetric in wet montane forests; 1300–2900 m. Mexico; Guat, Hond, Salv, Nic (Martı´nez 1806, MEXU). Selected Specimens Examined. Chis (Croat 40678, UC). Col (Goldsmith 42, UC). DF (Lyonnet 1913, MEXU, US). Dgo (Sa´nchez 727, MEXU). Gro (Lorea 855, ENCB). Hgo (Moore 1972, UC). Jal (Breedlove 39285, CAS). Me´x (Rzedowski 22828, DS, ENCB). Mich (Breedlove 64340, CAS). Mor (Hitchcock & Stanford 7101, DS, UC). Nay (Breedlove 45516, CAS). NL (Reeves 6035, 6116, ASU). Oax (Wendt et al. 4987, CAS, CHAPA, MEXU). Pue (Orcutt 3993, DS). Qro (Breedlove 66652, CAS). SLP (Lynch 9031, TEX, det. uncertain). Sin (Gentry 5882, 6334, ARIZ, DS). Ver (Valdivia Q. 2216, UC, XAL).

Pleopeltis mexicana is closely allied to P. macrocarpa (which is unknown from Mexico but occurs in Central America, South America, the Antilles, and Africa) and has usually been treated as a variety of it. The former is distinct in its brown-centered soral scales and lanceolate, somewhat dimorphic blades. Some specimens from Guerrero, e.g., Rzedowski & McVaugh 108 (ENCB), Martı´nez S. 886 (IEB), and Lorea 1635 (IEB), have many of the abaxial laminar scales with a short tail, thus tending towards P. conzattii, which is also closely related. Pleopeltis mexicana is best distinguished from P. polylepis by the conspicuously comose rhizome scales, dimorphic blades, and dimorphic rhizome scales, those at the base of the blades being tan, more lanceolate, and not comose. Pleopeltis mexicana appears to hybridize with Polypodium thyssanolepis, forming ⫻Pleopodium tricholepis Mickel & Beitel (Figs. 235A–C; Amer. Fern J. 76: 17. 1987. Type. Mexico. Oaxaca: Distrito Etla–Cuicatla´n, 39 km N of Rte. 190 past Telixtlahuaca, 1900 m, Mickel 3873, NY!; isotype UC!). This hybrid between a simple-bladed species and a pinnatifid species is intermediate morphologically between the suspected parents, most obviously in blade shape: irregularly pinnatisect, 16–18 ⫻ 5–6 cm, broadest

473

proximally; lobes 4–6 pairs, 5–7 mm wide. Blade scales are also intermediate between the putative parents: adaxial surfaces with scattered to dense, deeply fimbriate scales, mostly lanceolate (1–1.5 mm long), some round (0.8–1 mm wide), with brown centers grading to whitish margins, midribs dark with lanceolate scales, 1.5–2 mm long, adaxial surfaces with sparse, deeply lacerate, lanceolate scales. The spores are malformed. The hybrid is known only from the type gathering and was collected with Polypodium thyssanolepis (see UC isotype). Although P. conzattii was the only simple-bladed species of Pleopeltis found at the same locality, the presence of round scales and the absence of blackcentered rachis scales makes that species unlikely as a possible parent. The laminar scales are deeply bicolorous as one would expect in hybrids involving P. mexicana. Pleopeltis astrolepis and P. crassinervata are generally species of lower elevations, while P. polylepis is usually found at higher elevations. The existence of this hybrid shows the very close relationship between Pleopeltis (mexicana) and Polypodium (thyssanolepis) and the inappropriateness of separating the two genera on the basis of blade dissection. A very similar hybrid in the Greater Antilles and South America (Jamaica, Hispaniola, Venezuela, Ecuador, Peru) is Pleopeltis ⫻leucospora (Klotzsch) Moore, which is Pleopeltis macrocarpa (Bory ex Willd.) Kaulf. ⫻ Polypodium thyssanolepis.

7. PLEOPELTIS POLYLEPIS Pleopeltis polylepis (Roemer ex Kunze) T. Moore, Index Fil. 348. 1862. Polypodium polylepis Roemer ex Kunze, Linnaea 13: 131. 1839. Syntypes. Mexico. Mineral del Monte, Ehrenberg s.n.; s. loc., Hegewisch s.n. (Herb. Roemer); s. loc., Karwinski s.n. (probably all LZ, destroyed).

Rhizomes creeping, 1–2 mm diam.; rhizome scales monomorphic, 1.5–3 mm long, linear-lanceolate or lanceolate, rarely comose, centers black, occluded or not, margins pale, minutely denticulate or erose, sometimes sparingly comose; fronds monomorphic, distant to approximate; stipes 1/6–1⁄3 the frond length, darkly castaneous, terete; blades simple, narrowly elliptic to narrowly oblanceolate, 6–30 ⫻ 0.6–2.2 cm, apices acute to longacuminate; midribs abaxially hidden (greenish) to brown or blackish only at blade bases or nearly to the blade apices; veins not readily visible; abaxial blade scales scattered to often dense, round, 0.3–1 mm diam., centers dark reddish brown, margins pale brown to whitish, erose to fimbriate; sori oblong, soral scales peltate, 0.3–1 mm diam., centers dark reddish brown to brownblack, margins pale brown to whitish, erose to fimbriate. The overlapping, round laminar scales are diagnostic for this species. The infraspecific taxonomy of this species follows Hooper (1994, 1995).

Key to the Mexican Varieties of Pleopeltis polylepis 1. Leaf blades narrowly elliptic, 1–1.5 cm wide, with conspicuously blackened abaxial midribs; laminar scales ca. 0.4 mm diam., rarely overlapping; soral scales ca. 0.6 mm diam., with conspicuously blackened centers having occluded cell lumina, often persistent in mature sori. ............................................................................................................................................................................. 7c. var. interjecta.

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PLEOPELTIS

1. Leaf blades linear-elliptic to narrowly oblanceolate, usually less than 1 cm wide, midribs abaxially blackish to greenish; laminar scales ca. 0.6 mm diam., usually overlapping; soral scales ca. 0.7 mm diam., with dark brown centers (similar to laminar scales), rarely persistent in mature sori. 2. Stipes ca. 1/5 the blade length; leaf blades 0.7–0.8(–1.5) cm wide, 8–10 times longer than wide; midribs abaxially green or blackened; plants nearly always epiphytic. ............................................................................................................................................ 7a. var. polylepis. 2. Stipes ca. 1⁄2 the blade length; leaf blades 0.8–1 cm wide, 5–7 times longer than wide; midribs abaxially green; plants nearly always epipetric. ................................................................................................................................................................................... 7b. var. erythrolepis.

7a. Pleopeltis polylepis var. polylepis

Figs. 229J–M.

Polypodium peltatum Cav., Descr. Pl. 244. 1801, non Alderw. (1908), nec Pleopeltis peltata Scort. ex Alderw. 1909. Lectotype (chosen by Hooper, 1995: 77). Mexico. Ne´e s.n., right-center specimen of sheet 476120 in the Cavanilles herbarium (MA, photo UC!). Drynaria vestita Fe´e, Me´m. Foug. 5: 271. 1852. Type. Mexico. Galeotti s.n. (P).

Distinguished from other varieties by the stipes ca. 1/5 the blade length; leaf blades 0.7–0.8(–1.5) cm wide, 8–10 times longer than wide; midribs abaxially green or blackened; plants nearly always epiphytic; 2n⫽68 (NL).

e.g., Rzedowski 32441 (IEB, TEX, XAL), and we wonder whether this could be the result of some hybrid involvement with a fully and regularly pinnatifid scaly Polypodium species. In these particular specimens the sori and spores are too young to be informative. Polypodium bartlettii Weath., Amer. Fern J. 25: 56. 1935, Fig. 228M [Type. Mexico. Tamaulipas: Vicinity of San Jose´, above Mesa de Tierra, Bartlett 10286 (US!, photo US!; frag. GH!, US!)], is the putative hybrid between Pleopeltis polylepis and perhaps Polypodium polypodioides; it is known only from the type locality. A synonym is ⫻Pleopodium bartlettii (Weath.) Mickel & Beitel, Amer. Fern J. 77: 20. 1987. Field study is needed to verify its parentage. 7b. Pleopeltis polylepis var. erythrolepis (Weath.) Wendt, Amer. Fern J. 70: 9. 1980. Figs. 229E–H. Polypodium erythrolepis Weath., Contr. Gray Herb. 65: 11. 1922. Pleopeltis erythrolepis (Weath.) Pic. Serm., Webbia 23: 189. 1968. Type. Mexico. Chihuahua: Portrero [Potrero] Peak, Pringle 825 (GH!; isotypes DS!, GH!, LL, MO!, NY!–2 sheets, US!–3 sheets).

Distinguished from other varieties by the stipes ca. 1⁄2 the blade length; leaf blades 0.8–1 cm wide, 5–7 times longer than wide; midribs abaxially green; plants nearly always epipetric. Distribution. Epiphytic, or sometimes epipetric or in rock crevices, in pine-oak-alder forests; 1500–3600 m. Mexico. Selected Specimens Examined. Ags (McVaugh 18403, MEXU). BCS (Carter 2340, DS, MEXU, UC). Chih (Knobloch 1906, LL, MEXU). Coah (Ginzbarg 199, MEXU, TEX). DF (Lyonnet 898, CAS). Dgo (Breedlove 63043, CAS). Dgo/Sin (McGill et al. 9518, ASU). Gto (Rzedowski 42377, IEB). Hgo (Mexia 2769, CAS, UC). Jal (McVaugh 25916, MEXU). Me´x (Koch 75163, CAS, MO). Mich (Mason 1705, ARIZ, UC). Mor (Barkley et al. 7350, TEX). NL (Hinton et al. 17371, UC). Oax (Bartholomew 3303, CAS). Pue (Diggs 2110, MEXU, UC). Qro (Ferna´ndez N. 2521, CHAPA, ENCB). SLP (Breedlove 66597, CAS, UC). Tam (Stanford et al. 650, ARIZ, UC). Tlax (Sua´rez R. 38, ENCB, MEXU). Ver (Brandegee 319, UC). Zac (Breedlove 63971, CAS). Unverified, Doubtful, or Mistaken Reports. Gro (Catala´n et al. 508b and Dreckmann 59, FCME cited by Lorea-Herna´ndez & Vela´zquez M., 1998, but not verified).

Occasionally this variety grows on rocks or mossy road banks (e.g., Knobloch 1906, from Chihuahua; Ginzbarg 199, from Coahuila; McGill et al. 9518, from Durango/Sinaloa). The distributions of var. polylepis and var. erythrolepis appear to overlap somewhat in northwestern Mexico, and occasional intermediates are encountered. A few specimens have sinuately lobed or pinnatifid blades,

Distribution. Epipetric on limestone rocks and ledges, rarely epiphytic; 1800–2700 m. USA (Tex); Mexico. Selected Specimens Examined. Chih (Nesom & Vorobik 5599, TEX, UC; Correll & Gentry 23026, LL; Correll & Johnston 21642, LL; McVaugh 11509, MEXU). Coah (Johnston 11828, CAS, LL, MEXU; Wynd & Mueller 599, ARIZ, US). Dgo (Breedlove 59101, CAS; Goodding 2146, UC). Son (Hartman 346, UC; Mason 2118, ARIZ, MEXU).

Barkley et al. 7420 (TEX), from Edo. Me´xico, was cited as this variety by Wendt (1980), but it is var. polylepis. Var. erythro-

POLYBOTRYA

lepis was recorded by Hinton and Hinton (1995) from Nuevo Leo´n, but these specimens are all P. polylepis var. polylepis. Occasional specimens are reported as epiphytic, e.g., Nesom & Vorobik 5599 (MEXU), from Chihuahua. There also appear to be varietal intermediates with var. polylepis, e.g., Wendt 478, MEXU, from Coahuila, and Correll & Gentry 23090 (UC), from Chihuahua. 7c. Pleopeltis polylepis var. interjecta (Weath.) E. A. Hooper, Amer. Fern J. 85: 79. 1995. Figs. 229A–D. Polypodium peltatum Cav. var. interjectum Weath., Amer. Fern J. 34: 17. 1944. Pleopeltis macrocarpa (Bory ex Willd.) Kaulf. var. interjecta (Weath.) A. R. Sm., Amer. Fern J. 70: 26. 1980. Pleopeltis interjecta (Weath.) Mickel & Beitel, Amer. Fern J. 77: 21. 1987. Type. Guatemala. Chimaltenango, Standley 60957 (F).

Distinguished from other varieties by the wider (1–1.5 cm) and often longer leaf blades with conspicuously blackened abaxial midribs, laminar scales scattered and rarely overlapping, ca. 0.4 mm diam.; and soral scales ca. 0.6 mm diam., conspicuously blackened in centers with occluded cell lumina, often persistent in mature sori.

Distribution. Epiphytic on trunks, also on rocks and banks; (2050–)2450–3100 m. Mexico; Guat, Salv. Selected Specimens Examined. Chis (Breedlove 15094, 22092, DS, MEXU; Little & Sharp 9829, DS, US). Oax (Davidse & Davidse 9775, CAS, MEXU; Mickel 3752, 5332, UC). Pue (Arse`ne 2179, US; Smith et al. 3856, US). Ver (Barrington 422, MEXU; Nee & Taylor 25893, CAS, XAL; Turra 825, ENCB). Unverified, Doubtful, or Mistaken Reports. Jal (reported by Va´zquez G. et al., 1995, but not verified, probably based on a misidentification). Mich (reported by Pe´rez-Calix, 1996, but probably based on a misidentification).

Records of this taxon from the states of Me´xico, Morelos, and Hidalgo by Mickel and Beitel (1988) are based on misidentified specimens. An unusual specimen with irregular lobes (Oaxaca, Mickel 3784, NY) agrees with var. interjecta in its laminar and rhizome scales, but is sterile, so soral scales and spores cannot be assessed. Martı´nez V. 68 (CHAPA), from Veracruz, is said to be from 900 m, but the stated locality (15 km carr. Jalapa–Perote) is surely much higher, probably ca. 2000 m.

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Pleopeltis polylepis var. interjecta is closely allied to P. mexicana but is distinguished by a combination of characters: blackcentered soral scales, scattered fimbriate laminar scales, and black-centered, non-comose or sparingly comose rhizome scales. Excluded Species Polypodium fallacissimum Maxon, Contr. U.S. Natl. Herb. 17: 567. 1916. This was considered by Mickel and Beitel (1987, crediting Wendt (1980)) to be the putative hybrid between Pleopeltis polylepis var. erythrolepis and Polypodium guttatum. However, we now consider this taxon to be a good species and treat it in Polypodium, which see.

100. POLYBOTRYA Polybotrya Humb. & Bonpl. ex Willd., Sp. Pl., ed. 4, 5(1): 99. 1810. Type: Polybotrya osmundacea Humb. & Bonpl. ex Willd. Hemiepiphytic or terrestrial; rhizomes long-creeping, scaly, in cross section with 4–10 circularly arranged meristeles, each meristele surrounded by a dark sclerenchymatous sheath; fronds holodimorphic; stipes scaly, grooved; sterile fronds 1–3-pinnate, subcoriaceous to coriaceous; veins free to anastomosing; axes grooved adaxially, grooves coalescent; fertile fronds greatly contracted, not foliaceous, 2–4-pinnate, sporangia in masses on lower surface, indusia absent; spores bilateral with a thick, folded surface; x⫽41. Polybotrya is a neotropical genus of 35 species of lowland and montane rain forests, with two rare or uncommon species in Mexico. A third species, P. osmundacea Willd., is widespread in the neotropics and occurs in Guatemala and Belize; it may be found eventually in Mexico. The genus Olfersia (which see) is closely to related and often combined with Polybotrya. Olfersia cervina has parallel veins that fork near the midrib and fuse at the margin (marginal commissural vein), a conform terminal pinna, and sporangia that appear to be on both surfaces of the fertile pinnae, whereas Polybotrya lacks a commissural vein and the veins are not parallel most of their length. At a higher level, Polybotrya and Olfersia are related to the genera Maxonia and Cyclodium in Mesoamerica and South America. At still higher levels, the relationships are to Lastreopsis, Bolbitis, Elaphoglossum, Ctenitis, Stigmatopteris, Dryopteris, Polystichum, and Phanerophlebia, all members of the large, pantropical, subtropical, and temperate dryopteroid clade (Hasebe et al., 1995; Cranfill, unpubl.). Reference Moran, R. C. 1987. Monograph of the neotropical fern genus Polybotrya (Dryopteridaceae). Illinois Nat. Hist. Surv. Bull. 34: 1–138.

Key to the Mexican Species of Polybotrya 1. Sterile blades 2-pinnate-pinnatifid; veins of sterile blades free. ....................................................................................... 1. P. caudata.

476

POLYBOTRYA

1. Sterile blades 1-pinnate, the pinnae serrate-margined to crenately lobed; veins of sterile blades anastomosing.

2. POLYBOTRYA POLYBOTRYOIDES

...................................................................................... 2. P. polybotryoides.

Polybotrya polybotryoides (Baker) Christ, Bull. Herb. Boissier, se´r. 2, 1: 70. 1901. Figs. 230A–C.

1. POLYBOTRYA CAUDATA Polybotrya caudata Kunze, Linnaea 9: 23. 1834. Figs. 230D–H. Olfersia caudata (Kunze) Kunze, Linnaea 21: 206. 1848. Type. Peru. [Hua´nuco:] Pampayaco, Poeppig s.n. (LZ, destroyed; isotype B). Polybotrya villosula Christ, Bull. Herb. Boissier, se´r. 2, 6: 168. 1906. Type. Costa Rica. Santa Clara, Las Delicias, Pittier 10688 (P?). For additional synonymy, see Moran (1987).

Rhizomes densely clothed with brown, linear-lanceolate scales to ca. 1 cm ⫻ 1 mm, margins erose-denticulate; sterile fronds to ca. 1.3 m long, with stipes ca. 1⁄3 the blade length, to ca. 5 mm diam., scaly at stipe bases; sterile blades 2-pinnate-pinnatifid at base, 2-pinnate and ultimately pinnatifid distally; pinnae to ca. 35 ⫻ 12 cm, with pinnules lanceolate, truncate or cuneate at bases, more developed on acroscopic side, pinnules to ca. 8 ⫻ 2.5(–4) cm, larger ones lobed ca. 1⁄3 on acroscopic side, less on basiscopic side, lobes truncate at tip, ca. 5 mm wide; veins pinnate in ultimate segments, free, proximal pairs from adjacent segments meeting margins at the sinuses; laminae hairy on rachises, costae, and costules abaxially and in adaxial grooves, hairs hyaline, acicular, patent to ca. 0.15 mm; fertile blades 2-pinnatepinnatifid, pinnules long-caudate, to ca. 10 cm ⫻ 2–3 mm.

Acrostichum polybotryoides Baker, J. Bot. 19: 207. 1881. Type. Colombia. [Norte de Santander:] Ocan˜a, Kalbreyer 1254 (K; photo GH, MO). Polybotrya aucuparia Christ, Bull. Herb. Boissier, se´r. 2, 6: 166. 1906. Type. Costa Rica. [Cartago:] Valle del Rı´o Navarro, Werckle´ s.n. [Herb. Inst. Costar. 16770] (P; photos F, NY, UC!).

Rhizomes densely clothed with orange to castaneous, subulate scales 10–15 ⫻ 0.5–1 mm, margins denticulate towards scale tips; sterile fronds to ca. 1.3 m long, with stipes 1⁄2 the blade length, scaly at stipe bases; sterile blades pinnate (to bipinnate in Costa Rica), 70–100 cm long, apices pinnatifid; pinnae 10–25 ⫻ 2–6 cm, lanceolate, bases inequilateral, apices cuneate to obtuse, glabrous on both surfaces, margins serrate to crenate or shallowly lobed; veins pinnately branched with arcuate veins, branches anastomosing at acute angles; laminae glabrous abaxially and in grooves adaxially; fertile blades bipinnate, pinnules 1–2 cm ⫻ 2–3 mm; 2n⫽82 (CR).

Distribution. Hemiepiphytic in wet forests; 200–1300 m. Mexico; Guat, Bel, Hond, CR, Pan; Col, Ec, Peru. Selected Specimens Examined. Chis (Breedlove 21886, DS, NY, 38011, DS; Purpus 6761, BM, MO, NY, UC, US, 7245, BM, NY, UC, US). Oax (Herna´ndez G. 821, CHAPA, NY, 964, ENCB, CHAPA, NY, 1966, CHAPA, NY; Torres C. 11183, MEXU). Ver (Croat 40029, MO, UC; Ross 154, US-photo XAL).

Distribution. Hemiepiphytic in lowland and montane rain forests; 300–1200 m. Mexico; Guat, Bel, Hond, Nic, CR, Pan; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, n Braz, Bol. Specimens Examined. Chis (Breedlove 34101, DS, 57629, CAS, ENCB; Martı´nez S. 15945, IEB, 25532, MEXU). Oax (Va´zquez T. 1484, XAL).

Polybotrya caudata is readily distinguished from P. polybotryoides by having 2-pinnate-pinnatifid sterile blades, free veins, and usually hairy axes abaxially.

This species is distinguished from P. caudata by the less dissected, usually 1-pinnate sterile blades, and the anastomosing veins; also, the blades are glabrous or nearly so abaxially. Some forms of this species, especially in Costa Rica and Panama, have the sterile blades fully 2-pinnate. Mexican forms seem to be strictly 1-pinnate. Excluded Species Polybotrya scandens Fe´e, Me´m. Foug. 5: 47. 1852. Type. Alleged to be from Mexico, but possibly from Venezuela (Moran, 1987, p. 119); it may be a synonym of P. osmundacea Willd.

POLYPODIUM

1 0 1 . PO L YPO D I UM Polypodium L., Sp. Pl. 2: 1082. 1753. Lectotype (chosen by J. Sm., Hist. Fil. 88. 1875): Polypodium vulgare L. Marginaria Bory, Dict. Class. Hist. Nat. 6: 587. 1824. Type: Marginaria ceteraccina (Michx.) Bory [⬅ Polypodium ceteraccinum Michx.] ⫽ Polypodium polypodioides (L.) Watt. Goniophlebium sect. Lepicystis J. Sm., J. Bot. (Hooker) 4: 56. 1841. Lepicystis (J. Sm.) J. Sm., London J. Bot. 1: 195. 1842. Type: Lepicystis incana (J. Sm.) J. Sm. [⬅ Goniophlebium incanum J. Sm.] ⫽ Polypodium polypodioides (L.) Watt]. Pseudocolysis L. D. Go´mez, Brenesia 10/11: 116. 1977. Type: Pseudocolysis bradeorum (Rosenst.) L. D. Go´mez [⬅ Polypodium bradeorum Rosenst.]. Microphlebodium L. D. Go´mez, Phytologia 59: 58. 1985. Type: Microphlebodium muenchii (Christ) L. D. Go´mez [⬅ Polypodium muenchii Christ].

Epiphytic or epipetric, infrequently terrestrial; rhizomes short- to long-creeping, freely branched, often glaucous, root proliferations absent; rhizome scales concolorous to bicolorous, clathrate or not, peltately attached, surfaces glabrous or hairy, margins entire or toothed; blades pinnatifid to pinnate, infrequently more divided, monomorphic to subdimorphic, stipitate, articulate on phyllopodia; stipes stramineous, rarely dark, channelled (never terete); blades glabrous, hairy, or scaly; veins free or netted, if netted, areoles regular and each with a single, free, included vein; sori round to oblong, exindusiate, paraphyses (abortive sporangia, or branched or filamentous hairs) present or absent, each sorus terminal on lowest acroscopic vein or free included vein if areolate, in one to several rows between costae and margins; sporangia with paraphyses or glabrous; spores bilateral, verrucate, tuberculate, rugose or papillate; x⫽37. Polypodium, as defined in the narrow sense, comprises about 100–125 species, mostly of the New World tropics with a few species in the temperate regions of America, Europe, Asia and Africa. The species have pinnatifid to pinnate (rarely simple) fronds, peltate rhizome scales, free or netted venation with single free included veins in areoles, sori terminal on ends of veins or

477

on free included veins, channelled stipes, filamentous or branched paraphyses, and branched rhizomes. The limits of the various generic segregates of Polypodium here recognized (q.v.) and keyed below (Campyloneurum, Microgramma, Niphidium, Pecluma, Phlebodium, and Pleopeltis) are problematic. Within Polypodium itself, there are many problematic species complexes, including those of P. subpetiolatum, P. plesiosorum, P. furfuraceum, P. falcaria, P. plebeium, P. polypodioides, and P. thyssanolepis. The group of free-veined, hairy polypodiums is badly in need of revision. The scaly-bladed species are sometimes included in Pleopeltis, as in the recent Flora of North America treatment (Andrews & Windham in FNA Ed. Comm., 1993), but circumscription of Pleopeltis, thus redefined, and the nonscaly species of Polypodium, should await further study. Recent molecular evidence suggests that generic circumscriptions are in need of serious revision (Schneider et al., 2004). References de la Sota, E. R. 1966. Revisio´n de las especies americanas del grupo Polypodium squamatum L. Revista Mus. La Plata, Secc. Bot. 10: 69–186; Evans, A. M. & J. T. Mickel. 1969. A re-evaluation of Polypodium bradeorum and P. colysoides. Brittonia 21: 255–260; Go´mez, L. D. 1977. Contribuciones a la pteridologia centroamericana: 2. Novitates. Brenesia 10/11: 115–119; Go´mez, L. D. 1985. Microphlebium, un nuevo nombre generico (Polypodiaceae). Phytologia 59(1): 58; Haufler, C. H. & T. A. Ranker. 1995 [1996]. RbcL sequences provide phylogenetic insights among sister species of the fern genus Polypodium. Amer. Fern J. 85: 361–374; Haufler, C. H., D. E. Soltis & P. S. Soltis. 1995. Phylogeny of the Polypodium vulgare complex: insights from chloroplast DNA restriction site data. Syst. Bot. 20: 110–119; Hensen, R. V. 1990. Revision of the Polypodium loriceum complex (Filicales, Polypodiaceae). Nova Hedwigia 50: 279–336; Maxon, W. R. 1903. A study of certain Mexican and Guatemalan species of Polypodium. Contr. U.S. Natl. Herb. 8: 271–280; Maxon, W. R. 1916. Studies of tropical American ferns. No. 6. Contr. U.S. Natl. Herb. 17: 541–608; Tejero-Dı´ez, J. D. & L. Pacheco. 2004. Taxa nuevos, nomenclatura, redefinicio´n y distribucio´n de las especies relacionadas con Polypodium colpodes Kunze (Polypodiaceae, Pteridophyta). Acta Bot. Mex. 67: 75–115; Weatherby, C. A. 1939. The group of Polypodium polypodioides. Contr. Gray Herb. 124: 22–35; Wendt, T. 1980. Notes on some Pleopeltis and Polypodium species of the Chihuahuan Desert Region. Amer. Fern J. 70: 5–11.

Key to the Mexican Species of Polypodium 1. Blades scaly, at least abaxially. 2. Sori conspicuously elongate, 2–7 times as long as wide, often interrupted and then variously round, oblong, or linear. .... 4. P. bradeorum. 2. Sori round to slightly oblong, not more than 2 times as long as wide. 3. Blades undivided to pinnatisect with only 1–2 pinna pairs, scales sparse, cordate at bases; laminae strongly coriaceous. 4. Blades undivided; scales absent from sori; Gto. ...................................................................................................... 34. P. microgrammoides. 4. Blades pinnatisect, with 1–2 pinna pairs; scales present in sori; Oax, Chis. .................................................................... 35. P. muenchii. 3. Blades pinnatifid to bipinnate, with 6–40⫹ pinna pairs, or if only 1–3, blades densely scaly; laminae firm to subcoriaceous. 5. Rhizome scales 3–6 mm long, pale and concolorous, or occasionally darkened at point of attachment. 6. Blades 2–3-pinnate, lanceolate; Chiapas. ................................................................................................................. 30. P. lindenianum. 6. Blades pinnatisect to 1-pinnate, narrowly oblong to linear, or nearly deltate. 7. Segments entire, perpendicular to rachises, with or without lime-dots; sori round. 8. Abaxial blade scales round to ovate-lanceolate, occasionally linear-lanceolate in w. Mexico, deeply lacerate, cut half way to the center; stipe scales appressed to slightly spreading, round-fimbriate to ovate-entire; adaxial blade scales stellate to somewhat acicular. ............................................................ 23. P. furfuraceum. 8. Abaxial blade scales linear-lanceolate to acicular; stipe scales patent, lanceolate or linear-lanceolate; adaxial blade scales all distinctly acicular. .................................................................................................... 55. P. villagranii.

478

POLYPODIUM 7.

Segments undulate, crenulate or serrulate, often slightly falcate, generally with lime-dots; sori round or oblong. 9. Blades linear to narrowly oblong; blade scales fimbriate; pinnae 1–2 mm wide, excavate at bases on basiscopic side; stipes (1⁄9)1⁄6–1⁄3 the frond length; sori round; adaxial blade surfaces with scattered scales; rhizome scales 4–5 mm long with reddish brown point of attachment; 100–1000(–1550) m elevation. ......................................................................................................................................................................... 11. P. cryptocarpon.

9.

Blades narrowly deltate; blade scales subentire; pinnae 2–5 mm wide, not excavate at basiscopic bases; stipes 1⁄3–1⁄2 or more the frond length; sori oblong; adaxial blade surfaces glabrous; rhizome scales 5–6 mm long with black sclerotic point of attachment; 1400–2700 m elevation. .......................................... 36. P. platylepis. 5. Rhizome scales 0.5–4 mm, bicolorous with dark, usually well defined central streak of sclerotic cells, or scales concolorous. 10. Blades with scattered to sparse scales abaxially; scales absent adaxially; veins free. 11. Stipes narrowly alate nearly to rhizomes; fronds dimorphic, the fertile fronds slightly to far exceeding the sterile in length and the fertile pinnae distinctly narrower than the sterile. ..................................................... 44. P. remotum. 11. Stipes not alate; fronds monomorphic, the fertile fronds not far exceeding the sterile in length and the fertile pinnae not distinctly narrower than the sterile. 12. Pinnae 7–17 cm ⫻ (8–)10–12 mm, constricted at bases, 2–4 mm wide; pinna apices long-acuminate. .................................................................................................................................................................... 49. P. rzedowskianum.

12. Pinnae 2–6 cm ⫻ 2–8 mm, not especially constricted at bases; pinna apices obtuse to acuminate. 13. Rhizome scales appressed, margins plane to slightly crisped; pinna apices acuminate. ................ 1. P. alansmithii. 13. Rhizome scales spreading, margins strongly crisped and bent out of scale plane; pinna apices obtuse to acuminate. 14. Blade scales sparse, inconspicuous, ca. 0.5 mm long; distal pinnae somewhat ascending. ..... 37. P. plebeium. 14. Blade scales conspicuous, 0.5–1.2 mm long; distal pinnae perpendicular to rachises. 15. Blade scales mostly acicular, the tips often flexuous, pale with dark point of attachment; pinnae 2–3 mm wide; blades usually dark green. ................................................................. 32. P. madrense. 15. Blade scales ovate-lanceolate, orange-tan to brown, without dark point of attachment; pinnae (3–)4–7 mm wide; blades often yellowish green. .................................................... 24. P. guttatum. 10. Blade scales dense abaxially, rarely only scattered; scales scattered or sparse adaxially, rarely absent; veins anastomosing. 16. Rhizome scales black-centered with tan membranous margins. 17. Rhizomes 1–2 mm diam.; blades generally 1.5–6 cm wide, with 6–13(–16) pinna pairs; pinnae 2–3 mm wide. ............................................................................................................................................................. 40. P. polypodioides. 17. Rhizomes generally 3–7 mm diam.; blades generally 6 cm or more wide, if less than 6 cm wide, having only 0–4 pinna pairs; pinnae 5 mm or more wide. 18. Stipes densely scaly, covering the blade surface; veins mostly netted; fronds mostly 10–40(–50) cm long, rarely only 3.5 cm. ...................................................................................................................... 53. P. thyssanolepis. 18. Stipes sparsely scaly, not covering the blade surface; veins mostly free; fronds 3–13 cm long. 19. Blade scales imbricate, ovate- to deltate-caudate, with marginal cells pale and oblique/ ascending from scale center, cells of tail elongate, margins fimbriate with pronounced teeth, often incised through the pale margin; pinnae perpendicular to rachises. ......................... 47. P. riograndense. 19. Blade scales scattered, ovate with cordate bases, black-centered at blade bases to concolorous, cells of margins pale, perpendicular to scale center, scales shallowly toothed, not incised through the pale margins; pinnae oblique to rachises. ......................................................... 19. P. fallacissimum. 16. Rhizome scales concolorous or if bicolorous, reddish brown to castaneous-centered, never with blacksclerotic median portion. 20. Abaxial laminar scales 2–2.5 mm long; longest pinnae 2.5–6.5 cm long. 21. Rhizome scales 1–1.5 mm long, appressed; laminar scales whitish with reddish centers; longest pinnae 5–6.5 cm long. ......................................................................................................................... 50. P. sanctae-rosae. 21. Rhizome scales 3–4 mm long, spreading; laminar scales white with light brown centers; longest pinnae 2–5 cm long. 22. Rhizome scales 3–4 mm long, with black base and linear maroon tail-like tips; generally eastern Mexico. ............................................................................................................................... 43. P. pyrrholepis. 22. Rhizome scales ca. 4 mm long, red-brown, concolorous (weakly bicolorous with dark central streaks with age), linear-lanceolate; generally western Mexico. ........................................................ 48. P. rosei. 20. Abaxial laminar scales (0.3–)1–1.3 mm long; longest pinnae 7–12 cm long. 23. Rhizome scales orbicular, imbricate, 0.5–0.8 mm long; stipe scales round, 0.3 mm diam., to longacicular 1 mm long. .......................................................................................................................................8. P. collinsii. 23. Rhizome scales 2–2.5(–4) mm long, linear to linear-lanceolate, often strongly reflexed; stipe scales with round basal portion 0.3–0.5 mm wide, and long, twisted, hair-like tip 2–4 mm long. ............................................................................................................................................................... 28. P. lepidotrichum.

POLYPODIUM

479

1. Blades glabrous or hairy, not scaly. 24. Veins free, or with casual or irregular anastomoses. 25. Rhizome scales tightly appressed. ................................................................................................................................................38. P. plesiosorum. 25. Rhizome scales spreading, or if somewhat appressed, the scale apices conspicuous. 26. Laminae, rachises, and stipes essentially glabrous abaxially; pinna margins glabrous. 27. Rhizome scales comose; pinnae adnate or cuneate to petiolulate at base; fronds 21–65 cm long. 28. Pinnae adnate; northwestern Mexico (Jal–Son). .......................................................................................... 41. P. praetermissum. 28. Pinna bases cuneate, often petiolulate; widespread. ........................................................................................... 21. P. fraternum. 27. Rhizome scales not comose; pinnae adnate fronds 9–22 cm long. 29. Costae puberulent adaxially; veins free to casually anastomosing; widespread but rare. ............................ 5. P. californicum. 29. Costae glabrous adaxially; veins free; BCN, Chih. ............................................................................................. 26. P. hesperium. 26. Laminae, rachises, and/or stipes with conspicuous hairs abaxially; pinna margins with scattered hairs. 30. Pinna margins serrate to serrulate, rarely entire; pinnae not adnate, at least the proximal ones, or pinnae broadly adnate. 31. Rhizome scales 3–5 mm long, with short scattered hairs on surfaces; sporangia long-setose, the setae 0.3–0.5 mm long. 32. Pinnae narrowly attached, only the distal 1 or 2 pinna pairs broadly adnate. ........................................ 16. P. echinolepis. 32. Pinnae all broadly adnate, basal pinnae occasionally excavate basiscopically. ....................................... 42. P. puberulum. 31. Rhizome scales 4–10 mm long, glabrous or with tuft of long, brown hairs 1–2 mm long at point of attachment (scales comose); sporangia glabrous to short-setose, the setae 0.1 mm long. 33. Pinna margins sharply serrate. .......................................................................................................................... 20. P. fissidens. 33. Pinna margins shallowly serrulate to entire. 34. Sporangia long-setose, setae 0.2–0.3 mm (long as capsules); pinna margins entire or subentire to crenulate proximally. ...................................................................................................................................... 2. P. alavae. 34. Sporangia short-setose, setae 0.1 mm (shorter than capsules); pinna margins crenulate-serrulate to serrulate. 35. Stipes brown; pinnae all adnate and not constricted at their bases, except rarely the proximal pinnae; fronds 65–120 cm long. .................................................................................................... 7. P. christensenii. 35. Stipes stramineous, rarely to brown; pinnae mostly constricted at their bases; fronds 30–75 cm long. .............................................................................................................................................. 52. P. subpetiolatum. 30. Pinna margins entire (slightly undulate to straight); all pinnae adnate. 36. Rhizome scales 10–12 ⫻ 3 mm; sporangia with 0.5–0.8 mm long setae; adaxial laminar surfaces with scattered hairs 1 mm long. ....................................................................................................................................... 33. P. martensii. 36. Rhizome scales 1–8 ⫻ 1–1.5 mm; sporangia glabrous or with minute (less than 0.1 mm long) setae; adaxial laminar surfaces glabrous or with hairs 0.3–0.5 mm long. 37. Sori round; pinnae 3.5–6 mm wide; rhizome scales comose; stipes and rachises with dense hairs 0.8–1 mm long; abaxial costae with hairs 0.3–0.5 mm long; abaxial laminar surfaces with hairs 0.1–1 mm long. 38. Abaxial laminar surfaces with erect 0.1–0.3 mm hairs. .................................................................. 12. P. diplotrichum. 38. Abaxial laminar surfaces with lax 1 mm long hairs. .............................................................................. 6. P. chiapense. 37. Sori oblong; pinnae (6–)8–16 mm wide; rhizome scales not comose; stipes and rachises glabrous or with dense hairs 0.1–0.3 or 1–1.5 mm long; abaxial costae with hairs 0.1 mm long; abaxial laminar surfaces glabrous. 39. Sporangia setose with minute hairs 0.1–0.2 mm or less long; rhizome scales 4–8 mm long with long-attenuate, hair-like tips. 40. Stipes glabrous; rachises with hairs 0.3 mm long; laminar hairs scattered, 0.1 mm; fronds 15– 60(–79) cm long. ....................................................................................................................... 25. P. hartwegianum. 40. Stipes with spreading 1–1.5 mm hairs and rachises with 0.5–1 mm hairs; laminar hairs dense, 1– 1.5 mm long on axes and laminar surfaces; fronds 60–120 cm long. ...................................... 29. P. liebmannii. 39. Sporangia glabrous; rhizome scales 1.5–4 mm long, deltate-lanceolate, lacking hair-like tips. 41. Pinnae mostly constricted at base (narrowly adnate), often auriculate basiscopically (rarely also acroscopically); proximal pinnae not excised basicopically, distal pinnae roughly equally dilated. ...................................................................................................................................... 31. P. longepinnulatum. 41. Pinnae mostly adnate, not auriculate. 42. Proximal pinnae excised basiscopically, strongly surcurrent and adnate acroscopically, distal pinnae surcurrent. .................................................................................................... 10. P. conterminans. 42. Proximal pinnae not excised basiscopically, not strongly decurrent acroscopically. .............. 14. P. dulce. 24. Veins regularly anastomosing, at least in proximal parts of blades and pinnae. 43. Pinnae narrowed at their bases, narrowly adnate to distinct. 44. Sporangia glabrous; rhizome scales 1–3.5 mm wide.

480

POLYPODIUM 45. Sori in 2–5 rows on each side of the costae; rhizome scales clathrate with thickened dark brown walls in the central portion. 46. Rhizomes short-creeping; rhizome scales ovate-acuminate, spreading, especially at apices; distal pinnae broadly adnate. ............................................................................................................................................................ 54. P. triseriale. 46. Rhizomes long-creeping; rhizome scales round to slightly ovate, appressed; distal pinnae weakly or not at all adnate. .................................................................................................................................................................. 22. P. fraxinifolium. 45. Sori in 1 row on each side of the costae; rhizome scales concolorous or with darkened central portion, not clathrate. 47. Sori 1–1.5 mm diam., medial to submedial but not crowding the costae; proximal pinnae with cuneate bases; rhizome scales 3–7 mm long, concolorous, pale brown, brittle, lacking hairs adaxially; margins entire. .. 21. P. fraternum. 47. Sori over 2 mm diam., crowded along costa; proximal pinnae strongly excavate basiscopically; rhizome scales 8–10 mm long, with reddish brown central portion, lighter margins, non-brittle, with minute hairs abaxially on axes and laminae; margins minutely notched. ........................................................................... 39. P. pleurosorum. 44. Sporangia setose, with setae 0.3–0.5 mm long; rhizome scales 1.5–2 mm wide. 48. Rhizome scales minutely pubescent; stipes, rachises, laminar surfaces, and margins all with dense hairs 0.1–0.3 mm long. .......................................................................................................................................................................... 16. P. echinolepis. 48. Rhizome scales glabrous; stipes and rachises glabrous; abaxial surfaces with hairs rare on veins and scattered on costae, 0.3–0.5 mm long, and dense short hairs 0.3 mm long on costae; marginal hairs rare, 0.3–0.5 mm long. ................................................................................................................................................................................. 17. P. eperopeutes. 43. Pinnae not narrowed at their bases, broadly adnate. 49. Stipes winged (1 mm wide) almost to bases. .............................................................................................................. 45. P. rhachipterygium. 49. Stipes unwinged. 50. Rhizome scales clathrate; blade margins entire; pinnae falcate. 51. Sori in a single row between costae and pinna margins; rhizome scales 2.5–3 mm long, appressed, tips acute; laminar surfaces and costae with scattered, minute trichomidia ca. 0.1 mm; 1200–2700 m. .......................... 18. P. falcaria. 51. Sori in 2-4 rows between costae and pinna margins; rhizome scales 7–12 mm long, spreading, with long, slender tips; laminar surfaces with dense hairs 0.5–0.8 mm long, costae with dense hairs 1 mm long; 350– 950 m. .......................................................................................................................................................................... 13. P. dissimile. 50. Rhizome scales not clathrate; blade margins serrate to entire; pinnae straight. 52. Blades strongly coriaceous; Guadalupe Island. ........................................................................................................ 51. P. scouleri. 52. Blade texture chartaceous to firm; continental Mexico. 53. Rhizome scales tightly appressed, concolorous tan to brown, thin, ovate to occasionally lanceolate; laminar surfaces glabrous or densely hairy. 54. Laminar surfaces and margins with hairs 0.3–0.5 mm long; sporangia setose with setae 0.1–0.3 mm long; 300–900 m. .................................................................................................................................. 27. P. hispidulum. 54. Laminar surfaces and margins glabrous; sporangia glabrous; (600–)900–2800 m. 55. Adaxial and abaxial costae and rachises glabrous; veins, rachises, and sometimes stipes reddish. ......................................................................................................................................... 46. P. rhodopleuron. 55. Adaxial and abaxial costae with rare to dense hairs 0.1–0.3 mm long; rachises with dense short hairs 0.1–0.3 mm long; veins, rachises, and stipes stramineous to light brown. ...................38. P. plesiosorum. 53. Rhizome scales spreading, or appressed to slightly spreading, often bicolorous with reddish brown central portion. 56. Proximal pinnae rounded to cordate-clasping basiscopically; pinna margins entire; pinnae 4–8 pairs; veins forming two or more rows of areoles on each side of costa. .............................................. 15. P. eatonii. 56. Proximal pinnae adnate basiscopically, not clasping; pinnae 3–26 pairs; pinna margins crenulateserrulate; veins forming one row of areoles (rarely two) on each side of costa. 57. Stipes and rachises castaneous to brown, at least abaxially; adaxial costae with dense 0.1 mm long hairs; rachises with erect hairs 0.1–0.3 mm long. .................................................................... 9. P. colpodes. 57. Stipes and rachises stramineous to brown; adaxial costae and rachises glabrous except in northern Mexico. ................................................................................................................................. 3. P. arcanum.

1. POLYPODIUM ALANSMITHII Polypodium alansmithii R. C. Moran, Ann. Missouri Bot. Gard. 77: 845. 1990. Type. Mexico. Chiapas: 22 km from San Cristo´bal de las Casas on rd to Tenejapa, then right 3 km on rd to Matzam, Huft et al. 2173 (MO; isotype UC!). Figs. 236J–N.

Rhizomes creeping, 2–4 mm diam.; rhizome scales appressed, 2–2.5 ⫻ 0.5–0.8 mm, bicolorous with black-sclerotic central stripe (0.3 mm wide at base), and margins brown, thin, denticulate, plane to slightly crisped; fronds (10–)16–30 cm long, distant; stipes nearly 1⁄2 the frond length, non-alate, castaneous, grooved, glabrescent to very sparsely scaly; costae pale, nearly flush with the laminar surface; blades broadly ovate, pinnatisect, 5–12 cm wide, proximal pinnae not reduced, somewhat descend-

POLYPODIUM ARCANUM

ing, distal pinnae ascending; pinnae 10–16 pairs, linearlanceolate, acuminate or occasionally obtuse, 2.5–6 cm ⫻ 4–8 mm, broadest in middle, bases largely decurrent, acroscopically subauriculate; adaxial surfaces glabrescent, with lime-dots often in two rows on each side of costa, costae and rachises pale, stramineous, prominent; abaxial surfaces with scattered, brown, lanceolate, round (0.5 mm diam.) to linear-lanceolate (1 mm long), clathrate scales; pinna margins serrate to crenulate; sori round, supramedial; sporangia glabrous.

Distribution. Epiphytic in wet montane forests; 1500–3000 m. Mexico; Guat, Hond, Salv, Nic. Selected Specimens Examined. Chis (Mickel 1242, NY). Gro (Rzedowski 18533, ENCB, 18577, NY). Oax (Ishiki 1290, IEB; Mickel 4342, 4573, 4612, 4928, 5520, 5257, NY).

Polypodium alansmithii is closely allied to P. plebeium but generally occurs at higher elevations and has appressed rather than patent rhizome scales with plane rather than ruffled margins. These characters do not hold strictly, and further work is needed. This species has been called P. montigenum Maxon (type from Costa Rica) (Field Mus. Nat. Hist., Bot. Ser. 17: 306. 1938) in Smith (1981) and Mickel & Beitel (1988), but Moran (in Davidse et al., 1995) pointed out that P. montigenum differs in having pinnae that are obtuse at apices, widest at middle, and somewhat constricted near bases. Polypodium montigenum is confined to Costa Rica and Panama.

2. POLYPODIUM ALAVAE Polypodium alavae A. R. Sm., Amer. Fern J. 70: 21, f. 11–12. 1980. Type. Mexico. Chiapas: ca. 3 mi from Jola along trail from Jola to Chanal, Alava 1287 (UC!; isotypes DS!, MEXU!, MICH!). Figs. 243N, O. Rhizomes creeping, 4–8 mm diam., whitish; rhizome scales tan, deltate-lanceolate, entire to minutely denticulate, 5–7 ⫻ 1.5–2 mm, comose, dark brown and lustrous at point of attachment; fronds 40–90 cm long, distant; stipes ca. 1⁄3 the frond length, stramineous to brown, with short, erect hairs 0.1 mm long; blades pinnatisect, deltate-lanceolate, 14–30 cm wide, the proximal pin-

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nae narrowly attached, distal 1⁄3–2⁄3 of blades with broadly adnate pinnae; pinnae 16–40 pairs, 6–16 cm long, linear, acuminate, 5–13 mm wide, entire or subentire to crenulate proximally; abaxial surfaces pilose on costae and laminar surfaces; adaxial surfaces with erect hairs on costae, sparse on veins and margins; venation free except for occasional anastomoses; sori round, medial, ca. 2 mm diam.; sporangia with 4–8 capsular hairs 0.2–0.3 mm, about as long as the capsules.

Distribution. Epiphytic in pine-oak-Liquidambar forests; 1300– 2000 m. Mexico; Hond. Specimens Examined. Chis (Breedlove 27188, NY, 22337, 32126, 37005, DS; Me´ndez Ton 9284, TEX).

This species is closely related to P. subpetiolatum, but differs in having narrower, more parallel-sided, subentire pinnae, longhaired sporangia, and more coriaceous laminar tissue. It may also be related to P. puberulum but lacks the dense hairs on both laminar surfaces and has proximal pinnae constricted at their bases. Matuda 27227 (MEXU, Edo. Me´xico) resembles P. alavae in its narrow subentire pinnae but has thin blade texture and few short capsular hairs and is probably P. subpetiolatum.

3. POLYPODIUM ARCANUM Polypodium arcanum Maxon, Proc. Biol. Soc. Wash. 51: 38. 1938. Polypodium deltoideum Liebm., Mexic. Bregn. 190 (reprint 38). 1849, hom. illeg., non Sw., 1788. Lectotype (chosen by Tejero, Acta Bot. Mex. In press). Mexico. Puebla: Chinautla, Liebmann s.n. [Pl. Mex. 2598, Fl. Mex. 156]) (C!; isotype GH!).

Rhizomes creeping, 4–7 mm diam.; rhizome scales white to reddish brown with reddish brown bases, deltate, with longattenuate often flexuous-contorted tips, denticulate, especially on the long tips, 4–6 ⫻ 1.5 mm, often with a few minute hairs on the scale surfaces; fronds (7–)15–40 cm long, distant; stipes 1 ⁄3–1⁄2 the frond length, stramineous to brown, glabrous; rachises glabrous except for rare long hairs 0.8–1 mm and rare, clavate brown hairs 0.1–0.3 mm, mostly in the axils of the pinnae; blades

482

POLYPODIUM

pinnatisect, oblong to oblong-deltate, 6–18 cm wide; rachises glabrous or with minute hairs, usually with a few small, contorted, hair-like, reduced scales at bases of proximal pinnae; pinnae 3–26 pairs, linear-deltate, equally dilated at base, 3–9 cm ⫻ 8–18 mm, acute to acuminate, laminae chartaceous, often reddish, especially near costules; margins serrulate to serrulatecrenulate; veins netted, generally with one row of areoles on each side of costae; sori round, medial, one row on each side of costae; sporangia with minute, inconspicuous to conspicuous hairs 0.1 mm. There is considerable variation in number of pinnae in P. arcanum, and this seems to be correlated with geography. Specimens from western Mexico (Michoaca´n, Jalisco, and southwestern Oaxaca) have (8–)12–26 pairs of pinnae whereas those to the east have (3–)4–8(–10) pairs. The former have been called P. plesiosorum var. bakeri based on pinna pair number (10–29 in P. plesiosorum), but clearly resemble P. arcanum in the conspicuous whitish (to tan) rhizome scales. We believe the rhizome scales are more consistent and taxonomically reliable in this group. Polypodium arcanum is similar to P. eatonii, which occurs in Chiapas and Veracruz. The smaller scales, toothed pinna margins, narrower pinnae, and lack of cordate-clasping bases on the proximal pinnae distinguish P. arcanum. Polypodium arcanum is also very closely related to P. colpodes and P. plesiosorum, differing from the former in lacking dark stipes, and from the latter in having fewer pinnae (mostly 4–10 in the east and north, (12–26 in var. bakeri in the west) vs. 15–30(–39) in P. plesiosorum) and having larger, more spreading rhizome scales (vs. appressed and somewhat deciduous in P. plesiosorum). The blades are glabrous or with short minute hairs on the stipes and rachises and generally have sparse to rare small contorted reduced hair-like scales along the rachises at the bases of the proximal pinnae (Fig. 245K), reminiscent of the scale tips on the rhizomes of some specimens. The sporangia have short inconspicuous to conspicuous hairs 0.1 mm. Blades may have a reddish color when young.

Distribution. Epiphytic in pine-oak forests; 1700–2550 m. Mexico. Selected Specimens Examined. Gro (Calonico Soto 4531, FCME). Hgo (Gimate L. 1064, NY). Oax (Mickel 1174, 4054, NY). Pue (Marcks & Marcks 753, NY; Liebmann s.n. [Pl. Mex. 2598, Fl. Mex. 155, 156] C, GH, K; Purpus 2689, UC). Ver (Ventura A. 652, NY).

3b. Polypodium arcanum var. bakeri (Davenp.) Mickel & Tejero, Brittonia 56: 117. 2004. Figs. 245Q, R. Polypodium plesiosorum Kunze var. bakeri Davenp., Gard. & Forest 4: 556. 1891. Type. Mexico. Michoaca´n: Hills of Pa´tzcuaro, Pringle 3352 (GH!; isotypes GH!, K!–2 sheets, MICH!, MO, NY!, P!, UC!, US!).

Differs from var. arcanum in having a greater number of pinnae (8–26 pairs) and a more western distribution.

Key to the Mexican Varieties of Polypodium arcanum 1. Stipes, rachises, and costae sparsely to moderately pubescent abaxially with hairs less than 0.1 mm; pinna pairs (5–)7–15; Hgo to Gto and Coah. ............... 3c. var. septentrionale. 1. Stipes, rachises, and costae glabrous abaxially except for sparse to rare contorted hairs/reduced scales at the bases of proximal pinnae. 2. Pinna pairs (3–)4–8(–10); Oax to Gro and Hgo. ................................................................................... 3a. var. arcanum.

2. Pinna pairs 10–26; western Oax to Jal. ................... 3b. var. bakeri.

Distribution. Mostly epiphytic on oaks in forests of pine and oak; 2100–2500 m. Mexico. Selected Specimens Examined. Jal (Diaz Luna 5144, UC; Pe´rezRulfo T. 17167, UC). Mich (Beaman 2420, MEXU, UC; Rzedowski 2125, XAL, 3995, IEB, 45712, UAMIZ; Santos Martı´nez 2125, IEB, UAMIZ). Oax (Mickel 1174, ISC, UC, 3904, NY).

See var. arcanum for further discussion. 3a. Polypodium arcanum var. arcanum

Figs. 238N–P.

Var. arcanum is distinct by its few pinna pairs and southeastern distribution.

3c. Polypodium arcanum var. septentrionale Mickel, var. nov. Type. Mexico. Queretaro: Mpio. Landa, 2.5 km al suroeste de El Aguacate, Rubio 1243 (holotype NY!). Figs. 245M–P.

POLYPODIUM CALIFORNICUM A var. arcano laminis pilosis necnon distributione boreali diversa. (L., septentrionalis, northern, alluding to the more northern distribution of this variety).

Var. septentrionale differs from var. arcanum in having abaxial laminar hairs less than 0.1 mm, generally (5–)9–15 pinna pairs (vs. (3–)4–8(–10)), and a more northern distribution; 2n⫽74. From P. colpodes, var. septentrionale differs in the stramineous stipes and rachises.

Distribution. Epiphytic or occasionally on rocks in pine-oak or Liquidambar forests; 1400–1730 m. Mexico. Specimens Examined. Coah (Marsh 1926, TEX). Gto (Carranza & Pe´rez 4649, IEB, NY). Hgo (Chase 7307, MICH; Clark 7023, NY; Dunn & Dunn 19063, NY; Pringle 13294, CAS, TEX). NL (Mueller & Mueller 965, US; Rodrı´guez L. 136, TEX). Qro (Carranza G. 2323, NY; Gonza´lez 1350, IEB, MEXU; Gonza´lez P. 352, NY; Herrera 121, MEXU; Rzedowski 46687b, NY; Zamudio & Carranza 6598, NY). Tam (Sharp et al. 52036, US). Tex-Mex border (Wright 758, 2178, NY).

4. POLYPODIUM BRADEORUM Polypodium bradeorum Rosenst., Repert. Spec. Nov. Regni Veg. 10: 279. 1912. Figs. 231A–D, 232A–D, 241A, B. Pseudocolysis bradeorum (Rosenst.) L. D. Go´mez, Brenesia 10/11: 116. 1977. Type. Costa Rica. Prov. Alajuela: Llanuras de S. Carlos, Brade & Brade 460 (S; isotypes UC!, US!). Polypodium pleolepis Maxon & Copel. ex Copel., Univ. Calif. Publ. Bot. 19: 293, t. 46. 1941. Type. Guatemala. Barranca del Rubelcruz, Tu¨rckheim s.n. [Donn. Sm. 673] (US!; isotype MICH!). Polypodium colysoides Maxon & Copel. ex Copel., Univ. Calif. Publ. Bot. 19: 292, t. 45. 1941. Type. Mexico. Veracruz: Atoyac River, Ojo de Agua, Copeland herb. 135 (MICH!; isotypes GH!, NY!, P!, UC!, US!).

Rhizomes creeping, 2–3 mm diam.; rhizome scales lustrous, slightly bicolorous, orange-clathrate center with thick-walled cells, margins paler, thin-walled, lanceolate to linear-lanceolate, erose, ca. 3 ⫻ 0.8–1 mm; fronds 26–40 cm long, 3–10 mm distant; stipes of sterile fronds 1⁄3–1⁄2 the frond length, those of fertile fronds 1⁄2–2⁄3 the frond length, stramineous, with scattered to abundant scales 1–4 mm long (as on rhizomes); sterile blades simple to pinnatisect, simple ones 2–3.2 cm wide; pinnae 0–5 pairs, sterile ones oblong, narrowed at bases, sterile pinnae 1.5– 2.6 cm wide, apices acuminate to attenuate; fertile fronds taller

483

and with narrower pinnae, fertile pinnae 1–1.5 cm wide; abaxial surfaces with scales similar to those on stipes and rhizomes, lustrous, orange, denticulate to fringed, 0.5–2 mm long; adaxial surfaces glabrous; venation netted; sori oblong to linear, 2–10 mm long, often interrupted and the sori then round; sporangia glabrous.

Distribution. Epiphytic and on shaded limestone rocks in wet montane forests; 700–1350 m. Mexico; Bel, Hond, Nic, CR, Pan. Selected Specimens Examined. Chis (Breedlove 22284, DS, F, MEXU, NY, 22496, 23552, DS). Oax (Mickel 7299, NY; Torres C. 5768, MEXU). Ver (Finck 45, US, NY, 199, UC; Palacios-Rios 2900, IEB; Ventura A. 9531, ENCB, IEB).

Evans and Mickel (1969) placed P. colysoides in synonymy under P. bradeorum, though Mickel and Beitel (1988) recanted. Moran (in Davidse et al., 1995) followed the Oaxaca treatment in treating the two as distinct species and even placed them in different genera (bradeorum in Pseudocolysis), believing that P. colysoides had only oblong sori rather than linear interrupted sori. Sori are variable in the species (see Figs. 231B, 232C, 241A, B), however, and the type of P. colysoides has interrupted linear sori. Scales of the rhizomes, stipes, and blades are identical in the two. Generally, specimens with 0–3 pinna pairs have linear, usually interrupted sori, whereas those with 3–5 pinna pairs have oblong sori. Moran (in Davidse et al., 1995) suggested that Polypodium guatemalense Klotzsch (Allg. Gartenzeitung 23: 33. 1855) is a possible synonym of Pseudocolysis bradeorum (Polypodium bradeorum). The tripartite frond form is similar, but the type (Guatemala, Warszewicz s.n., B, digital image at NY!) has very large round sori in rows between pronounced primary veins, as in Niphidium, so the name does not seem to apply to P. bradeorum.

5. POLYPODIUM CALIFORNICUM Polypodium californicum Kaulf., Enum. Filic. 102. 1824. Type. U.S.A. California, Chamisso s.n. (LZ destroyed; isotypes LE!, UC!, also with drawing of LE type). Figs. 238K–M. Rhizomes creeping, 5–10 mm diam.; rhizome scales brown, concolorous, ovate-lanceolate with twisted tips, 5–6 mm long,

484

POLYPODIUM

margins denticulate; fronds 10–22 cm long, clumped; stipes 1⁄3– 1⁄2 the frond length, stramineous, glabrous; blades pinnatifid, lanceolate, 4.5–9 cm wide; pinnae 8–12 pairs, 2–5(–7.5) cm ⫻ 7– 10 mm, tips broadly acute; laminae thin, herbaceous, glabrous except for rare jointed hairs 1 mm long in pinna axils and rare minute, clavate hairs 0.1 mm long along rachises; margins toothed; veins free or with occasional anastomoses; sori round, in a single row on each side of costae; sporangia glabrous; 2n⫽74 (Calif).

Distribution. Epiphytic in lower montane rain forests; (150–) 600–1000 m. Mexico. Specimens Examined. Chis (Breedlove 20281, 21671, DS, 31231, DS, NY, 31537, 32006, DS; Cabrera 7970, MEXU). Oax (Wendt et al. 4764, CHAPA, NY).

7. POLYPODIUM CHRISTENSENII Distribution. Epipetric at high elevations; 100–600 in Baja California, otherwise 2150–3950 m. USA (Calif); Mexico. Selected Specimens Examined. BCN (Van Devender 92-318, ARIZ; Wiggins 4236, CAS, MICH). BCS (Coronados Is., Anthony s.n., in 1895, UC). DF (Lyonnet 85, MEX). Gto (Rzedowski 47916, IEB). Guad (Rose 16015, NY, US). Me´x (Rose & Painter 7946, US). Oax (Mickel 7426, NY). Pue (Reeves R5804, ASC). Qro (Rzedowski 44464, IEB). Rev (Mason 1533, CAS, US). Ver (Beaman 2171, MSC).

Polypodium californicum belongs to the circumboreal P. vulgare complex. It is distinct in its glabrous, pinnatisect blades with free to netted veins. In Mexico, it is most closely related to P. hesperium, which differs in its smaller, narrower, totally glabrous blades, and P. scouleri, confined in Mexico to Guadeloupe Island, which has fewer pinnae and coriaeous blades. Molecular evidence shows that P. plesiosorum is also closely related.

6. POLYPODIUM CHIAPENSE Polypodium chiapense A. M. Evans & A. R. Sm., Amer. Fern J. 70: 23, f. 16–17. 1980. Figs. 244E, F. Pecluma chiapensis (A. M. Evans & A. R. Sm.) M. G. Price, Amer. Fern J. 73: 113. 1983. Type. Mexico. Chiapas: Mpio. Ocozocoautla de Espinosa, Selva del Ocote, 32 km NW of Ocozocoautla, Breedlove 27453 (DS!; isotype NY!)

Similar to P. diplotrichum but with uniform, lax, white hairs 1 mm, rather than minute erect hairs, on abaxial laminar surfaces.

Polypodium christensenii Maxon, Contr. U.S. Natl. Herb. 13: 10. 1909. Type. Guatemala. Alta Verapaz: near Coba´n, Tu¨rckheim II 2179 (US!; isotypes BR!, NY!, US! frag. UC!). Figs. 243H, J–M. Rhizomes creeping, 3–4 mm diam.; rhizome scales orange-tan, lanceolate, attenuate, the margins inrolled, entire, 6–7 ⫻ 1–1.2 mm; fronds 65–120 cm long, 1 cm or more distant; stipes 1⁄4–1⁄2 the frond length, dark brown, with short, whitish hairs, 0.1 mm; blades pinnatisect, apices usually pinnatifid to blade tips, narrowly oblong, 15–30 cm wide; pinnae 23–30 pairs, broadly adnate to rarely narrowly adnate proximally, linear, 7–18 cm ⫻ 1.4–2.2 mm; margins crenulate-serrulate; abaxial surfaces with dense erect hairs ca. 0.5 mm long, hairs on costae 0.6–1 mm; adaxial surfaces with hairs ca. 0.5 mm long; venation free, with rare anastomoses, vein ends conspicuous; sori round, medial; sporangia with hairs 0.1 mm; 2n⫽74 (Chis).

POLYPODIUM COLPODES

Distribution. Epiphytic or on mossy banks, margins of montane rain forests, evergreen cloud forests; 2000–2400 m. Mexico; Guat. Specimens Examined. Chis (Breedlove 32006, DS, NY, 34803, 41087, CAS, DS).

Polypodium christensenii is very similar to P. puberulum but seems to differ in the larger fronds to 1.2 m long; stipes dark brown; pinnae irregularly crenulate-serrate along margins; and perhaps in rhizome scales without a greatly expanded base, not as long-attenuate, spreading. This group of free-veined, hairy polypodiums (including P. subpetiolatum) is badly in need of revision.

8. POLYPODIUM COLLINSII Polypodium collinsii Maxon, Contr. U.S. Natl. Herb. 17: 583, t. 41. 1916. Figs. 235D–F. Goniophlebium collinsii (Maxon) Conz., Fl. Tax. Mex. 1: 106. 1939. Type. Mexico. Chiapas: Near Pantepec, Collins & Doyle 227 (US!). Polypodium mickelii de la Sota, Revista Mus. La Plata, Secc. Bot. 10: 163. 1966. Type. Mexico. Oaxaca: Distrito Villa Alta, 3–5 mi S of Villa Alta on trail to Betaza, Mickel 1130 (ISC!; isotypes LP, NY!).

Rhizomes creeping, 5–7 mm diam.; rhizome scales tightly appressed, round to slightly lance-attenuate, 0.5–0.8 mm long, bicolorous, centers dark reddish brown, occluded, margins light reddish, fimbriate; fronds 50–85 cm long, distant, 1–3 cm apart; stipes 1⁄3–2⁄5 the frond length, castaneous, densely clothed with appressed, round (0.3 mm) to long-acicular, fimbriate scales 1 mm long; blades ovate-lanceolate, pinnatisect, 15–22 cm wide; pinnae 10–15(–27) pairs with terminal pinna conform, lineardeltate, 6–9 mm wide, bases broadly adnate, surcurrent, with nectary on surcurrent edge, apices acuminate; costae castaneous, prominent; adaxial surfaces with scattered acicular fimbriate scales; abaxial surfaces more densely scaly, some scales round, 0.3 mm, others 1 mm long, bicolorous, centers reddish brown, tips whitish; veins not evident; sori round to slightly oblong, pushing through the overlying scales, forming a cup, depressed and making adaxial surfaces embossed; sporangia glabrous; 2n⫽74 (Chis).

485

Distribution. Epiphytic or epipetric in pine-oak forests; 825– 1900 m. Mexico. Specimens Examined. Chis (Breedlove 24924, 23117, DS, NY; Rovirosa 845, NY). Oax ( Galeotti 6512, BR; Mickel 1600, 4821, NY). Ver (Calzada 00261, 00270, CAS).

Polypodium collinsii is distinct from P. sanctae-rosae, the most abundant member of the complex, by having longer pinnae and smaller scales on the rhizomes and abaxial laminar surfaces.

9. POLYPODIUM COLPODES Polypodium colpodes Kunze, Linnaea 23: 176, 316. 1850. Type. Guatemala. Warszewicz 54 (LZ destroyed; isotype B!). Figs. 238Q–S. Polypodium fuscopetiolatum A. R. Sm., Amer. Fern J. 70: 24, f. 13–15. 1980. Type. Mexico. Chiapas: 6–8 km WNW of Soyalo, Breedlove 37155 (DS!).

Rhizomes creeping, 4–6 mm diam.; rhizome scales appressed to slightly spreading, lanceolate, central portion reddish brown, tips whitish, filiform, minutely denticulate, 3–4 mm long; fronds (8–)25–70 cm long, distant; stipes 1⁄5–2⁄5 the frond length, castaneous to brown, at least abaxially, rarely stramineous, glabrous or with rare hairs; blades ovate-deltate to ovate-lanceolate, 5–25 cm wide, deeply pinnatisect to pinnate, apices pinnatifid or terminal pinna subconform; rachises castaneous, pubescent with erect hairs 0.1–0.3 mm; pinnae 8–18 pairs, narrowly deltate, 9– 16 mm wide, bases somewhat surcurrent, apices acute to acuminate, infrequently with sparse to rare contorted hairs (dissected scales) at bases of proximal pinnae (similar to P. arcanum); costae abaxially sparsely to densely hairy, hairs 0.3 mm long, adaxially with dense 0.1 mm long hairs; laminar surfaces glabrous or with stiff erect hairs 0.1 mm; margins crenulate-serrulate; veins netted; sori round, medial or submedial; sporangia glabrous or with inconspicuous single short hairs 0.1 mm.

486

POLYPODIUM

Distribution. Epiphytic and epipetric in oak and pine-oak woods; 300–2600 m. Mexico; Guat, Hond, Salv, Nic, CR, Pan; Col, Ven. Selected Specimens Examined. Chis (Breedlove 22867, NY). Col (McVaugh 16103, NY). Gro (Lorea 1436, US). Jal (Gonza´lez T. 388, NY). Me´x (Hinton 1737, NY, UC). Mich (Hinton et al. 12249, NY). Nay (McVaugh 18898, NY). Oax (Mickel 6294, NY). Qro (Rzedowski 45071, IEB). Sin (Oliver et al. 739, NY).

Polypodium colpodes is distinguished from P. hispidulum by the narrower, somewhat bicolorous, spreading, denticulate rhizome scales and the lustrous, castaneous stipes and rachises (vs. appressed, concolorous scales). It is closest to P. arcanum, from which it differs in the darker stipes and/or rachises and alternate pinnae. This complex is in need of more study.

10. POLYPODIUM CONTERMINANS Polypodium conterminans Liebm., Mexic. Bregn. 201 (reprint 49). 1849. Lectotype (chosen by Smith, 1981: 183). Mexico. Veracruz: “Aguas Santas pr. Mirador,” Liebmann s.n. [Pl. Mex. 2493, Fl. Mex. 139] (C!, photo US!; isolectotype K!). Figs. 244K, L. Polypodium confluens Liebm., Mexic. Bregn. 191 (reprint 39). 1849, hom. illeg., non R. Br., 1810, nec Colla, 1836, nec Hook., 1863, nec Fe´e, 1869. Type. Mexico. [Puebla:] Chinautla, 7000’, Liebmann s.n. [Pl. Mex. 2716, Fl. Mex. 695, 696] (C!, frag. US!, photos NY!, US!). Polypodium surcurrens A. R. Sm., Proc. Calif. Acad. Sci., ser. 4, 40: 223, f. 5A–C. 1975. Type. Mexico. Chiapas: Mpio. Cintalapa, 16 km NW of Rizo de Oro, SW of Cerro Baul, Breedlove 31311 (DS!).

Rhizomes short-creeping, 5–6 mm diam.; rhizome scales lustrous, dark brown, lanceolate-deltate, entire, 1.5–3 ⫻ 1–1.5 mm; fronds 43–60 cm long; stipes 1⁄3–1⁄2 the frond length, castaneous, glabrous abaxially, adaxially with abundant hairs 0.2– 0.3 mm; blades pinnatisect, lanceolate, 16–21 cm wide, thin; pinnae 15–25 pairs, 8–11 cm ⫻ 10–16 mm, linear, strongly surcurrent; abaxial surfaces with scattered hairs 0.1 mm on costae; adaxial surfaces with hairs on rachises, costae, and margins; venation mostly free to mostly netted, dark in at least the proximal portions; sori oblong, medial; sporangia glabrous.

Distribution. Epiphytic or epipetric in wet montane forests; 800–1600 m. Mexico; Hond. Selected Specimens Examined. Chis (Breedlove 21795, DS, ENCB, NY, 24922, DS, NY, 32493, 33118, DS). Pue (Knobloch 2171, MSC; Liebmann s.n., C, photo NY). Ver (Smith 2207, UC; Galeotti 6414, BR; Ventura A. 12990, ENCB, NY, 16831, IEB). Unverified, Doubtful, or Mistaken Reports. Qro (Ferna´ndez 2103, ENCB, cited by Dı´az-Barriga & Palacios-Rios, 1992, but not verified and doubtful).

Polypodium conterminans is distinct by its strongly surcurrent pinna beses.

11. POLYPODIUM CRYPTOCARPON Polypodium cryptocarpon Fe´e, Me´m. Foug. 8: 88. 1857. Type. Mexico. [Veracruz:] Co´rdoba, Schaffner 194 (RB). Figs. 234O–R. Polypodium incanum Sw. β umbrosum Liebm., Mexic. Bregn. 212 (reprint 60). 1849. Type. Mexico. Oaxaca: Dist. Chinantla, Teotalcingo, Liebmann s.n. [Pl. Mex. 2507, Fl. Mex. 110] (C!– mixed with P. furfuraceum). Polypodium skinneri Hook., Sp. Fil. 4: 214. 1862. Type. Guatemala. Skinner s.n. (K!, frag. NY!). Polypodium furfuraceum Schtdl. & Cham. var. rufum, E. Fourn., Mexic. Pl. 1: 83. 1872. Syntypes. Mexico. Veracruz: Cordoba, Bourgeau 1440 (P!), Schaffner 194 (P?); Portero, Hahn s.n. (P?); Mirador, San Francisco, Mu¨ller 144 (P?). Polypodium bernoullii Baker in Hooker & Baker, Syn. Fil., ed. 2, 510. 1874. Type. Guatemala. Dept. Retalhuleu: Bernoulli 442 (K!, frag. NY!; isotypes BR!, NY!, P!–2 sheets, US!).

Rhizomes creeping, 1.5–3 mm diam.; rhizome scales tan, entire to minutely denticulate, lanceolate, 4–5 ⫻ 1–1.5 mm with reddish brown point of attachment; fronds 12–30 cm long, distant; stipes (1⁄9–)1⁄6–1⁄3 the frond length, castaneous to atropurpureous, with a mixture of small, round, appressed scales and larger ovatelanceolate, spreading scales; blades pinnatisect, linear to narrowly oblong, 1.5–4 cm wide; pinnae 25–60 pairs, 1–2 mm wide, obtuse, often excavate on the basiscopic side; adaxial blade surfaces glabrescent with sparse acicular scales; abaxial blade surfaces with scattered whitish acicular scales 1 mm long, bases fimbriate, cut less than halfway to center, dilated, centers black or sometimes brownish, margins white, sometimes brown; veins obscure; sori

POLYPODIUM DISSIMILE

487

round, appearing to extend beyond the pinna margins; sporangia glabrous.

Distribution. Epiphytic in wet montane forests; 1375–1450 m. Mexico. Distribution. Epiphytic in wet montane forests; 100–1000(– 1550) m. Mexico; Guat, Hond. Selected Specimens Examined. Chis (Breedlove 26349, NY; Ventura & Lo´pez 527, XAL). Oax (Herna´ndez G. 944, CHAPA; Mickel 6159, NY). Pue (Lyonnet 460900008, MEXU). Ver (Copeland herb. 140, NY; Couch M32, US; Mickel 735b, NY; Va´zquez T. et al. V-2415, CHAPA; Wendt et al. 2737, CHAPA, ENCB). Unverified, Doubtful, or Mistaken Reports. Mich (Rzedowski 37392, IEB, cited by Dı´az-Barriga & Palacios-Rios, 1992, but not verified and doubtful). Qro (Ferna´ndez N. 3417, IEB, cited by Dı´azBarriga & Palacios-Rios, 1992, but not verified and doubtful).

Polypodium cryptocarpon has abaxial scales that vary from lanceolate to acicular and are more shallowly fimbriate than those of P. furfuraceum, being cut ca. 1⁄4 the way to the middle. In addition, the pinnae are nearly glabrous adaxially. Polypodium cryptocarpon has undulate (vs. straight) pinna margins and narrow, often slightly falcate or slightly ascending pinnae. Abaxial scales are scattered with short teeth (vs. scales dense, larger, with hair-like teeth in P. furfuraceum), the adaxial blade scales sparse to absent (vs. scattered).

12. POLYPODIUM DIPLOTRICHUM Polypodium diplotrichum Mickel & Beitel, Mem. New York Bot. Gard. 46: 297. 1988. Type. Mexico. Oaxaca: Distrito Juquila, 28–29 km N of San Gabriel, Mickel 6181 (NY!; isotype UC!). Figs. 235L, M. Rhizomes creeping, 6–7 mm diam.; rhizome scales dark brown, lanceolate, 2–3 mm long, spreading, comose; fronds 20–25 cm long, distant; stipes and rachises with dense long hairs 0.8–1 mm long; blades pinnatisect, 5–6 cm wide; pinnae 25–30 pairs, 4–6 mm wide, shorter at blade bases, adnate, apices rounded-acute, margins entire; abaxial laminar surfaces with dense hairs 0.1–0.3 mm long, costae with dense hairs 0.3–0.5 mm long; adaxial surfaces, margins, and midribs with scattered hairs 0.3–0.5 mm long; veins free; sori round; sporangia glabrous.

Specimens Examined. Chis (Purpus 6754, NY, UC p.p., US, 7225, UC, US). Oax (only the type collection).

Similar to P. hartwegianum and P. longepinnulatum, P. diplotrichum differs in its long hairs on stipes, rachises, and laminar surfaces. It resembles P. chiapense in most features, including frond form, comose rhizome scales, and long-hairy stipes, but is distinct in having numerous short erect hairs 0.1–0.3 mm long on the abaxial laminar surfaces and hairs 0.3–0.5 mm long on the costae. Polypodium chiapense has scattered hairs 0.8–1 mm long on its costae and abaxial laminar surfaces. That species was transferred to Pecluma by M. G. Price (Amer. Fern J. 73: 109– 116. 1983); however, both P. chiapense and P. diplotrichum have the essential Polypodium characters of long-creeping rhizomes and channelled axes, and lack dark axes and clavate soral paraphyses, characteristic of Pecluma.

13. POLYPODIUM DISSIMILE Polypodium dissimile L., Syst. Nat., ed. 10, 2: 1325. 1759. Type. Jamaica. Browne s.n. (LINN 1251.24). Lellinger (Amer. Fern J. 67: 100. 1977, op. cit. 70: 30. 1980) recently discussed the earlier misapplication of this name. Figs. 239K–M. Polypodium chnoodes Spreng., Neue Entd. Pflanzenk. 3: 6. 1822. Goniophlebium chnoodes (Spreng.) Fe´e, Me´m. Foug. 5: 255. 1852. Type. Martinique. Kohaut s.n. (Sieber exsicc.) (LZ destroyed).

Rhizomes creeping, 6–14 mm diam.; rhizome scales clathrate, 7–12 mm long, long-attenuate; fronds 20–100 cm long, distant; stipes ca. 1⁄4 the frond length, stramineous, with abundant short white hairs 0.3–0.5 mm long; blades 1-pinnate, narrowly oblong, 6.5–22 cm wide, apices pinnatifid, margins entire; pinnae (12–) 30–36 pairs, linear-deltate, falcate, acuminate, broadly adnate, generally not decurrent or surcurrent, proximal pinnae constricted basiscopically; laminae on both sides evenly and densely pubescent with short white hairs 0.5–0.8 mm long, costae and rachises with longer dense hairs 1 mm long, margins slightly undulate; veins netted with 3–4 rows of areoles between costae and margins; sori round, in (2–)3(–4) rows between costae and margins; sporangia glabrous; 2n⫽74 (Jam).

488

POLYPODIUM Specimen Examined. Chis (Croat 47759, MO, cited by Moran in Davidse et al., 1995, but not verified; Stevens & Martı´nez S. 25859, NY).

15. POLYPODIUM EATONII Polypodium eatonii Baker in Hooker & Baker, Syn. Fil., ed. 2, 511. 1874. Figs. 245C–E. Polypodium ghiesbreghtii D. C. Eaton, Proc. Amer. Acad. Arts 8: 618. 1873, hom. illeg., non Hook. & Baker, 1867. Goniophlebium eatonii (Baker) Maxon, Contr. U.S. Natl. Herb. 16: 60. 1912. Type. Mexico. Chiapas: Ghiesbreght 273 (YU!–2 sheets, photos BM!, P!; isotypes BM!, GH!, K!, NY!, PH!). Goniophlebium pringlei Maxon, Proc. U.S. Natl. Mus. 27: 954. 1904. Type. Mexico. Veracruz: near Jalapa, Pringle 11855 (US!; isotypes ACU, ASU!, CAS!, F, GH!, MICH, MO, P!, UC!).

Distribution. Epiphytic in wet montane forests; 350–950 m. Mexico; Guat, Bel, Hond, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Ec. Specimens Examined. Chis (Breedlove 27949, NY; Stevens & Martı´nez S. 25859, NY, UC). Oax (Hallberg 1281, NY, US, 1573, NY; Mickel 6348, NY).

Polypodium dissimile is distinguished by its clathrate rhizome scales, densely hairy blades, and broadly adnate pinnae.

14. POLYPODIUM DULCE

Rhizomes creeping, 4–5 mm diam.; rhizome scales whitish with orange bases, lanceolate-acuminate, subentire with sparse minute teeth, spreading, 5–7 ⫻ 1 mm at bases; fronds 24–34 cm long, distant; stipes 1⁄4–2⁄5 the frond length, stramineous to tan, glabrous; blades pinnatisect, deltate to lanceolate, 10–13 cm wide; pinnae 4–6(–8) pairs, 5–8 ⫻ 1.5–2 cm, oblong, apices broadly acute, proximal pinnae and occasionally the next with basiscopic bases rounded to cordate, usually clasping the rachises; margins entire; abaxial and adaxial surfaces glabrous; venation netted, forming 2 or more rows of areoles between costae and margins; sori round, medial, in one row on each side of costae; sporangia glabrous; 2n⫽74 (Ver).

Polypodium dulce Poir. in Lamarck, Encycl. 5: 523. 1804. Figs. 232E, F. Chrysopteris dulcis (Poir.) Fe´e, Me´m. Foug. 5: 265. 1852. Lectotype (chosen by Proctor & Lourteig, Bradea 5: 387. 1990). Plumier, Traite´ Foug. Ame´r., pl. 80, 1705, illustrating a plant from Martinique. Polypodium sororium Humb. & Bonpl. ex Willd., Sp. Pl., ed. 4, 5(1): 191. 1810. Type. [Venezuela.] Nova Andalusia prope Caripe, Humboldt & Bonpland s.n. (B-Willd. 19684-1).

Similar to P. longepinnulatum in the thin pinnatisect blades with free to casually anastomosing veins and round to oblong sori, but distinct in the pinnae being broadly adnate without basal auricles.

Distribution. Epiphytic in pine-oak forest, 1200–1700 m. Mexico. Specimens Examined. Chis (Breedlove 6210, DS, US; Mu¨nch 61, DS; Thorne & Lathrop 41060, DS). Pue (Tenorio L. 12600, IEB). Ver (Ventura A. 7110, 8918, CHAPA, ENCB, NY, 16699, IEB).

Some Chiapas and Veracruz specimens do not have the cordate-clasping segment bases mentioned by Maxon but otherwise agree with his characterization. See P. arcanum for further discussion.

16. POLYPODIUM ECHINOLEPIS Distribution. Epiphytic in wet forests; 0–1500 m (Moran in Davidse et al. 1995). Mexico; Guat, Bel, Hond, Nic, CR; Col, Ven, Ec, Peru, Bol.

Polypodium echinolepis Fe´e, Me´m. Foug. 8: 87. 1857. Type. Mexico. [Veracruz:] Cordoba, Schaffner 188 (RB). Figs. 235N, O.

POLYPODIUM FALCARIA Polypodium sessilifolium Liebm., Mexic. Bregn. 192 (reprint 40). 1849, hom. illeg., non Desv., 1827. Type. Mexico. Veracruz: “Barranca de Huitamalco,” Liebmann s.n. [Fl. Mex. 697] (C!, frag. US!). Polypodium insigne Liebm., Mexic. Bregn. 192 (reprint 40). 1849, hom. illeg., non Blume, 1828. Polypodium plectolepis Hook., Sp. Fil. 5: 30. 1863. Lectotype (chosen by Smith, 1981: 185). Mexico. Veracruz: “Hacienda de Mirador,” Liebmann s.n. [Pl. Mex. 2719, Fl. Mex. 700) (C!; isotypes B, frag. NY!, K!, frag. US!). Goniophlebium plectolepis Fe´e, Me´m. Foug. 8: 95. 1857. Type. Mexico. [Veracruz:] “Pre`s d’Orizaba,” Schaffner 187 RB). Polypodium plectolepis Fe´e var. incisum E. Fourn., Mexic. Pl. 1: 78. 1872. Syntypes. Mexico. Veracruz: Orizaba, Bourgeau 3361 (P!–3 sheets), Schaffner 187 (P?), Weber s.n. (P?); Hacienda de Mirador, Liebmann s.n. (P?); Cordoba, Bourgeau 1430, 1791, (P?).

Rhizomes creeping, 6–10 mm diam.; rhizome scales orangebrown, deltate-attenuate, margins of tips curled, minutely pubescent on outer surfaces, spreading, 3–5 ⫻ 1.5–2 mm; fronds 30– 85 cm long; stipes 1⁄5–2⁄5 the frond length, stramineous to light brown, with scattered to dense white hairs 0.1–0.3 mm long; blades 1-pinnate, 15–26 cm wide, terminal pinna pinnatifid; pinnae 9–20 pairs, 1.4–2.5 cm wide, linear-lanceolate; rachises, costae, and laminar surfaces subglabrous or with dense hairs 0.1–0.3 mm long; margins serrulate with hairs 0.1–0.3 mm long; veins free to casually anastomosing; sori round, medial, in one row on each side of costae; sporangia setose with hairs 0.3 mm long; 2n⫽74 (Ver).

489

17. POLYPODIUM EPEROPEUTES Polypodium eperopeutes Mickel & Beitel, Mem. New York Bot. Gard. 46: 298. 1988. Type. Mexico. Oaxaca: Distrito Mixe, vicinity of Zacatepec, along trail ca. 1 mi. N of city, Mickel 1595 (NY!; isotype NY!). Figs. 239D–F. Similar to P. echinolepis except: rhizome scales glabrous with planar margins; stipes and rachises glabrous; abaxial laminar surfaces glabrous except for sparse hairs 0.3–0.5 mm long on veins and scattered hairs 0.3–0.5 mm long on costae; adaxial laminar surfaces with scattered hairs 0.3–0.5 mm long on surfaces and scattered to dense hairs 0.3 mm long on costae; pinnae mostly net-veined; sporangia with occasional setae.

Distribution. Epiphytic, epipetric, and terrestrial in wet montane forests; 1250–1800 m. Mexico. Specimens Examined. Chis (Croat 40719, UC). Me´x (Matuda 27227, US). Oax (only the type collection). Ver (Copeland herb. 133, US; Croat 39547, UC; Ventura A. 4904, NY).

Distribution. Epiphytic in wet forests; 500–1600 m. Mexico; Guat, Hond, Nic, CR, Pan. Selected Specimens Examined. Chis (Breedlove 29135, NY, 32906, MEXU). Mich (Escobado 1369, NY). Mor (Lyonnet 500500002, MEXU). Oax (Mickel 1430, 7266, NY). Pue (Riba et al. 282, NY). Ver (Bourgeau 1430, 3070, NY; Ventura A. 15797, MEXU).

See key and description of P. eperopeutes for comparison. Polypodium echinolepis is most closely related to P. subpetiolatum but is distinct by its variable venation (often anastomosing or casually so, but sometimes totally free) and rhizome scales having minute surficial hairs. Rarely the rhizome scales of P. subpetiolatum are slightly papillose, obscuring the difference between the two species. More work is needed. Polypodium echinolepis resembles P. puberulum in its papillose rhizome scales, but the latter is distinct by its broadly adnate pinnae.

Polypodium eperopeutes is distinct by the usually regular vein anastomoses, glabrous rhizome scales, and glabrous blades. We now regard most of the specimens originally cited under P. eperopeutes (Mickel & Beitel, 1988) as P. longepinnulatum; they were mistakenly placed under P. eperopeutes because of the variable character of net venation. Net venation is apparently more frequent in larger specimens of P. longepinnulatum than previously thought.

18. POLYPODIUM FALCARIA Polypodium falcaria Kunze, Linnaea 18: 316. 1844 [1845]. Lectotype (chosen by Hensen 1990: 300). [Mexico. Veracruz:] Leibold 90b (LZ destroyed; isotypes B!, G, photo US!, P!). Figs. 238D–F. Goniophlebium calaguala Fe´e, Me´m. Foug. 8: 93. 1857. Type. Mexico. Valle´e du Mexique, Schaffner 184 (P?; isotype K). Goniophlebium invertens Fe´e, Me´m. Foug. 9: 25. 1857. Type. Mexico. [Veracruz:] Orizaba, Schaffner 494 (P?)

490

POLYPODIUM

Rhizomes creeping, 3–5 mm diam.; rhizome scales clathrate with pale narrow margins, thin, lanceolate-deltate, appressed, peltate, 2.5–3 ⫻ 1–1.5 mm, rather loosely attached, the rhizomes slightly exposed with age; fronds (15–)30–85 cm long; stipes ca. 1⁄4–2⁄5 the frond length, stramineous to brown, glabrous except for scattered trichomidia, or with very sparse clathrate scales near bases; blades pinnatisect, narrowly deltate, 5–25 cm wide, proximal pinnae reflexed; pinnae 10–32 pairs, broadly adnate, lineardeltate, falcate, (5–)8–11(–15) mm wide, apices acuminate, bases surcurrent, surfaces glabrous except for scattered trichomidia 0.1 mm long on abaxial surfaces and costae adaxially; margins slightly undulate, without teeth, with scattered hairs 0.3 mm; veins netted, with one row of areoles between costae and margins; sori round, submedial, in one row between costae and margins; sporangia glabrous.

⫻Pleopodium fallacissimum (Maxon) Mickel & Beitel, Amer. Fern J. 77: 20. 1987. Type. Mexico. Coahuila: San Lorenzo Canyon, 6 mi southeast of Saltillo, Palmer 426 (US!; isotypes F!, GH!, NY!, UC!–2 sheets).

Rhizomes creeping, 1 mm diam.; rhizome scales bicolorous, each with black-clathrate center and narrow pale fringed margins, tan, lanceolate, 1.5–2 ⫻ 0.5 mm; fronds 3.5–9.5 cm long, subapproximate; stipes ca. 1⁄3 the frond length, stramineous, with scattered scales grading from black-centered at base to concolorous but still with thicker walled isodiametric cells in center and elongate cells at fringed margins, lanceolate with cordate bases; blades pinnatifid, elliptic, (0.5–)1–2 cm wide, decurrent at bases; pinnae (0–)1–2 pairs, 8–20 ⫻ 2.5–4 mm, narrowly oblong and ascending at ca. 45⬚ to rachises; abaxial surfaces with scattered to dense scales as on stipes, ovate to lanceolate, concolorous but with thick-walled isodiametric cells; adaxial surfaces glabrous, lime-dots absent; veins obscure; sori round, medial; sporangia glabrous.

Distribution. Epiphytic in wet montane forests, largely on the Atlantic side; 1200–2700 m. Mexico; Guat, Hond, Salv. Selected Specimens Examined. Chis (Breedlove 22637, NY). Gro (Rzedowski & McVaugh 28, NY). Oax (Mickel 1527, 5028, NY; Rzedowski 33367, NY). Pue (Rzedowski 29977, NY; Ventura A. 20488, NY). Ver (Bourgeau 3187, NY; Pringle 6083, NY; Ventura A. 9213, NY).

Among the net-veined polypodiums with adnate pinnae, P. falcaria is distinct by its falcate pinnae and tightly appressed clathrate rhizome scales. Hensen (1990) treated P. falcaria as synonymous with P. latipes Langsd. & Fisch. (type from Brazil), but the latter has less falcate, often apically rounded pinnae and longattenuate rhizome scales. The name P. loriceum L. (type from Martinique) has generally been applied to this plant. Material from the Antilles, Venezuela, Colombia, and north to Nicaragua has round, appressed rhizome scales and often two rows of sori, whereas Mexiconorthern Central American plants have lanceolate rhizome scales and single rows of sori. More work is needed on this complex.

Distribution. Epipetric on rocky slopes in open oak-ash woods and matorral; 2150–2200 m. Mexico. Selected Specimens Examined. Coah (Arce s.n., 31 Aug 1986, ANSM; Cowan 3806, TEX, UC; de la Rosa I. & Villarreal 1607, ANSM, MSC, XAL, UC; Koch & Gonza´lez L. 8655, NY; Lyonnet 43100001, MEXU, TEX, UC; Palmer 425, F, UC; Villarreal & Carranza 1729, ANSM). NL (Hinton 20138, TEX; Ramirez A. s.n., 1 Feb 1980, UNL).

This species closely resembles Polypodium riograndense and small specimens of P. thyssanolepis, but is distinct in its pinnae being oblique to the rachis and the large laminar scales. Polypodium fallacissimum is like P. guttatum in its yellow-green blades and fringed blade scales. Polypodium riograndense also differs in having caudate blade scales with pronounced teeth.

20. POLYPODIUM FISSIDENS

19. POLYPODIUM FALLACISSIMUM

Polypodium fissidens Maxon, Contr. U.S. Natl. Herb. 8: 275. 1903. Type. Mexico. Chiapas: Ghiesbreght 244 (YU!; isotypes BM!, GH!, K!, NY!, PH!). Figs. 243C–G.

Polypodium fallacissimum Maxon, Contr. U.S. Natl. Herb. 17: 567. 1916. Figs. 242F–H, J.

Rhizomes creeping, 4 mm diam., glaucous; rhizome scales orange-tan, lanceolate, entire, 4–5 ⫻ 2 mm; fronds 17–62 cm long, distant; stipes 1⁄4–1⁄3 the frond length, stramineous to cas-

POLYPODIUM FRAXINIFOLIUM

491

taneous, glabrous; blades pinnatisect, deltate, 5.5–25 cm wide, apices pinnatifid; pinnae 13–30 pairs, 2.5–13 cm ⫻ 5–11 mm, linear, broadly adnate, the proximal pinnae slightly constricted at bases, margins sharply serrate; abaxial blade surfaces with erect hairs 0.3–0.5 mm scattered on surfaces, dense on costae and rachises; adaxial blade surfaces with sparse hairs; veins free, ending in coarse teeth; sori round, medial; sporangia setose with hairs 0.1 mm.

Distribution. Epiphytic in wet forests; (300–) 900-2200 m. Mexico; Guat, Bel, Hond, Nic, CR. Selected Specimens Examined. Chis (Breedlove 23279, NY). Gro (Otonio O. 280, FCME). Hgo (Go´mez M. 2145, NY). Jal (McVaugh 16180, MICH). Mich (Labat 2490, IEB). Nay (Feddema 932, MICH). Oax (Mickel 4747, NY). Pue (Ventura A. 171, NY). Qro (Ferna´ndez N. 4433, NY). SLP (Pringle 4001, NY). Ver (Mu¨ller 1812, NY).

Distribution. Epiphytic in evergreen cloud forests; (2400–) 2700–2900 m. Mexico; Guat, Hond, Salv. Selected Specimens Examined. Chis (Breedlove 21996, 22006, 23021, 32009, DS, NY, 32061, 32068, DS, 69852, CAS; Gonza´lez-Espinosa et al. 580, CHAPA; Mickel 1241, NY; Mu¨nch 66, DS).

Polypodium fissidens is distinct by the sharp teeth of the pinna margins.

21. POLYPODIUM FRATERNUM Polypodium fraternum Schltdl. & Cham., Linnaea 5: 608. 1830. Type. Mexico. [Veracruz:] Near Jalapa, Schiede & Deppe [753] (B!; isotype LE!). Figs. 237K, L, O, P. Goniophlebium revertens Fe´e, Me´m. Foug. 8: 94. 1857. Polypodium revertens (Fe´e) E. Fourn., Mexic. Pl. 1: 79. 1872. Syntypes. Mexico. [Veracruz:] Huatusco, Schaffner 189 (P; isotype K!), Orizaba, Schaffner 450 (presumably P, not found; isotype K!). Polypodium petiolatum Davenp., Bot. Gaz. 19: 394. 1894. Type. Mexico. San Luis Potosı´: Las Canoas, Pringle 4001 (GH!, VT, photo US).

Rhizomes creeping, 6–7 mm diam., often glaucous; rhizome scales pale reddish brown, ovate-lanceolate to long-attenuate, 3–6 mm long, margins irregularly denticulate; fronds 25–65 cm long, distant; stipes ca. 1⁄3 the frond length, pale brown, glabrous; blades broadly oblong, pinnate, 22–40 cm wide with subconform terminal pinna; pinnae 6–15 pairs, somewhat ascending, linearelliptic, 8–22 ⫻ 0.8–1.8 cm, coriaceous, often petiolulate, the petiole to 8 mm long, glabrous, costae lighter (rarely darker) than the laminae, bases cuneate, apices acuminate; margins entire to subcrenulate; veins netted to free; sori round to oblong, 1–1.5 mm diam., in one row on each side of costae, submedial; sporangia glabrous; 2n⫽74 (SLP).

Polypodium fraternum is highly variable in the degree of vein anastomosis, but is distinct by its glabrous blades, cuneate-based pinnae, sori in a single row on each side of the costae and noncomose rhizome scales.

22. POLYPODIUM FRAXINIFOLIUM Polypodium fraxinifolium Jacq., Collectanea 3: 187. 1789 [1791]. Figs. 240A–D. Goniophlebium fraxinifolium (Jacq.) T. Moore, Index Fil. lxxiv. 1857. Type. Cultivated specimen, Hort. Scho¨nbrunn, from specimen said to have originated from Caracas (W; photo BM!).

Rhizomes long-creeping, 5–6 mm diam.; rhizome scales brown-clathrate with pale margins, round to slightly ovate, entire, 1–2 mm long; fronds 45–85 cm long, 1–3.5 cm distant; stipes 2 ⁄5 the frond length, stramineous to orange, glabrous; blades 1pinnate, lanceolate-deltate, 20–30 cm wide; pinnae 5–9 pairs, 12– 18 cm ⫻ 18–33 mm, lanceolate to elliptic, distal pinnae not at all or only slightly adnate, margins entire; abaxial and adaxial blade surfaces glabrous; venation meniscioid, with 3–5 rows of areoles between costae and margins; sori round, medial, in 3–5 rows between costae and margins; sporangia glabrous; 2n⫽74 (CR).

492

POLYPODIUM

adaxial surfaces glabrescent, often with lime-dots; sori round, medial, sporangia glabrous.

Distribution. Terrestrial in Mexico or epiphytic in pine-oak forests; 1300 m. Mexico; Guat, Bel (Davidse & Holland 36729A, MO), Nic, CR, Pan; Col, Ven, Guy, Sur, Fr Gui, Ec, Peru, s Braz, Bol. Specimens Examined. Chis (Breedlove 32151, DS, MICH).

This species is close to P. triseriale except: distal pinnae not or only slightly adnate, pinnae elliptic with 3–5 rows of sori/ areoles between costae and pinna margins (vs. only 2 or occasionally 3), rhizome scales appressed, with rounded apices (vs. spreading, lanceolate, attenuate), and longer-creeping rhizomes, (2–)4–10 cm between stipe bases, rhizomes thinner, ca. 5–6 mm diam. (vs. 7–10 mm).

23. POLYPODIUM FURFURACEUM Polypodium furfuraceum Schltdl. & Cham., Linnaea 5: 607. 1830. Type. Mexico. [Veracruz:] Near Jalapa, Schiede & Deppe [750] (B!; isotypes LE!, P!). Figs. 234A–D, 241K–M. Polypodium nivosum Fe´e, Me´m. Foug. 8: 89. 1857. Type. Mexico. [Veracruz:] Huatusco, Schaffner 193 (presumably P, not found; isotype K!). Polypodium furfuraceum Schltdl. & Cham. var. coronulatum E. Fourn., Mexic. Pl. 1: 83. 1872. Type. Mexico. [Veracruz:] Jalapa, Hahn s.n. (P!). Polypodium furfuraceum Schltdl. & Cham. var. rufum E. Fourn., Mexic. Pl. 1: 83. 1872. Type. Mexico. [Veracruz:] Potrero, 19 August 1869, Hahn s.n. (P!). Polypodium margallii Rovirosa, Pteridogr. Sur Me´xico 206, t. 38a, f. 1–5. 1909. Type. Mexico. Chiapas: Mesa de Coapilla, Rovirosa 1061 (isotype PH!).

Rhizomes creeping, 1.5–2 mm diam.; rhizome scales concolorous tan, ovate to lanceolate, entire to minutely denticulate, 3–5 ⫻ 1 mm; fronds 23–48 cm long, distant to subapproximate; stipes 1 ⁄6–1⁄4 the frond length, castaneous to atropurpureous, with some scales lanceolate and spreading as on rhizomes, others small, appressed, round, peltate, laciniate, as on blades; blades pinnatisect, narrow to linear-oblong, 2–7 cm wide; pinnae 27–40 pairs, 12–28 mm long, linear, costae and veins not visible, sterile pinnae 2–4 mm wide, fertile pinnae ca. 2 mm wide; abaxial surfaces with dense, round to ovate-lanceolate (occasionally linearlanceolate in w. Mexico), fimbriate scales cut 1⁄2 or nearly to center, white with dark center, to 1.5 ⫻ 0.5–1 mm; rachises with similar and also larger ovate scales with fimbriate margins;

Distribution. Epiphytic in wet montane forests, dry oak forests; 750–2300 m. Mexico; Guat, Hond, Salv, Nic, CR, Pan; Peru, Bol. Selected Specimens Examined. Chis (Breedlove 22158, NY). Col (Espero´n 18291, GUADA). Dgo (Bashor 665, NY). Gro (Rzedowski 22652, NY). Hgo (Clark 7022, NY). Jal (Palafox 5407, NY). Me´x (Rzedowski 20777, NY). Mich (Pringle 3357, NY). Mor (Rzedowski 19778, NY). Nay (Feddema 931, NY). Oax (Mickel 3701, NY). Qro (Ferna´ndez N. 3417, NY). Sin (Breedlove 18246, NY). SLP (Rzedowski 24557, NY). Ver (Purpus 2167, NY).

Polypodium furfuraceum most closely resembles P. cryptocarpon in its densely scaly blades and pale, concolorous rhizome scales but is distinguished by the segments perpendicular to the rachises (vs. somewhat falcate or ascending) and the deeply lacerate abaxial blade scales (vs. cut less than halfway to costa); entire margins (vs. often crenulate); broader pinnae (sterile 2–4 mm wide (vs. 1.5–2 mm). The species is highly variable with blades ranging from very narrow to broad, with or without lime-dots, scaly to nearly glabrous adaxially, and with scales dark or light at point of attachment. Material from Peru and Bolivia has much wider blade scales, especially adaxially, and may represent a distinct taxon. Material from Guerrero and western Oaxaca is distinct in having a high percentage of spreading, linear-lanceolate scales on the adaxial laminar surfaces (Fig. 241M), e.g., Gro (Calonico Soto 3881, FCME; Clark 7145, NY; Keil 9132, ASU; Mexia 9017, NY; Paxson et al. 17M790, TEX; Rzedowski 22541, 22652, NY). Oax (Diggs 3989, NY; Mickel 4208, 6121, 6982, NY).

24. POLYPODIUM GUTTATUM Polypodium guttatum Maxon, Contr. U.S. Natl. Herb. 17: 575, pl. 40. 1916. Type. Mexico. Coahuila: near Saltillo, Palmer 65 (US!; isotypes GH!–2 sheets, NY!, UC!). Figs. 244A–D. Rhizomes creeping, 2–3 mm diam.; rhizome scales 1–1.5 mm long, bicolorous, each with narrow black center and broad, thin, membranous, ruffled margins, cell lumina not occluded in the dark central streak; fronds 15–25 cm long; stipes 1⁄3–1⁄2 the length

POLYPODIUM HESPERIUM

of the fronds, reddish brown to atropurpureous, glabrous; blades pinnatisect, oblong-deltate, not narrowed at bases, 4.5–7 cm wide; pinnae 12–15 pairs, linear, (3–)4–7 mm wide, not or barely constricted near bases, obtuse to rarely acute at apices, the margins entire to minutely crenulate; distal pinnae slightly ascending; veins not visible; abaxial blade scales (0.5–)1–1.2 mm long, ovatelanceolate, orange-tan to brown; adaxial surfaces glabrescent with hydathodal pits; sori round, medial; sporangia glabrous.

493

attenuate, with an occasional marginal hair; fronds 15–60(–79) cm long, 3–8 mm distant; stipes 1⁄6–1⁄3 the frond length, castaneous, glabrous; rachises with dense hairs 0.3 mm long adaxially, scattered hairs abaxially; blades 1-pinnate, apices pinnatifid; pinnae 22–49 pairs, sessile, acuminate, linear-oblong, 30–50 ⫻ 6–10 mm, only slightly reduced proximally, the basal pinnae at least 1⁄2 as long as the longest pinnae; hairs sparse abaxially, scattered to abundant on adaxial surfaces, 0.1–0.3 mm, and along margins, 0.3–0.5 mm; margins slightly undulate; veins free; sori distinctly elongate-oblong; sporangia with sharp-pointed setae less than 0.1 mm long.

Distribution. Epipetric on steep, rocky mountainsides in oak or oak-pine forests; 1575–3400 m. Mexico. Selected Specimens Examined. Ags (Corres 15303, GUADA). BCS (Brandegee 653, NY). Chih (Correll & Gentry 23049, UC, US). Coah (McPherson 893, NY). Dgo (Tenorio L. 8083, MEXU). Gto (McVaugh 24112, NY). Hgo (Rivera 58, NY). Me´x (Rzedowski 24040, ARIZ, ASC, NY). Mich (Rzedowski 42223, IEB). NL (Hinton et al. 22239, NY). Oax (Ernst 2367, MEXU). Pue (Ventura A. 8357, NY). Qro (Ferna´ndez N. 1550, NY). SLP (Parry & Palmer 973, NY). Tam (Clausen 7401, NY). Ver (Chazaro B. et al. 6096, IEB). Zac (Johnston et al. 11570, CAS). Unverified, Doubtful, or Mistaken Reports. Mor (Cardoso 6, UAMIZ, cited by Riba et al., 1986, but specimen is Polypodium madrense). Sun(reported by Vega et al., 1995, but not verified).

This species is distinct by the obtuse to broadly acute pinnae, large laminar scales, and yellow-green blades.

25. POLYPODIUM HARTWEGIANUM Polypodium hartwegianum Hook. in Bentham, Pl. Hartw. 54. 1840. Type. Mexico. [Hidalgo:] “Monte Sumate” [Zumate], 9500’, Hartweg [415] (K–2 sheets!, frag. & drawing NY!; isotypes BM!, G photo US!, L photos NY! & UC!, P!). Figs. 237D, E. Polypodium ellipsoideum Fe´e, Me´m. Foug. 7: 57, t. 21, f. 1. 1857. Syntypes. Mexico. Popocate´petl, Schaffner 270 (P!); “San Angel,” Schaffner 211 (presumably P, not found; isosyntype K!). Polypodium pubescens Fe´e, Me´m. Foug. 8: 87. 1857, hom. illeg., non L., 1759. P. fournieri C. Chr., Index Filic. 528. 1906. Type. Mexico. Veracruz: Huatusco, a` Dos Puentes, Schaffner 181, 1854 (P?).

Rhizomes creeping, 4–6 mm diam.; rhizome scales widely spreading, dark reddish brown, 4–6 ⫻ 1 mm, margins paler, narrow, tips linear to uniseriate, ovate-lanceolate to lanceolate-

Distribution. Epiphytic, rarely terrestrial or epipetric, in cool, wet pine-oak forests; 1850–3100 m. Mexico; Guat, Hond, Salv. Selected Specimens Examined. BCS (Pray 1773, LAM). Chih (Correll & Gentry 22953, NY). Chis (Breedlove 22758, NY). DF (Pringle 11796, US). Dgo (Reveal 3573, NY). Gro (Hinton et al. 14729, NY). Gto (Zamudio & Murillo 6398, IEB). Hgo (Dunn et al. 23091, NY). Jal (Rzedowski 26164, MICH). Me´x (Rzedowski 26447, NY). Mich (Hinton et al. 15347, NY). Mor (Lyonnet & Chavez 3397, US). Nay (Flores F. 2204, US). Oax (Mickel 4333, NY). Pue (Arse`ne 7065, US). Qro (Rzedowski 45738, IEB). SLP (Schaffner s.n. in 1879, NY). Ver (Ugent & Flores C. 2367, NY). Unverified, Doubtful, or Mistaken Reports. NL (Favela 33, UNL, cited by Arreguı´n-Sa´nchez & Aguirre-Clavera´n, 1985, but not verified).

This is the most common species of Polypodium at higher elevations. It is distinct by the elongate sori and pilose blades. Polypodium liebmannii is similar but densely hairy on the abaxial surfaces, and the fronds are larger (60–115 cm).

26. POLYPODIUM HESPERIUM Polypodium hesperium Maxon, Proc. Biol. Soc. Wash. 13: 200. 1900. Figs. 243A, B. Polypodium vulgare L. var. hesperium (Maxon) A. Nelson & J. F. Macbride, Bot. Gaz. 41: 30. 1916. Type. U.S.A. Washington: Lake Chelan, Coyote Canyon, Gorman 642 (US!). Polypodium prolongilobum Clute, Fern Bull. 18: 97. 1910. Type. U.S.A. Arizona: S side of Mt. Lemmon, Santa Catalina Mts, Oct 1910, Ferris s.n. (isotype US!). Polypodium vulgare L. var. columbianum Gilbert, Working List N. Amer.

494

POLYPODIUM

Pterid. 38. 1901. Polypodium vulgare subsp. columbianum (Gilbert) Hulte´n, Kongl. Svenska Vetenskapsakad. Handl. 8(5): 176. 1962. Syntypes. Canada. British Columbia: Columbia River, Arrow Lake, Hill s.n., Sept 1899, Rocky Mts, near Okanagan Lake, Gilbert s.n. (NYS?)

Rhizomes creeping, 2–3 mm diam., occasionally whitish pruinose; rhizome scales concolorous, orange-tan, deltate-lanceolate, 2.5–3 ⫻ ca. 1 mm, entire to irregularly denticulate near apices; fronds 9–22 cm long, subapproximate; stipes 1⁄3–2⁄5 the frond length, stramineous, glabrous; blades pinnatisect, narrowly lanceolate, 2.2–4.5 cm wide; pinnae 5–10 pairs, 12–22 ⫻ 5–8 mm, oblong, rounded at apices, subcoriaceous; abaxial and adaxial blade surfaces glabrous; venation free but obscure, vein ends evident; sori round, medial; sporangia glabrous; 2n⫽148 (USA). Distribution. Epiphytic in wet forests; 300–900 m. Mexico; Guat, Bel (Hawkins 1078, UC), Hond, Salv, Nic. Specimens Examined. Chis (Breedlove 22436, NY). Oax (Hallberg 1693, NY; Herna´ndez G. 1858, NY; Vera Santos 2625, US). Unverified, Doubtful, or Mistaken Reports. Ver (Riba 1148a, UAMIZ; F. Ramı´rez 514, XAL, both cited by Palacios-Rios, 1998, but not verified).

Distribution. Crevices and cliffs on a variety of noncalcareous substrates, rarely on limestone; 2100–2500 m. Canada (BC), w USA; Mexico. Specimens Examined. BCN (Boyd 2714, UC; Brandegee s.n., 13 May 1893, UC, US; Moran 25612, UC; Moran & Thorne 14319, MICH, UC; Wiggins & Demaree 5000, UC, US). Chih (Correll & Gentry 23218, MSC, US).

This species is distinct by its small size, short pinnae, and glabrous blades.

27. POLYPODIUM HISPIDULUM Polypodium hispidulum Bartlett, Proc. Amer. Acad. Arts 43: 48. 1907. Type. Guatemala. Izabal: Los Amates, Deam 117 (GH!). Figs. 235H, I. Rhizomes creeping, 3–5 mm diam.; rhizome scales tan to brown, tightly appressed, ovate to lanceolate, wrinkled, lustrous, 3–5 ⫻ 0.8–1 mm; fronds 10–23 cm long, distant; stipes ca. 1⁄4 the frond length, stramineous, glabrous; blades pinnatisect, oblong, 5–8 cm wide; pinnae 5–13 pairs, linear-deltate, equally dilated at base, acuminate, with abundant white hairs 0.3–0.5 mm long on rachises, costae, margins, and laminar surfaces; margins shallowly serrulate; veins netted, with one row of areoles between costae and pinna margins; sori round, medial, in one row between costae and margins; sporangia with setae 0.1–0.3 mm long; 2n⫽74 (Nay).

Polypodium hispidulum has the tightly appressed rhizome scales of P. plesiosorum and P. rhodopleuron, but differs from those in the densely hairy blades. In Mexico, P. hispidulum appears to be confined to Oaxaca and Chiapas, whereas P. plesiosorum is more widespread.

28. POLYPODIUM LEPIDOTRICHUM Polypodium lepidotrichum (Fe´e) Maxon, Contr. U.S. Natl. Herb. 17: 591. 1916. Figs. 233A–D. Goniophlebium lepidotrichum Fe´e, Me´m. Foug. 8: 93. 1857. Lectotype (chosen by Maxon, 1916: 591). Mexico. Veracruz: Orizaba, Schaffner 451 (RB).

Rhizomes creeping, 5–7 mm diam.; rhizome scales pale, concolorous except with cell walls near center dark brown, stramineous to lustrous-castaneous, linear to linear-lanceolate, longattenuate, remotely serrulate, 2–2.5(–4) ⫻ 0.5–0.8 mm, often reflexed; fronds 30–65 cm long, distant; stipes 1⁄3–1⁄2 the frond length, stramineous, lanose with pale acicular scales with round basal portion 0.3–0.5 mm wide, centers dark red, tips 2–4 mm long, twisted, white; blades deltate-ovate or obovate, pinnatisect, with terminal pinna conform, 16–23 cm wide; pinnae 10–18 pairs, to 12 ⫻ 0.7–1.5 cm, linear-deltate, broadly adnate, surcurrent to equally surcurrent and decurrent, occasionally slightly constricted at bases, nectaries on surcurrent edges, apices acuminate; costae prominent, stramineous; adaxial laminar surfaces with scattered minute acicular fimbriate scales; abaxial surfaces with more and larger scales, centers red, tips white, 1–1.3 mm long; veins not evident; sori slightly oblong, pushing through scale cover, impressed, adaxial surfaces embossed; sporangia glabrous.

POLYPODIUM LINDENIANUM

495

Distribution. Epiphytic, in montane wet forests of oak and Liquidambar; 300–1850 m. Mexico.

Distribution. Epiphytic in moist oak forests; 1700–2150 m. Mexico.

Selected Specimens Examined. Hgo (Rzedowski 28695, NY). Oax (Mickel 4352b, 7293, NY). Pue (Pe´rez Garcı´a 755, MEXU; Sa´nchez M. 669, US). Ver (Bourgeau 2853, NY; Nee & Taylor 29063, NY; Taylor et al. 131, NY; Ventura A. 2357, 10280, NY).

Selected Specimens Examined. Hgo (Moore 3183, UC, US). Pue (Liebmann s.n., C; Orcutt 3985, NY, US, 3989, DS, NY). Ver (Calzada 9402, IEB; Martı´nez & Va´zquez 476, IEB; Ventura A. 4672, 5604, NY, 11703, CHAPA, ENCB, IEB, 22106, IEB).

Unverified, Doubtful, or Mistaken Reports. Qro (Dı´az-Barriga 6656, IEB, cited by Dı´az-Barriga & Palacios-Rios, 1992, but not verified and doubtful, probably Polypodium rosei).

This species is closely related to P. hartwegianum, differing in blades, stipes, and rachises being very densely hairy on both sides

Among members of the P. squamatum L. complex, P. lepidotrichum is distinct by its long pinnae, strongly reflexed rhizome scales, and the long-tipped, acicular stipe scales.

30. POLYPODIUM LINDENIANUM

29. POLYPODIUM LIEBMANNII Polypodium liebmannii C. Chr., Index Filic. 539. 1906. Figs. 244G, H, J. Polypodium griseum Liebm., Mexic. Bregn. 198 (reprint 46). 1842, hom. illeg., non Schkuhr, 1806. Type. Mexico. Puebla: Chinautla, Liebmann s.n. [Pl. Mex. 2525, Fl. Mex. 125, 715] (C!, frag. US!; isotypes K!, P!).

Rhizomes short-creeping, 4–5 mm diam.; rhizome scales lustrous, brown, entire, linear-lanceolate, ca. (4–)8 ⫻ 0.8–1 mm, tips long-attenuate; fronds 60–120 cm long, ca. 5 mm distant; stipes 1⁄8–1⁄3 the frond length, castaneous, with spreading, lax, tan to whitish hairs 1–1.5 mm long; blades pinnatisect, linearlanceolate, 8–24 cm wide, broadly adnate, contiguous or separated; pinnae 30–65 pairs, 4–11 ⫻ 0.8–1.3 cm, entire; hairs dense, 0.3–0.5 mm on blades and rachises, hairs to 0.8 mm on pinna margins, erect abaxially, appressed adaxially; venation free; sori oblong, medial; sporangia setose.

Polypodium lindenianum Kunze, Farrnkra¨uter 2: 83. 1851. Syntypes. Mexico. Chiapas: Linden s.n., Linden “5539” [probably 1549] (isosyntypes of both collections at P!). Figs. 241C, D. Polypodium cancellatum Fe´e, Me´m. Foug. 5: 242. 1852. Type. “Cuba” [probably incorrect, most likely Chiapas]. Linden s.n. (RB; isotypes P–2 sheets!). Polypodium verapax Christ, Bull. Herb. Boissier, se´r. 2, 5: 253. 1905, nom. nud. Based on Guatemala. Alto Verapaz: entre Coba´n et Paitio, Bernoulli 287, 448; Coba´n, Tuerckheim 4400; Cubilquitz, Tuerckheim 7726 (all presumably at P).

Rhizomes creeping, ca. 3 mm diam.; rhizome scales tan, lanceolate, entire to minutely denticulate, 5–6 ⫻ 1–1.5 mm; fronds 22–70 cm long, 2–5 mm distant; stipes ca. 1⁄3 the frond length, castaneous to atropurpureous, with abundant tan to whitish, concolorous, round to acicular, fimbriate scales, 0.2–0.5 mm, and larger, ovate to ovate-lanceolate, fringed scales 1–2 mm; blades 2–3-pinnate, lanceolate, 4–10 cm wide; pinnae 20–30 pairs, 2.5–7 mm long, veins not visible, ultimate segments 1–2 mm wide, firm; abaxial blade surfaces with scattered, round to acicular, maroon-centered, fimbriate scales, scales most dense over sori; adaxial blade surfaces glabrous except on rachises and costae; rachises green-winged, with wings extending onto distal part of stipes; venation obscure; sori round, medial; sporangia glabrous.

496

POLYPODIUM

Distribution. Epiphytic in montane rain forests, pine-oak forests; 800–2100 m. Mexico; Guat, Bel, Hond, Salv, Nic, CR.

Distribution. Epiphytic in wet montane forests; 1000–3000 m. Mexico; Guat, Hond.

Selected Specimens Examined. Chis (Breedlove 21364, 25714, 34597, NY; Calzada et al. 8746, UC; Ghiesbreght 257, NY; Hammel et al. 15705, UC, US; Heath & Long 23A, CAS; Me´ndez Ton 5335, NY, UAMIZ; Purpus 7227, NY, UC; Sundberg 2374, UC).

Selected Specimens Examined. Chis (Breedlove 15360, NY; Little & Sharp 9945, US). Gro (Rzedowski 18599, ENCB). Hgo (Rzedowski 32681, CHAPA). Jal (McVaugh 14241, US). Oax (Mickel 6597, NY). Pue (Rzedowski 31755, ENCB; Vera Santos 3670, US). Qro (Rubio 2495, IEB). Ver (Ventura A. 4792, NY).

This species is closely related to P. friedrichsthalianum Kunze, from Costa Rica, which differs in being fully tripinnate.

31. POLYPODIUM LONGEPINNULATUM Polypodium longepinnulatum E. Fourn., Mexic. Pl. 1: 77. 1872. Type. Mexico. Oaxaca: Distrito Villa Alta, Galeotti 6510 (P!, photo BM; isotype BR!, photos UC, US!). Figs. 237M, N, 244M, N. Polypodium biauritum Maxon, Contr. U.S. Natl. Herb. 13: 9. 1909. Type. Guatemala. Baja Verapaz: Between Purulha´ and Panzal, Tu¨rckheim s.n. [Donn. Sm. 1688] (US!).

Rhizomes creeping, 3–5 mm diam.; rhizome scales deltatelanceolate, brown, lustrous, 3–4 mm long; fronds 30–70 cm long, distant; stipes 1⁄6–1⁄3 the frond length, light to medium brown, minutely glandular with hairs less than 0.1 mm; rachises with dense hairs 0.1 mm long; blades narrowly to broadly oblong, tapering to pinnatifid apices, (10–)12–26(–30) cm wide, very thin; pinnae linear-lanceolate to linear-deltate, often constricted at bases and with a slight superior auricle or with both superior and inferior auricles at pinna bases; adaxial laminar surfaces with dense hairs 0.3–0.5 mm on costae, a few hairs also on veins and margins; abaxial laminar surfaces less abundantly hairy, costae with hairs 0.1 mm; margins entire to slightly undulate; veins mostly casually anastomosing proximally, especially in larger specimens, distally free; sori slightly oblong, rarely round, medial to supramedial; sporangia glabrous; 2n⫽74 (Chis).

Unverified, Doubtful, or Mistaken Reports. Ags (Siqueiros 4348, HUAA, cited by Siqueiros-Delgado & Gonza´lez-Adame, 2004, but not verified).

Polypodium longepinnulatum is distinguished by the pinnae usually constricted at the bases, each pinna often with small superior auricle or with both a superior and inferior auricle.

32. POLYPODIUM MADRENSE Polypodium madrense J. Sm. in Seemann, Bot. Voy. Herald 338, t. 73. 1856. Type. Mexico. Sierra Madre, Seemann s.n. (BM!, frag. NY!). Figs. 236A–E. Polypodium oulolepis Fe´e, Me´m. Foug. 8: 86. 1857. Syntypes. Mexico. Veracruz and Me´xico: Orizaba & Popocate´petl, Schaffner 191 (K!), 192, 274 p.p. (P!).

Rhizomes creeping, 3–5 mm diam.; rhizome scales strongly bicolorous, spreading, 2.5–3 mm long, each with a black narrow central streak and margins broad, thin, tan to brown, ruffled, erose-denticulate; fronds 18–40 cm long, subdistant; stipes 1⁄2 the frond length, dark castaneous to atropurpureous, with scattered lanceolate scales similar to those of rhizomes; rachises with scattered bicolorous scales with central black streak; blades pinnatisect, oblong to narrowly deltate, not reduced proximally, 4–8(– 9) cm wide; pinnae 7–18 pairs, to 3–4 cm ⫻ 2–3 mm, linear, apices obtuse, crenulate distally; abaxial surfaces with conspicuous round scales 0.5 mm diam., varying to acicular scales 1 mm long, serrulate, pale, each with dark point of attachment; adaxial surfaces glabrescent, with hydathode pits often in double row near pinna bases; veins free, not visible; sori round, medial; sporangia glabrous.

POLYPODIUM MICROGRAMMOIDES

497

adaxial blade surfaces and margins with scattered, lax, white hairs 0.8–1 mm long; abaxial costae and veins with scattered hairs; veins free; sori round, submedial; sporangia with setae 0.5–0.8 mm long.

Distribution. Epiphytic; 1800–3250 m. Mexico. Selected Specimens Examined. Ags (Correa 172, MICH). Chih (Correll & Gentry 23100, LL). Col (Goldsmith 43, US). DF (Rusby 330, NY). Dgo (Keil 8877, NY). Gro (Rzedowski 26301, NY). Gto (Ventura y Lo´pez 9318, IEB). Hgo (Breckon et al. 2323, NY). Jal (McVaugh 25915, NY). Me´x (Mickel 671, NY). Mich (Pringle 3351, NY). Mor (McPherson 962, NY). Oax (Mickel 4062, NY). Pue (Dı´az Luna 4939, UC). Qro (Rzedowski 45739, MEXU). Sin (Gentry 7223, US). SLP (Parry & Palmer 973, NY). Tlax (Herna´ndez X. 430, US). Unverified, Doubtful, or Mistaken Reports. Ver (Nee & Taylor 25896, F, NY; Ventura 10281, ENCB, MO, both cited by PalaciosRios, 1992, but the former is P. plebeium and the latter not verified but probably also P. plebeium).

Polypodium madrense closely resembles P. plebeium but is distinct by its long-attenuate, contorted blade scales (vs. scales small, ovate-lanceolate, and appressed), narrower pinnae, held perpendicular to the rachises (vs. distal pinnae somewhat ascending), and by its occurrence at higher elevations of 1800– 3250 m (vs. (450–)1000–2800 m).

33. POLYPODIUM MARTENSII Polypodium martensii Mett., Abh. Senckenberg. Naturf. Ges. 2: 61. 1857. Figs. 237A–C. Polypodium affine M. Martens & Galeotti, Me´m. Foug. Mexique 37. 1842, hom. illeg., non G. Forst., 1786, nec Blume, 1829. Type. Mexico. Veracruz: Orizaba, Galeotti 6453 (BR!; isotype BR!). Polypodium callolepis Fe´e, Me´m. Foug. 8: 86. 1857. Syntypes. “Mexique, sur le Popocate´petl, pre`s de Mecameca, a` 2700 me`tres,” Schaffner 271 (P!–2 sheets; isotype K!), 272 (presumably P, not found).

Rhizomes creeping, 7–10 mm diam.; rhizome scales pale reddish brown, ovate-lanceolate to lance-attenuate, spreading, 10– 12 ⫻ 3 mm, bases reddish brown, tips pale, fimbriate-denticulate, tangled; fronds (15–)23–42 cm long, distant; stipes 1⁄6–1⁄4 the frond length, brown, with dense minute glandular hairs 0.1 mm long; rachises dark reddish brown, with dense glandular hairs 0.1 mm long and rare, long, jointed hairs 0.8 mm long; blades linearlanceolate to ovate-lanceolate, pectinate, 5–9(–14) cm wide; pinnae linear-oblong to linear-deltate, 5–10 mm wide, proximal pinnae often greatly reduced, bases equally broadly adnate, not strongly surcurrent or decurrent, obtuse to acute; costae dark;

Distribution. Epiphytic in pine-oak woods; 1800–3100 m. Mexico. Selected Specimens Examined. Chih (Spellenberg et al. 12090, UC). DF (Pringle 11798, US). Gto (Rzedowski 44770, IEB). Hgo (Fisher 37101, US). Jal (Rzedowski 27518, NY). Me´x (Rzedowski 21711, NY). Mich (Hinton et al. 13282, NY). Mor (Lyonnet 2477, MEXU, US). NL (Hinton et al. 23268, TEX). Oax (Pringle 4917, GH, LE, NY, US). Pue (Purpus 4042, NY, US). Qro (Rzedowski 47980, IEB). SLP (Palmer 174, NY). Tam (Lof 704, UC). Ver (Nee 23091, NY).

Among the free-veined, hairy-bladed polypodiums, P. martensii is distinguished by its entire margins, large rhizome scales, and long-setose sporangia. Disjunct material from Chihuahua has smaller fronds 7–20 cm long, deltate or deltate-lanceolate blades, pinnae not reduced proximally as in other parts of the range, and pinnae only 3–5 mm wide.

34. POLYPODIUM MICROGRAMMOIDES Polypodium microgrammoides Mickel & A. R. Sm., Brittonia 52: 236. 2000. Type. Mexico. Guanajuato: Mpio. de Ocampo, 4 km al S de Santa Ba´rbara, Carranza et al. 4320 (UC!). Figs. 240E–H. Rhizomes creeping, glaucous, 2–3 mm diam.; rhizome scales concolorous orange-brown, lanceolate, minutely denticulate along attenuate tips, appressed, 4 ⫻ 1.5 mm; fronds 6–13 cm long, distant, coriaceous; stipes 1⁄6–1⁄5 the frond length, castaneous, glabrous; blades simple, narrowly elliptic, entire (but minutely and regularly notched along margins, which are thickened and slightly revolute), 0.7–1 cm wide, strongly coriaceous, bases attenuate, apices acute; abaxial blade surfaces with scattered round to lanceolate, orange to tan scales, each with dark center and pale margins, becoming whitish with age; adaxial blade surfaces glabrous; venation obscure, anastomosing, veins ending near blade margins in conspicuous pits (hydathodes); sori round to slightly oblong, to 3 mm diam., medial, one row on each side of midrib, lacking scales in the sori but with a few larger scales

498

POLYPODIUM

slightly enveloping young sori at their periphery; sporangia glabrous.

ovate-lanceolate, deeply cordate (pseudopeltate), entire to erose; fronds 8–25 cm long, distant, usually 1–3 cm apart; stipes ca. 1⁄2 the frond length, castaneous to darkly castaneous, with sparse, appressed, ovate scales; blades pinnatisect, 4–10 cm wide, with a conform terminal pinna; rachises and costae pale to castaneous, prominent; pinnae 1–2 pairs, 4.5–6.5 cm ⫻ 7–14 mm; adaxial laminar surfaces glabrous; abaxial laminar surfaces with sparse, minute, round to lanceolate scales with reddish centers, margins pale, serrulate, 0.8–1 ⫻ 0.3 mm; blade texture coriaceous; veins netted but not evident; nectaries borne on acroscopic side of pinna bases; sori large, round, medial, soral scales peltate, 0.8 mm wide, centers reddish brown, margins pale brown, erose; sporangia glabrous.

Distribution. On rocks on hillside; 2250–2500 m. Mexico. Known only from the type collection. Polypodium microgrammoides differs from P. muenchii, heretofore the sole member of the segregate genus Microphlebodium, primarily by the simple blades. The distribution of P. muenchii is in southern Mexico (Oaxaca and Chiapas), Guatemala, Honduras, and El Salvador. From Microgramma nitida (J. Sm.) A. R. Sm., which it resembles in the simple blades and creeping rhizomes, P. microgrammoides differs by the glaucous rhizomes beneath the scales (as in P. muenchii); the monomorphic (vs. dimorphic) blades; the broadly ovate, bright orangish brown, concolorous rhizome scales (similar to P. muenchii); longer stipes, to ca. 3 cm (vs. stipes absent or to ca. 0.5 cm); the lack of hair-like scales in the sori; prominent hydathodes adaxially; notched (vs. entire) blade margins; more numerous and ovate (vs. lanceolate to filiform) scales; more obscure venation (veins not at all or only faintly visible in P. microgrammoides, usually readily visible adaxially and sometimes abaxially in M. nitida, and the petioles and midribs at the base of the blades dark purplish brown, exactly as in P. muenchii but unlike M. nitida. Polypodium microgrammoides also grows at much higher elevations than does M. nitida (0–900 m), which has not been found in Guanajuato. Polypodium muenchii also has notched pinna margins and prominent hydathodes, as in P. microgrammoides, and the two species are closely allied.

35. POLYPODIUM MUENCHII Polypodium muenchii Christ, Bull. Herb. Boissier, se´r. 2, 3: 147. 1903 [“munchii”]. Figs. 231E–H. Pleopeltis muenchii (Christ) A. R. Sm., Proc. Calif. Acad. Sci., ser. 4, 40: 230. 1975. Microgramma muenchii (Christ) Copel., Amer. Fern J. 43: 15, t. 2. 1953. Microphlebodium muenchii (Christ) L. D. Go´mez, Phytologia 59: 58. 1985. Type. Mexico. Chiapas: San Cristo´bal Las Casas, Mu¨nch 90 (P!; isotype DS, frag. US).

Rhizomes long-creeping, 2.5–3(–4) mm diam., glaucous; rhizome scales 4–6 ⫻ 1.5–2 mm, lustrous golden to reddish brown,

Distribution. Epiphytic in cloud forests at higher elevations; 2400–3000 m. Mexico; Guat, Hond, Salv. Selected Specimens Examined. Chis (Breedlove 6692, 7811, US; Carlson 1680, US; Laughlin 162, US; Little & Sharp 9948, US; Mickel 1229, US). Oax (Campos V. 882, NY; Mickel 1141, 4330, NY; Torres C. 10916, NY).

Polypodium muenchii is distinct in its pinnatifid blades, lustrous brown rhizome scales, and occurrence at high elevations. Young sori with roundish scales caused Smith (1981) and Mickel and Beitel (1988) to place it in Pleopeltis, but the soral scales in fact are not peltate but cordate-based. The distinctness of this species caused Go´mez (1985) to erect the genus Microphlebodium. Recently, a simple-bladed species, P. microgrammoides (which see), was discovered in this group.

36. POLYPODIUM PLATYLEPIS Polypodium platylepis Mett. ex Kuhn, Linnaea 36: 137. 1869. Lectotype (chosen by Maxon, Contr. U.S. Natl. Herb. 17: 563. 1916). Guatemala. Skinner s.n. (K!, frag. NY!). Figs 234E–H, J. Rhizomes creeping, 2–3 mm diam.; rhizome scales tan, darksclerotic at point of attachment, lanceolate, minutely denticulate, 5–6 mm long; fronds 16–37 cm long, distant; stipes ca. 1⁄2 the frond length, castaneous to atropurpureous, with spreading, ovate scales, and appressed, round scales, both types denticulate

POLYPODIUM PLESIOSORUM

to erose; blades pinnatisect, narrowly deltate, 3.5–7 cm wide; pinnae 10–25 pairs, 2–7 mm wide, margins serrulate; scales on abaxial surfaces round (0.8 mm diam.) to ovate-lanceolate, longacuminate (2 mm long), denticulate; adaxial surface glabrescent, lime-dots present; nectaries borne on acroscopic bases of pinnae; sori oblong, 1–1.5 mm long; sporangia glabrous.

Distribution. Epiphytic in oak forests, pine-oak forests; 1400– 2750(–3325) m. Mexico; Guat; Ven. Selected Specimens Examined. Chis (Breedlove 27695, NY). DF (Lyonnet 470700001, MEXU). Gro (Rzedowski 18525, NY). Jal (McVaugh 26021, NY). Me´x (Hinton 7474, US; Rzedowski 33649, NY). Mich (Hinton et al. 13283, NY). Mor (Lyonnet 511000002, MEXU). Oax (Mickel 4540, NY). Qro (Carranza G. 1390, IEB). Unverified, Doubtful, or Mistaken Reports. Ver (reported by Smith, 1981, but not verified and doubtful).

Polypodium platylepis resembles P. furfuraceum and P. cryptocarpon in the scaly blades and pale, concolorous rhizome scales, but is distinct in its broader pinnae, narrowly deltate blades, longer stipes, glabrous adaxial blade surfaces, and oblong sori.

37. POLYPODIUM PLEBEIUM Polypodium plebeium Schltdl. & Cham., Linnaea 5: 607. 1830. Type. Mexico. [Veracruz:] Jalapa, Schiede & Deppe [746] (B!; isotypes LE!–2 sheets). Figs. 236F–H. Polypodium cheilostictum Fe´e, Me´m. Foug. 8: 87. 1857. Type. Mexico. [Veracruz:] Orizaba, Schaffner 453 (RB). Marginaria karwinskyana C. Presl, Suppl. Tent. Pterid. 188. 1836. Polypodium karwinskyanum (C. Presl) A. Braun ex Kunze, Linnaea 23: 279, 319. 1850. Type. Mexico. Santa Rosa, Jun 1803, Karwinski s.n. (B!).

Rhizomes creeping, 3–5 mm diam.; rhizome scales bicolorous, each with narrow, black center, and broad, thin, membranous, ruffled margins, 2–3 mm long, lumina occluded in central streak; fronds 12–35 cm long, distant; stipes 1⁄4 to nearly 1⁄2 (rarely over 1 ⁄2) the frond length, reddish brown to atropurpureous, nearly glabrous but with scattered, appressed scales with dark central streak and pale, thin margins; blades pinnate to pectinate, oblong-deltate, not reduced proximally, 5–10 cm wide, distal pinnae ascending; pinnae 10–17 pairs, linear to linear-deltate (to linear-lanceolate), 3–5 mm wide, not or barely constricted near

499

bases, apices acuminate to broadly acute; scales minute, brown, ca. 0.5 mm long, sparse, inconspicuous; adaxial surfaces glabrescent with hydathodal pits; margins entire to crenulate, modified, clear; veins not visible; sori round, medial; sporangia glabrous; 2n⫽74 (Oax).

Distribution. Epiphytic and terrestrial; 600–3050 m. Mexico; Guat, Hond, Salv, Nic, CR, Pan. Selected Specimens Examined. Chis (Breedlove 21148, NY). Gto (Ventura V. & Lo´pez P. 9571, IEB). Hgo (Rzedowski 27652, NY). Me´x (Rzedowski 24040, LL). Oax (Mickel 7144, NY). Pue (Rzedowski 23391, NY). Qro (Rubio 2085, MEXU). SLP (Pringle 3258, NY). Tam (von Rozynski 713, NY). Ver (Diggs & Nee 2917, NY). Unverified, Doubtful, or Mistaken Reports. Gro (Lorea 1507, FCME, cited by Lorea-Herna´ndez & Vela´zquez M., 1998, but not verified). Mor (cited by Riba et al., 1986, but specimens are P. madrense). Tab (reported by Magan˜a, 1992, but figured plant is Polypodium polypodioides).

Polypodium plebeium and its close relatives have blackcentered rhizome scales with tan, well differentiated, membranous marginal wings, but P. plebeium is distinct by the rhizome scales spreading with crisped margins and ascending distal pinnae.

38. POLYPODIUM PLESIOSORUM Polypodium plesiosorum Kunze, Linnaea 18: 313. 1844 [1845]. Figs. 238G–J, 245F–L. Goniophlebium plesiosorum (Kunze) Fe´e, Me´m. Foug. 9: 24. 1857. Polypodium loriceum L. var. plesiosorum (Kunze) Christ, Bull. Soc. Roy. Bot. Belgique 35(1): 221. 1896. Syntypes. Mexico. Herb. Roemer, Leibold 90, 91 (LZ, destroyed; isotypes P!–2 sheets). Polypodium confluens Liebm., Mexic. Bregn. 191 (reprint 39). 1849, hom. illeg., non R. Br., 1810. Lectotype (chosen by Smith, 1981: 189). Mexico. Puebla: Chinautla, Liebmann s.n. [Pl. Mex. 2716, Fl. Mex. 696] (C!, frag. US!; isolectotypes BM!, K!). Goniophlebium molestum Fe´e, Me´m. Foug. 8: 92. 1857. Polypodium anisomeron Fe´e var. δ molestum (Fe´e) E. Fourn., Mexic. Pl. 1: 81. 1872. Type. Mexico Veracruz: Near Huatusco, Cordoba, and Orizaba, Schaffner 180 (P!). Goniophlebium anisomeron Fe´e, Me´m. Foug. 8: 93. 1857. Polypodium anisomeron (Fe´e) E. Fourn., Mexic. Pl. 1: 81. 1872. Syntypes. Mexico. [Veracruz:] Orizaba, Schaffner 182 (P!; isosyntype K!), 458 (presumably P, not found). Gonophlebium calaguala Fe´e, Me´m. Foug. 8: 93. 1857. Type. Mexico. [Distrito Federal/ Edo. Me´xico:] Valley of Mexico, Schaffner s.n. (RB; isotype P!).

500

POLYPODIUM

Polypodium anisomeron (Fe´e) E. Fourn. var. γ elongatum E. Fourn., Mexic. Pl. 1: 81. 1872. Syntypes. Mexico. Veracruz: Ixhuatlancillo, Bourgeau 3061 (GH!), 3061bis (P!), Orizaba, Schaffner s.n. (P!; isosyntype K!). Polypodium anisomeron (Fe´e) E. Fourn. var. α genuinum E. Fourn., Mexic. Pl. 1: 81. 1872. Type. Mexico. Bourgeau 3062, (P!; isotype GH!). Polypodium anisomeron (Fe´e) E. Fourn. var. β pectinatum E. Fourn., Mexic. Pl. 1: 81. 1872. Type. Mexico. Veracruz: Cordoba valley, Bourgeau 1943 (GH!), 1943ter (P!; isotypes B!, BR, photo US!, K!). Polypodium anisomeron (Fe´e) E. Fourn. var. ε abortivum E. Fourn., Mexic. Pl. 1: 82. 1872. Type. Mexico. Veracruz: Orizaba, region of Orizaba, Bourgeau s.n. (P!). Polypodium feei E. Fourn., Mexic. Pl. 1: 82. 1872, hom. illeg., non (Bory) Mett., 1857. Syntypes. Mexico. Oaxaca: Cordillera, Galeotti 6412 (P!, frags. NY!, US!); Veracruz: Orizaba, Botteri & Sumichrast 1422 (MPU). Polypodium van-heurckii E. Fourn., Mexic. Pl. 1: 82. 1872. Syntypes. Mexico. Veracruz: region of Orizaba ca. Escamela, Botteri 1471 (P!); Veracruz: Orizaba, San Andre´s 1473 (P). Polypodium pachyrhizon Christ, Notul. Syst. (Paris) 1: 231. 1910. Lectotype (chosen by Smith, 1981: 190). Mexico. Michoaca´n: Barranca NW of Punguato, Arse`ne 2588 (P!). Polypodium rossii Christ ex H. Ross, Mem. Acad. Nac. Ci. “Antonio Alzate” 32: 191. 1912. Type. Mexico. Morelos: Cuernavaca, Santa Marı´a, Ross Fl. Mex. 281 (M, photo BM!). Goniophlebium stramineum Underw. ex Conz., Fl. Tax. Mex. 1: 105. 1939, nom. nud. Based on. Mexico. Puebla: Estacio´n Honey, Pringle 8980 (MEXU; isotypes CAS!, GH, US).

Rhizomes long-creeping, 3–5 mm diam.; rhizome scales brown, ovate-lanceolate, entire, 3–5 mm long, tightly appressed, somewhat caducous and partially exposing green rhizomes; fronds (10–) 15–65 cm long, distant; stipes 1⁄5–1⁄2 the frond length, stramineous to light brown, occasionally dark brown or atropurpureous, glabrous; rachises with dense hairs 0.1–0.3 mm long, at least adaxially; blades (narrowly oblong to) ovate-lanceolate, pinnatisect, 5–14(–19) cm wide; pinnae 10–29(–38) pairs, linear-deltate, 5–12 mm wide, equally dilated to slightly more surcurrent at bases, apices acuminate to acute or rounded, margins subentire to serrulate; abaxial surfaces with rare hairs 0.1–0.3 mm long only on costae, to densely hairy, hairs 0.1–0.3 mm on costae and veins; adaxial surfaces glabrous except for dense hairs 0.1–0.3 mm long on costae; veins netted (goniophlebioid), often casually so; sori round, medial, in one row each side of costa; sporangia glabrous; 2n⫽74 (SLP).

Distribution. Epiphytic, terrestrial, and epipetric in wet montane forests; 600–2800 m. Mexico; Guat, Hond, Salv, Nic, CR, Pan.

Selected Specimens Examined. Chis (Breedlove 28183, NY). Coah (Lyonnet 421000002, MEXU). DF (Schaffner 70, NY). Gro (Lorea 2268, US; Rzedowski & McVaugh 176, NY). Gto (Rzedowski 45457, IEB). Hgo (Dunn & Dunn 19030, NY). Jal (Rzedowski 27498, NY). Me´x (Mickel 678, NY). Mich (Pringle 3354, NY). Mor (Lyonnet 2426, MEXU, US). Nay (McVaugh 16353, NY). NL (Palmer 1374, NY). Oax (Mickel 4473, NY). Pue (Ventura A. 220, NY). Qro (Ferna´ndez N. 3588, NY). SLP (Rzedowski 24461, NY). Tam (Stanford et al. 1066, NY). Ver (Avendano R. et al. 00044, NY). Unverified, Doubtful, or Mistaken Reports. Sin (reported by Vega et al., 1995, but probably P. praetermissum).

Polypodium plesiosorum is similar to P. rhodopleuron, but the latter has glabrous rachises and pinnae, and reddish stipes and veins. The abaxial surfaces in P. plesiosorum vary greatly from having only rare hairs to being densely hairy. Polypodium plesiosorum and its close relatives (P. rhodopleuron and P. colpodes) are distinct by their greenish rhizomes with tightly appressed scales. There is great variation in plant size and pinna apices. One specimen, for example, U. C. Bot. Gard. 57.420–1, Kennedy s.n. (UC), from Veracruz, has fronds 47–53 ⫻ 17–23 cm.

39. POLYPODIUM PLEUROSORUM Polypodium pleurosorum Kunze ex Mett., Abh. Senckenberg. Naturf. Ges. 2: 101, t. 2, f. 6, 7. 1856. Type. Mexico. [Chiapas:] “entre St.Bartolo et Pueblo Nuevo,” Linden 1518 (LZ destroyed; isotypes K!, frag. & drawing NY!, P!–2 sheets. Figs. 239A–C. Phlebodium inaequale T. Moore, Gard. Chron. 1855: 660, t. 58, f. 4. 1855. Goniophlebium inaequale (T. Moore) J. Sm., Cult. ferns 3. 1857. Polypodium inaequale (T. Moore) Lowe, Ferns 2: t. 28. 1858, hom. illeg., non Link, 1833. Polypodium lowei C. Chr., Index Filic. 541. 1906. Type. Cultivated specimen, said to have been gathered in Guatemala by Loddiges [prob. collected by Skinner] (K!). Polypodium guatemalense Hook., Sp. Fil. 5: 29. 1863, hom. illeg., non Klotzsch, 1855. Type. Guatemala. Skinner s.n. (K!, frag. & drawing NY!).

Rhizomes creeping, 5–10 mm diam.; rhizome scales reddish brown with paler margins, lustrous with denticulate to fimbriate margins when young, erose to entire when old, spreading, ovatelanceolate, peltate, 8–10 ⫻ 2.5–3.5 mm; fronds 25–75 cm long, distant; stipes 1⁄3–1⁄2 the frond length, reddish brown, glabrous; blades 1-pinnate with terminal pinna conform, glabrous except for scattered, brown trichomidia 0.1–0.3 mm long, broadly oblong, 15–50 cm wide; pinnae 2–11 pairs, narrowly oblong, sessile, 7.5–25 ⫻ 1.2–2.5 cm, bases cuneate, apices long-attenuate, proximal pinnae strongly excavate basiscopically; margins with short teeth 0.1 mm long; veins netted with only an occasional included veinlet; sori in one row on each side of costae, round, over 2 mm diam., nearly costal; sporangia glabrous.

POLYPODIUM POLYPODIOIDES

501

Acrostichum polypodioides L., Sp. Pl. 2: 1068. 1753. Polypodium incanum Sw., Prodr. 131. 1788, hom. illeg. Marginaria incana (Sw.) C. Presl, Suppl. Tent. Pterid. 188. 1836. Goniophlebium incanum (Sw.) J. Sm., J. Bot. (Hooker) 4: 56. 1841. Marginaria polypodioides (L.) Tidestr., Torreya 5: 171. 1905. Pleopeltis polypodioides (L.) E. G. Andrews & Windham, Contr. Univ. Michigan Herb. 19: 46. 1993. Lectotype (chosen by Weatherby, Contr. Gray Herb. 124: 28. 1939). Jamaica. s. coll. s.n. (Herb. Plukenet, BM).

Distribution. Epiphytic in wet montane forests; 1800–3000 m. Mexico; Guat, Hond, Salv, Nic, CR. Selected Specimens Examined. Chis (Breedlove 22748, 29421, NY). Gro (Hinton et al. 14232, NY; Holmes 4445, NY; Rzedowski 18598, US). Oax (Calzada 19827, NY; Hellwig 234, NY; Mickel 4901, 6598, NY; Wendt et al. 4642, NY). Unverified, Doubtful, or Mistaken Reports. Mich (Dı´az-Barriga 1403, 1684, IEB, both cited by Dı´az-Barriga & Palacios-Rios, 1992, but not verified).

Polypodium pleurosorum is distinct from P. triseriale by its single, nearly costal row of large sori.

40. POLYPODIUM POLYPODIOIDES Polypodium polypodioides (L.) Watt, Canad. Naturalist & Quart. J. Sci., ser. 2, 3: 158. 1867.

Rhizomes long-creeping, 1–2 mm diam.; rhizome scales imbricate, linear-lanceolate, 3–4 mm long, denticulate-ciliolate, strongly bicolorous with black centers and pale denticulate margins; fronds 5–25 cm long, distant; stipes less than half the frond length, with scales appressed, linear-lanceolate, peltate; blades oblong-deltate to linear-oblong, pinnatifid, truncate proximally, 1.5–6 cm wide; pinnae 6–13(–16) pairs, obtuse, bases dilated, coriaceous, curling on drying; abaxial surfaces with dense, round to ovate-lanceolate, peltate scales with dark centers and pale erosedenticulate margins; adaxial surfaces with few scales with strongly fimbriate margins; veins obscure; sori supramedial, partly embossed; sporangia glabrous. Distribution. Epiphytic and epipetric in mesic woods; 0–2400 m. Se USA; Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Bah, Gr & L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol, Parag, Uru, Arg. Weatherby (1939) recognized five varieties, of which three occur in Mexico; in addition, a new variety is described here. Polypodium polypodioides is distinct from small specimens of P. thyssanolepis by the narrow rhizomes and blades, more and narrower pinnae, and absence of hydathodes.

Key to the Mexican Varieties of Polypodium polypodioides 1. Scales on abaxial laminar surfaces fimbriate-serrulate, non-acicular; adaxial surfaces glabrous (or rarely with a few scales); southern Mexico. ................................................................................................................................................................................................... 40a. var. polypodioides. 1. Scales on abaxial laminar surfaces entire to erose or minutely serrulate, acicular or not; adaxial surfaces glabrous to sparsely scaly. 2. Adaxial laminar surfaces glabrous; scales of abaxial laminar surfaces lacking long, acicular tips; northern Mexico and Chis. ........................................................................................................................................................................................................ 40d. var. michauxianum.

2. Adaxial laminar surfaces with sparse to scattered scales; some scales of abaxial laminar surfaces, at least near margins, with long, acicular tips. 3. Scales of the abaxial laminar surfaces a mixture of round, lanceolate, and acicular scales, the last concentrated near pinna margins; widespread in Mexico except for the northern tier of states and the Yucatan Peninsula. .................................... 40b. var. aciculare. 3. Scales of the abaxial laminar surfaces all acicular; Sin, Son, Chih. ................................................................................ 40c. var. knoblochianum.

502

POLYPODIUM

40a. Polypodium polypodioides var. polypodioides Figs. 231S, T. Distinct by the abaxial laminar scales being fimbriateserrulate, not acicular, and by the adaxial laminar surfaces being glabrous (or rarely with a few scales).

Distribution. Epiphytic and epipetric in mesic pine-oak woods at middle elevations; 500–2400 m. Mexico; Guat, Hond, Salv, Nic, CR, Pan.

Distribution. Rare epiphyte at low elevations; 50–300 m. Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Bah, Gr & L Ant; Col, Ven, Trin. Selected Specimens Examined. Chis (Gilly & Herna´ndez X. 185, US). Oax (Hallberg 1481, NY; Martı´nez-C. 792, US; Mickel 5127, 5791, 5813, 6851, 7218, NY). Tab (Zamudio 476, MEXU). Yuc (Gaumer 1110, NY). Unverified, Doubtful, or Mistaken Reports. Camp (Palacios-Rios et al. 3606, XAL, cited by Palacios-Rios, 2002a, but not verified). QR (Castillo R. & Tun G. 239, UADY, XAL cited by Palacios-Rios, 2002c, but not verified).

This variety is distinguished by its fimbriate-serrulate, nonacicular abaxial laminar scales and its occurrence at low elevations (0–300 m). Breedlove 22432, DS, from Chiapas, combines the characters of three varieties: lamina glabrous above as in var. michauxianum, abaxial laminar scales fimbriate as in var. polypodioides, and with some abaxial laminar scales lanceolate at elevation of 700 m as in var. aciculare; 2n⫽148 (Jam, Trin). Some specimens from Oaxaca and Chiapas appear intermediate with var. aciculare.

40b. Polypodium polypodioides var. aciculare Weath., Contr. Gray Herb. 124: 33. 1939. Figs. 231J–L, U, V. Pleopeltis polypodioides (L.) Watt var. acicularis (Weath.) E. G. Andrews & Windham, Contr. Univ. Michigan Herb., 19: 46. 1993. Type. Costa Rica. San Francisco de Guadalupe, Tonduz 8476 (GH!; isotypes NY!, US!).

Distinct by the abaxial laminar surfaces having a mixture of round, lanceolate, and acicular scales, the last concentrated near pinna margins, and by the adaxial laminar surfaces with sparse to scattered scales.

Selected Specimens Examined. Chis (Breedlove 27721, NY). Col (Espero´n 18292, GUADA). DF (Rusby 332, NY). Dgo (Bashor 694, NY). Gro (Hinton et al. 9435, NY). Gto (Rzedowski 40576, IEB). Hgo (Dunn & Dunn 19025, NY). Jal (McVaugh 16198, MICH). Me´x (Hinton et al. 4162, NY). Mich (Gonza´lez Quintero 1683, NY). Mor (Lyonnet 772, MEXU, NY). Nay (Solorzano 16912, GUADA). Oax (Mickel 3916, NY). Pue (Orcutt 4023, US). Qro (Ferna´ndez N. 3743, NY). Rev (Barkelew 236, NY, US). Sin (Breedlove 18184, NY—intermediate with var. knoblochianum). SLP (King 4439, NY). Ver (Purpus 5799, NY). Zac (Rzedowski & McVaugh 936, NY—intermediate with var. knoblochianum). Unverified, Doubtful, or Mistaken Reports. Ags (Siqueiros 4339, HUAA, and three other collections, cited by Siqueiros-Delgado & Gonza´lez-Adame, 2004 [without variety but presumed to be var. aciculare], but not verified).

This is the most widespread variety of P. polypodioides in Mexico, the other three varieties being rather limited in distribution: var. michauxianum in northeastern states and Chiapas, var. polypodioides in southeastern states, and var. knoblochianum in northwestern states. Var. aciculare has the adaxial blade surfaces sparsely scaly, abaxial surfaces at least somewhat scaly, especially along the margins, and the scales with small dilated bases and acicular tips. Var. michauxianum has the adaxial surfaces glabrous, the abaxial surfaces with scales round to deltate-ovate, obtuse or acute, none with acicular tips; it is diploid, 2n⫽74 (SLP). Var. aciculare can be distinguished by the mixture of acicular scales on the abaxial laminar surface, although the percentage of acicular scales is rather variable, sometimes only at the margins, sometimes with a few near the pinna center, and occasionally with some scattered, but never wholly acicular as in var. knoblochianum. Most specimens of P. polypodioides from Sinaloa, Durango, and Zacatecas are intermediate between vars. knoblochianum and aciculare. 40c. Polypodium polypodioides var. knoblochianum Mickel, var. nov. Type. Mexico. Sonora: Sierra del Nacori, Hartman 341 (holotype NY!; isotypes PH!, UC!). Figs. 231M–R.

POLYPODIUM PRAETERMISSUM Laminis abaxialiter paleis elongatis acicularibus densis obsita distincta. (Honoring the late Irving W. Knobloch, scholar of Chihuahuan ferns.)

Distinct by the abaxial laminar surfaces having only acicular scales, and by the adaxial laminar surfaces with sparse to scattered scales.

503

Guat, Hond, Nic, CR, Pan; Moran (in Davidse et al., 1995) reported it from Bahamas, Antilles, Colombia, Venezuela, and Trinidad. Specimens Examined. Chis (Breedlove 22432, DS, 38192, DS). Coah (Wind & Muller 368, NY). NL (Muller 2750, NY). SLP (Mickel 525, NY). Tam (King 4538, NY).

Var. michauxianum has an unusual distribution pattern, occurring primarily in the northern states but apparently also in Chiapas, Oaxaca, and Guatemala. It is possible the characters have been misinterpreted. More work is needed on this complex. Some collections from Veracruz and San Luis Potosı´ are intermediate between var. michauxianum and var. aciculare, lacking adaxial scales, but with acicular or at least some lanceolate scales below.

41. POLYPODIUM PRAETERMISSUM Distribution. On rocks. 1500–1700 m. Mexico. Specimens Examined. Chih (Knobloch 102, 115, 148, MSC, 5977, DS; Townsend & Barber 387, NY). Sin (Breedlove 17112, NY—intermediate with aciculare). Son (Lloyd 479, PH; Pennell 19651, CAS, 19669, PH; Phillips 515, MICH).

Var. knoblochianum is limited in its distribution to northwestern Mexico, being found primarily in Sonora and Chihuahua. No other varieties of P. polypodioides have been seen from those states. 40d. Polypodium polypodioides var. michauxianum Weath., Contr. Gray Herb. 124: 31. 1939. Figs. 231W, X. Pleopeltis polypodioides (L.) Watt var. michauxiana (Weath.) E. G. Andrews & Windham, Contr. Univ. Michigan Herb. 19: 46. 1993. Type. U.S.A. Virginia: Nansemond Co., Kilby, Fernald, Long & Fogg 4703 (GH).

Polypodium praetermissum Mickel & Tejero, Brittonia 56: 117. 2004. Type. Mexico. Nayarit: New lava flows of Volca´n Ceboruco, 6.5 mi NW of Ahuacatla´n, Mickel 1371 (NY). Figs. 240J–L. Rhizomes creeping, ca. 5 mm diam.; rhizome scales orange-tan, concolorous to somewhat paler at margin, ovate-lanceolate, comose, 2.5–3.5 mm long, margins denticulate; fronds 21–45 cm long, distant; stipes 1⁄3 the frond length, stramineous to castaneous, glabrous;blades pinnatisect, deltate to oblong, 6–12(–18) cm wide, apices pinnatifid or with a subconform terminal pinnae; pinnae (8–)16–28 pairs, linear-deltate, adnate to slightly narrowed at bases, attenuate, glabrous; veins partially and irregularly anastomosing; sori round, median in one row on each side of costae; sporangia glabrous.

Distinct by the abaxial laminar scales lacking long, acicular tips, and by the adaxial laminar surfaces being glabrous.

Distribution. On rocks, rarely epiphytic, wet north-facing cliffs and rock faces on lava flows and steep arroyos in pineoak forests; 1100–2150 m. Mexico.

Distribution. Epiphytic and epipetric in pine-oak and montane rain forests; 700–1700 m (150 in SLP). Se USA; Mexico;

Specimens Examined. Dgo (Reeder & Reeder 2473, MEXU, US). Jal (Correll 14362, US; Palmer 374, NY, US). Nay (Breedlove 4550, CAS, MEXU; Feddema 462, NY; Gonza´lez Ortega 4291, MEXU; Mickel 1370, NY). Sin (Breedlove 18573, NY, 44963, CAS; Pennell 20137, PH). Son

504

POLYPODIUM

(Fishbein 1934, ARIZ, 1937, ARIZ; Pennell 19657, PH; Van Devender 981025, NY).

The laminar surfaces generally have short hairs on the adaxial costae and rachises, abaxially scattered on laminae and axes; Sonoran specimens are essentially glabrous. Plants from Jalisco and Nayarit have 10–20 pairs of pinnae, whereas those from Sinaloa and Sonora have 8–10 pairs. Polypodium praetermissum is closest to P. plesiosorum in the casually netted veins, broadly adnate pinnae, and short hairs on the adaxial rachises, but differs in less appressed, comose rhizome scales. Polypodium praetermissum also closely resembles P. subpetiolatum, which can be distinguished by its abundant laminar hairs, toothed pinna margins, and free veins. Polypodium praetermissum is similar to P. fraternum in its glabrous (or subglabrous) laminae and the irregularly anastomosing veins, relatively few pinna pairs, attenuate pinna apices, and untoothed pinna margins. It differs in the more broadly adnate pinnae, thinner texture, rhizome scales being shorter and comose, presence of some stubby hairs on the costae and laminae, and more northwestern distribution.

42. POLYPODIUM PUBERULUM Polypodium puberulum Schltdl. & Cham., Linnaea 5: 607. 1830, non Baker, 1867. Type. Presumably Mexico. [Veracruz:] Schiede & Deppe [747] (isotype B, photo BM). Figs. 235J, K, 237H, J. Polypodium ehrenbergianum Klotzsch, Linnaea 20: 381.1847. Type. Mexico. Ehrenberg s.n. (B!; isotype S, photo US!). Polypodium olivaceum Liebm., Mexic. Bregn. 199 (reprint 47). 1849. Lectotype (chosen by Smith, 1981: 192). Fl. Mex. 128 (C!) Mexico. Veracruz: “Inter Huatusco et Orizaba,” Liebmann s.n. [Pl. Mex. 2536, Fl. Mex. 128] (C!, frags. NY!, US!). Polypodium olivaceum Liebm.var. elatum Liebm., Mexic. Bregn. 199 (reprint 47). 1849. Type. Mexico. Veracruz: “Hac. de Mirador,” Liebmann s.n. [Pl. Mex. 2537, Fl. Mex. 126] (C!, frag. US!). Polypodium pubescens Fe´e, Me´m. Foug. 8: 87. 1857, hom. illeg., non L., 1759. Polypodium fournieri C. Chr., Index Filic. 528. 1906. Type. Mexico. Veracruz: Huatusco a` Dos Puentes, Schaffner 181, in 1854 (RB). Polypodium galeottii Mett. ex Kuhn, Linnaea 36: 135. 1869. Type. Mexico. Galeotti 6552 (B). Polypodium glaberulum Mickel & Beitel, Mem. New York Bot. Gard. 46: 300. 1988. Type. Mexico. Oaxaca: Distrito Ixtla´n, 2 km E of Ixtla´n, 1–2 km N of Vivero Rancho Teja, Mickel 4982 (NY!).

Rhizomes creeping, 5–7 mm diam.; rhizome scales deltate, long-attenuate, 4–5 mm long, tan to dark lustrous brown, often sclerotic-centered, with abundant minute hairs on outer surfaces; fronds 30–60(–96) cm long, distant; stipes 1⁄4–1⁄2 the frond length, stramineous to light brown, with dense, small, white hairs 0.3– 0.5 mm long; blades pinnate, 11–18 cm wide, basal pinnae reflexed, excavate basiscopically; pinnae linear-deltate, 1.2–2 cm wide, broadly adnate, often surcurrent, rarely any free rachis, acuminate, margins irregularly coarsely serrulate; adaxial laminar surfaces, margins and veins with abundant white hairs 0.3–0.5 mm long; abaxial surface hairs 0.3–0.8 mm long on rachises,

costae, and laminar surfaces; veins free; sori round, medial; sporangia with setae 0.3–0.5 mm long; 2n⫽74 (Ver).

Distribution. Epiphytic in wet montane forests; 1600–2500 m. Mexico; Salv. Selected Specimens Examined. Chis (Breedlove 9411, 33516, 34803, DS). Oax (Galeotti 6552, BR; Mickel 4309, 6599, NY). Pue (Riba et al. 281, NY). Ver (Ventura A. 358, NY). Unverified, Doubtful, or Mistaken Reports. Qro (Gonza´lez P. 1133, IEB; Rubio 1134, IEB, both cited by Dı´az-Barriga and PalaciosRios, 1992, but not verified and doubtful).

The free-veined Polypodiums are in need of more work. Polypodium puberulum resembles specimens of P. echinolepis but is distinct by its broadly adnate pinnae.

43. POLYPODIUM PYRRHOLEPIS Polypodium pyrrholepis (Fe´e) Maxon, Contr. U.S. Natl. Herb. 17: 593. 1916. Figs. 233E–H. Goniophlebium pyrrholepis Fe´e, Me´m. Foug. 8: 94. 1857. Type. Mexico. [Veracruz:] Huatusco, Schaffner 197 (P?, not found).

Rhizomes creeping, 5–7 mm diam.; rhizome scales widely spreading, 3–4 mm long, bicolorous with black bases and maroon, long-attenuate, ciliate-serrulate tips, the teeth often bifid; fronds 25–50 cm long, distant, 1–2 cm apart; stipes 1⁄4–1⁄2 the frond length, castaneous, with long, acicular scales with somewhat twisted tips; blades pinnatisect, linear-lanceolate, 3.5–7 cm wide with very small terminal pinna; pinnae 22–40 pairs, linearoblong, to 2.5–3.5 cm long, bases broadly adnate, surcurrent, 4–6 mm wide, apices obtuse; nectaries on acroscopic surcurrent pinna bases; costae prominent, dark, at least in proximal half; adaxial surfaces with scattered acicular, fimbriate, white scales, with light brown centers; abaxial surfaces with dense scales 2–2.3 mm long, these with light brown centers, whitish margins and light brown tips; sori round, emerging through scales; sporangia glabrous; 2n⫽74 (Oax, Ver).

POLYPODIUM RHACHIPTERYGIUM

Distribution. Epiphytic in wet montane forests; 130–1500 m. Mexico. Selected Specimens Examined. Chis (Breedlove 25139, 34569, NY). Oax (Herna´ndez G. 356, NY; Mickel 6464, NY; Wendt et al. 3625, NY). Tab (Magan˜a 2122, ENCB). Ver (Bourgeau 1446, NY; Purpus 2166, NY; Wendt et al. 3612, 3968, NY). Unverified, Doubtful, or Mistaken Reports. Mich (Dı´az-Barriga 1338, IEB, XAL, cited by Dı´az-Barriga & Palacios-Rios, 1992, but not verified and doubtful, probably Polypodium rosei). Mor (Pe´rez Garcı´a 794, UAMIZ, cited by Riba et al., 1996, but specimen is Polypodium rosei).

Within the P. squamatum complex (de la Sota, 1966), P. pyrrholepis is distinct by its short pinnae with rounded apices, and strongly reflexed, reddish brown rhizome scales.

505

Distribution. Epiphytic, rarely epipetric or terrestrial in montane rain forests; 1220–1700 m. Mexico; Guat, Nic, CR, Pan; Jam, Hisp; Col, Ven, Ec, Peru, Bol. Specimens Examined. Chis (Breedlove 26080, DS, NY, 32673, CHAPA, DS, NY, 38955, CAS, ENCB). Ver (Calzada 8339, XAL).

Mexican specimens are not as strongly dimorphic as those from Central and South America. Smith (1981) reported that at least some specimens occurred on wet rocks adjacent to and even within streams, as opposed to the species’ usual epiphytic habitat. Smith (1981) also reported that at least occasionally the spores of this species germinate while still inside the sporangia, and thus the sori may appear green. Polypodium remotum belongs to the P. plebeium group having black-centered rhizome scales with broad tan margins.

45. POLYPODIUM RHACHIPTERYGIUM 44. POLYPODIUM REMOTUM Polypodium remotum Desv., Me´m. Soc. Linn. Paris 6: 232. 1827. Type. “habitat in America calidiori,” Desvaux s.n. (P! photo BM!). Figs. 242N, O. Polypodium leucosticton Kunze ex Klotzsch, Linnaea 20: 380. 1847. Syntypes. Venezuela. Hartweg 1499 and Moritz 336 (BM–2 sheets!).

Rhizomes horizontal, compact, 4–5 mm diam.; rhizome scales bicolorous, each with black center or sclerotic cells and tan, membranous, entire or erose margins, 2–3.5 ⫻ 1 mm; fronds 26– 70 cm long, the fertile far exceeding the sterile, approximate; stipes 1⁄4–1⁄3 the frond length, stramineous to atropurpureous, slightly winged to base, with scattered, bicolorous, linearlanceolate scales 2–4 mm long; blades pinnatisect, narrowly deltate, the sterile ones 8–15 cm wide, the fertile ones 6–10 cm wide; pinnae 18–37 pairs, the sterile ones narrowly oblong, 3.5–7 cm ⫻ 6–8 mm, the fertile ones 2.5–4 cm ⫻ 2–4 mm, subcoriaceous; abaxial blade surfaces nearly glabrous, with sparse linearlanceolate scales ca. 1 mm long on or near costae; adaxial blade surfaces glabrous, with lime-dots; venation not visible, veins ending in hydathodes, often with lime-dots; sori round, medial; sporangia glabrous.

Polypodium rhachipterygium Liebm., Mexic. Bregn. 191 (reprint 39). 1849. Figs. 239N–Q. Goniophlebium rhachipterygium (Liebm.) T. Moore, Index Fil. 396. 1862. Lectotype (chosen by Smith, 1981: 193). Mexico. Oaxaca: Chinantla, Teotalcingo, Liebmann s.n. [Fl. Mex. 160] (C!; isotype BM, frag. US!). Polypodium stenoloma D. C. Eaton, Proc. Amer. Acad. Arts 8: 618. 1873. Type. Mexico. Chiapas: Ghiesbreght 386 (YU!; isotypes GH!, K!). Polypodium donnell-smithii Christ, Bull. Herb. Boissier, se´r. 2, 6: 291. 1906. Type. Guatemala. Alta Verapaz: Tu¨rckheim s.n. [Donn. Sm. 8823] (isotype US!).

Rhizomes creeping, 7–10 mm diam., greenish; rhizome scales ovate, pale brown, lustrous, 3–4 ⫻ 1–1.5 mm, somewhat caducous, point of attachment dark brown; fronds 20–70 cm long, distant; stipes ca. 2⁄5 the frond length, brown abaxially, stramineous adaxially, narrowly winged to base with 1 mm wide green wing, glabrous; rachises narrowly winged, glabrous; blades oblong, pectinate, with somewhat large terminal pinna, 16–23(–31) cm wide; pinnae 3–10 pairs, 1.2–2.6(–3.1) cm wide, linearlanceolate, slightly constricted near and dilated at bases; rachises, costae, and larger veins prominent, glabrous except for rare hairs 0.3–0.5 mm long on abaxial costae and appressed trichomidia on adaxial costae; margins with small, flap-like teeth; veins regularly uniting near margins to form a single row of areoles, free

506

POLYPODIUM

at margins; sori distinctly elongate-oblong, 2–3 mm long, 1 row each side of costae, medial; sporangia glabrous.

Distribution. Epiphytic in wet forests; 800–2100 m. Mexico; Guat, Hond, Salv, Nic, CR, Pan. Distribution. Epiphytic or on rocks in streams in wet montane forests of Atlantic slope; 80–1600 m. Mexico; Guat. Selected Specimens Examined. Chis (Purpus 6753, UC, 6871, UC, US). Oax (Hallberg 1605, NY; Herna´ndez G. 1996, NY; Mickel 4230, 6416, 7417, NY). Ver (Calzada 1539, NY; Martı´nez C. 02191, NY; Va´zquez T. 2487, NY).

This is one of the few species of Polypodium with distinctly elongate-oblong sori. The netted veins, broadly adnate pinnae, glabrous blades, and occurrence by streams at low elevations serve to identify it.

46. POLYPODIUM RHODOPLEURON Polypodium rhodopleuron Kunze, Linnaea 18: 315. 1844 [1845]. Syntypes. Mexico. Veracruz: Xalapa, Schiede & Deppe [745] (isosyntype P!); Leibold 91 (isosyntypes K!, P! p.p.). Figs. 238A–C. Polypodium isomeron E. Fourn., Mexic. Pl. 1: 80. 1872. Type. Mexico. “Pr. Teguitlan,” Hahn s.n. (P!). Polypodium lesourdianum E. Fourn., Mexic. Pl. 1: 82. 1872. Mexico. Veracruz: Orizaba, Botteri & Sumickrast 1471 in herb. LeSourd (MPU).

Resembling P. plesiosorum except: stipe and veins reddish, especially when young, but more or less persisting; rachises and pinnae totally glabrous; 2n⫽74 (Ver).

Selected Specimens Examined. Chis (Breedlove 21647, NY; Rzedowski 603, NY). Gro (Lorea 3177, US). Hgo (Chase 7307, US). Oax (Mickel 1094, 6603, NY). Pue (Sa´nchez M. 665, US). Qro (Dı´az B. & Carranza 6647, NY). Ver (Bourgeau 2887, NY; Ventura A. 295, NY).

47. POLYPODIUM RIOGRANDENSE Polypodium riograndense (T. Wendt) Mickel, comb. et stat. nov. Figs. 242A–C. Polypodium thyssanolepis Klotzsch var. riograndense T. Wendt, Amer. Fern J. 70: 6. 1980. Pleopeltis riograndensis (T. Wendt) E. G. Andrews & Windham in Windham, Contr. Univ. Mich. Herb. 19: 46. 1993. Type. U.S.A. Texas: Presidio Co., N side of Chinati Mountains, lower Indian Cave Canyon, Butterwick & Lott 3897 (TEX; isotypes GH!, MEXU).

Rhizomes creeping, 1 mm diam.; rhizome scales bicolorous, centers clathrate and black, lanceolate to linear-lanceolate, 2–3 ⫻ 0.3–0.5 mm, margins tan, denticulate; fronds (3–)5–13 cm long, subapproximate; stipes 1⁄3(–1⁄2) the frond length, tan to stramineous, with scattered lanceolate scales 2 mm long and smaller ovate scales 0.5 mm long, centers bicolorous, clathrate, marginal cells pale, thin-walled, with coarse teeth, tips attenuate; blades pinnatisect, narrowly deltate to oblong, 1–4 cm wide, decurrent; pinnae 2–9 pairs, narrowly oblong, 0.5–2.4 cm ⫻ 2–4 mm; adaxial blade surfaces with scattered to slightly imbricate ovate to deltate scales, bases with thick-walled brown central cells and paler margins, with coarse, often forked teeth, apices often attenuate; adaxial blade surfaces glabrous; venation obscure; sori round, medial; sporangia glabrous.

POLYPODIUM RZEDOWSKIANUM

507

acicular, fimbriate, each with reddish black center and whitish margins, 2–2.5 mm long; abaxial surfaces and costae completely hidden by the scales, these acicular, like those on the adaxial surfaces; sori impressed slightly (but this not apparent on the adaxial surface), round to slightly oblong; sporangia glabrous; 2n⫽74 (Mexico-state not given).

Distribution. Rock crevices or gravelly soil in pine-oak woods; 1500–2400 m. USA (Ariz, Tex); Mexico. Selected Specimens Examined. Chih (Correll & Gentry 23052, US; Knobloch 5945, NY; LeSueur 416, US; Pringle 443, NY [some sheets of this number are P. thyssanolepis]; Reeder et al. 3477, ARIZ; Spellenberg et al. 12119, UC). Coah (Henrickson 11652, LL; Johnston et al. 11888b, LL). Son (Fishbein 1784, ARIZ; Harvey 1706, US).

In contrast to P. thyssanolepis, the venation in this species is mostly free, laminae less scaly (not concealing the blade surfaces), stipes are only sparsely scaly, and the proximal pinnae are alternate. The stipe scales are mostly ovate or lance-ovate (vs. mostly suborbicular). There are some intermediate specimens in Chihuahua (Wendt, 1980). Polypodium thyssanolepis is widespread in Mexico, reaching its northern limit in central Chihuahua, whereas P. riograndense is limited in Mexico to Coahuila, Chihuahua, and Sonora. This species is distinct from P. fallacissimum in the coarse, often forked scale teeth and pinnae more perpendicular to the rachises. From P. thyssanolepis, P. riograndense differs in the small plant size, sparse stipe scales, mostly free veins, and alternate proximal pinnae.

Distribution. Epiphytic or epipetric on rocky slopes in oakjuniper grasslands, pine-oak dry, forests, and moist ravines, montane, (600–)1000–3000 m. Mexico; reported from Honduras and Costa Rica by Moran (in Davidse et al., 1995), but doubtful. Selected Specimens Examined. Dgo (Breedlove 59125, NY). Gro (Hinton et al. 14498, NY). Jal (McVaugh 26022, NY). Me´x (Dunn & Dunn 18933, NY). Mich (Rzedowski 34627, ENCB). Mor (Lyonnet 859, MEXU, NY). Nay (Breedlove 44341, CAS). Oax (Campos V. 2201, MEXU). Sin (Breedlove 44890, NY). Zac (McVaugh 25789, NY). Unverified, Doubtful, or Mistaken Reports. Qro (Ferna´ndez N. 3755, ENCB, cited by Dı´az-Barriga & Palacios-Rios, 1992, but not verified).

This species is distinct from P. sanctae-rosae by the rhizome scales, which are larger (over 4–5 mm long instead of 0.8–1.5 mm).

48. POLYPODIUM ROSEI Polypodium rosei Maxon, Contr. U.S. Natl. Herb. 17: 594. 1916. Type. Mexico. Morelos: Sierra de Tepostla´n, Rose & Painter 7254 (US!). Figs. 242K–M. Polypodium hirsutissimum Raddi var. sericeum M. Martens & Galeotti, Me´m. Foug. Mexique 42. 1842. Type. Mexico. Michoaca´n: Vicinity of Morelia, Galeotti 6432 (BR).

Rhizomes creeping, 3–6 mm diam.; rhizome scales slightly reflexed, flaccid, linear-lanceolate with sparse teeth, light reddish brown, darker in the center with age and then weakly bicolorous, 4–5 mm long; fronds (10–)20–50 cm long; stipes 1⁄5–2⁄5 the frond length, with abundant scales, stramineous to orange-tan, with red-black point of attachment and acicular tip 4–7 mm long; blades pinnatisect, long-lanceolate, 4.5–11 cm wide, the longest pinnae 2.4–6 cm ⫻ 4–6 mm; pinnae 18–37 pairs, linear, the terminal pinna small to medium-sized; nectaries on acroscopic surcurrent pinna bases; adaxial surfaces heavily scaly, the scales

49. POLYPODIUM RZEDOWSKIANUM Polypodium rzedowskianum Mickel, sp. nov. Type. Mexico. Oaxaca: Dto. Teotitla´n, Mpio. de Tenango, midway point (Cerro del A´lamo) on trail from Jalapa de Dı´az to San Martı´n Caballero, Hallberg 1364 (holotype NY!). Figs. 236O–S. A P. plebeio pinnis latis longi-acuminatis basi valde constrictis differt. (Honoring the outstanding Mexican floristician Jerzy Rzedowski).

Rhizomes creeping, 3.5–5 mm diam.; rhizome scales reflexed, 2–3 ⫻ 0.5–0.8 mm, bicolorous, central stripe black, narrow, lumina visible, 0.1 mm wide, margins brown, thin, ruffled, erosedenticulate; fronds 30–58 cm long, distant; stipes 1⁄3–2⁄5 the frond length, reddish brown, non-alate, with appressed scales of various shapes, dark streak and broad thin margins; blades pinnatisect to pinnate, broadly ovate, 10–26 cm wide; pinnae 4–12(–18) pairs, linear-lanceolate, constricted toward base (2–4 mm wide)

508

POLYPODIUM

and dilated at rachises (sometimes more surcurrent or more decurrent), to 7–17 cm ⫻ (8–)10–12 mm, apices long-acuminate; adaxial surfaces glabrous; abaxial surfaces with scattered small, lanceolate, brown to black scales 0.5–1 mm long, hydathodes prominent, often in two rows on each side of costae; margins modified, clear, serrate to crenate or entire; apices entire; veins free, 3–4 forked, obscure; sori round, medial; sporangia glabrous; 2n⫽70 or 72 (Oax).

rhizome scales); blades pinnate to pinnatisect, with small to medium-sized terminal pinnae, lanceolate to deltate-lanceolate, 8–15 cm wide; pinnae 20–30 pairs, 5–6.5 cm ⫻ 5–8 mm wide, linear; nectaries on acroscopic surcurrent pinna bases; costae castaneous at bases, covered with scales; adaxial surfaces with sparse scales, glabrescent with age, the scales acicular and fimbriate with centers reddish and margins whitish, 2–2.5 mm long; abaxial surfaces with scales completely hiding the surfaces and costae, linear-lanceolate, like scales on adaxial surfaces; sori impressed slightly but not embossed, round to slightly oblong; sporangia glabrous; 2n⫽74 (Oax).

Distribution. Epiphytic, epipetric, or terrestrial in wet montane forests; 1200–1850(–2150) m. Mexico. Specimens Examined. Gro (Hinton et al. 11071, NY; Lorea 2062, NY; Rzedowski 18525, DS, 18605, NY). Jal (McVaugh 20399, IEB, NY). Oax (Geith 95-M040, NY; Hallberg 1454, 1455, NY; Mickel 6040, 6183, 6219, NY; Panero 5372, NY; Toriz 909, NY; Torres C. 11899, NY). Sin (Lehto 24391, ASU).

Polypodium rzedowskianum is a member of the P. plebeium complex but is distinct in its broad, long-acuminate pinnae that are conspicuously constricted at the bases. This species has been misconstrued as P. pleolepis Maxon & Copel. (Mickel & Beitel, 1988), whose type closely resembles this species in pinna form. That species, however, is a synonym of P. bradeorum and is distinct in its orange-tan, weakly bicolorous rhizome scales (vs. strongly bicolorous, black-centered scales) and oblong to linear sori (vs. round).

Distribution. Epiphytic in pine-oak and oak woods; 800–2700 m. Mexico; Guat, Salv, Hond, Nic. Selected Specimens Examined. Chis (Breedlove 25504, 27548, NY). Col (Va´zquez 1303, UC, photocopy of specimen at UCR). Jal (Boutin & Brandt 2489, NY; McVaugh 21365, MICH; Palafox 5409, NY). Me´x (Tejero-Dı´ez, 2435, IZTA). Nay (McVaugh 16445, NY). Oax (Mickel 867, 3734, NY; Ramı´rez G. 556, NY). Unverified, Doubtful, or Mistaken Reports. Mich (Arse`ne 5226, MEXU, cited by Dı´az-Barriga & Palacios-Rios, 1992, as Polypodium squamatum L., but that species occurs only in the West Indies; this specimen is most likely Polypodium sanctae-rosae or P. rosei).

Polypodium sanctae-rosae is the most abundant member of the P. squamatum complex (de la Sota, 1966) and is distinguished by its mostly appressed rhizome scales and pinnae 5–6.5 cm long.

50. POLYPODIUM SANCTAE-ROSAE 51. POLYPODIUM SCOULERI Polypodium sanctae-rosae (Maxon) C. Chr., Index Filic., Suppl. 1: 62. 1913. Figs. 233J–S. Goniophlebium sanctae-rosae Maxon, Contr. U.S. Natl. Herb. 13: 8. 1909. Type. Guatemala. Baja Verapaz: Near Santa Rosa, Tu¨rckheim II. 1607 (US!; isotypes F, MO, NY!, P, US!).

Rhizomes creeping, 5–7 mm diam.; rhizome scales appressed to slightly reflexed, broadly ovate to linear-lanceolate, fimbriate, 0.8–1.5 mm long, each with a dark red-brown center and whitish margins; fronds 40–55 cm long, distant; stipes 1⁄4–1⁄2 the frond length, with scales mostly appressed, of two types, one similar to rhizome scales, the other dark-centered with toothed acicular tips 2 mm long, spreading (even on specimens with only small

Polypodium scouleri Hook. & Grev., Icon. Filic. 1: t. 56. 1829. Type. U.S.A. Western North America, near Columbia River, Scouler s.n. (K). Figs. 245A, B. Rhizomes creeping, glaucous, 5–6 mm diam.; rhizome scales lustrous brown, lanceolate, subconcolorous (castaneous centers with slightly paler narrow margins), entire to minutely denticulate near apices, 3–5 ⫻ 1.5–2 mm; fronds 13–30 cm long, distant; stipes 1⁄3–1⁄2 the frond length, orange-tan, with scales lanceolate to deltate-attenuate, slightly bicolorous; blades pinnatisect, lanceolate-deltate, very coriaceous, deltate, 8–10 cm wide; pinnae 3–8 pairs, 4–5 ⫻ 1–1.6 cm, oblong, entire, apices rounded; ab-

POLYPODIUM THYSSANOLEPIS

axial blade surfaces glabrous except for sparse dark, deltate, attenuate scales 2–4 mm long; adaxial blade surfaces glabrous; venation anastomosing, obscure; sori round, near costae, ca. 4 mm diam.; sporangia glabrous; 2n⫽74 (USA).

509

oblong-deltate or lanceolate, 1-pinnate, pinnatifid at apices, 7– 23 cm wide; pinnae 16–28 pairs, linear-lanceolate, 9–18 mm wide, bases rounded to truncate, adnate only in upper part of blade, apices acuminate; abaxial surfaces glabrous with hairs 0.5 mm scattered on costae, veins, and margins; adaxial surfaces with rare to scattered white hairs; margins finely serrulate to crenulate to subentire; veins free or with occasional anastomoses; sori round to slightly oblong; sporangia not obviously setose but each with one or more short stubby hairs 0.1 mm long.

Distribution. Epiphytic in pine forests; 0–550 m. Canada (BC); USA (Calif, Ore, Wash); Mexico. Specimens Examined. Guad (Hanna s.n., 15 July 1922, CAS, US; Moran 6420, 6422, UC; Newcomb 169, UC, US; Palmer 104, NY; Rico C. 5, CHAPA; Wiggins & Ernst 46, UC).

Polypodium scouleri is distinct by its very leathery fronds with few, apically rounded, entire pinnae and sori over 3 mm diam. Its venation is mostly netted but quite variable.

52. POLYPODIUM SUBPETIOLATUM Polypodium subpetiolatum Hook. in Bentham, Pl. Hartw. 54. 1840. Type. Mexico. [Hidalgo:] Regla, Hartweg s.n. [414] (K!; isotype P!). Figs. 235G, 237F, G. Polypodium biserratum M. Martens & Galeotti, Me´m. Foug. Mexique. 38. 1842. Type. Mexico. Oaxaca: Llano Verde, Galeotti 6451 (BR!). Polypodium schaffneri Fe´e, Me´m. Foug. 9:22. 1857. Type. Mexico. [Distrito Federal:] San Angel, Schaffner 310; Popocate´petl, Schaffner 273 (P?). Polypodium guilleminianum E. Fourn., Mexic. Pl. 1: 77, t. 2. 1872. Type. Mexico. San Rafael, Guillemin s.n. (P!, photo BM!). Polypodium hahnii E. Fourn., Mexic. Pl. 1: 79. 1872. Type. Mexico. [Veracruz:] Misantla, Hahn s.n. (P!). Polypodium firmulum Maxon, Contr. U.S. Natl. Herb. 8: 274, pl. 61, f. 3a, 3b. 1903, hom. illeg., non Baker, 1893. Polypodium maxonii C. Chr., Index Filic. 543. 1906. Type. Mexico. San Luis Potosı´, Palmer 448 (US!; isotypes GH!, NY!). Polypodium adelphum Maxon, Contr. U.S. Natl. Herb. 8: 275, t. 62. 1903. Type. Mexico. Chiapas: Ghiesbreght 244 (GH! p.p., photo NY!; isotypes BM, K, NY, YU p.p.!). Polypodium teresae Maxon, Contr. U.S. Natl. Herb. 8: 274, t. 61, f. 2. 1903. Type. Mexico. [Nayarit:] Territorio de Tepic, Sierra Madre, near Santa Teresa, Rose 2205 (US! frag. NY!; isotypes GH!, NY!).

Rhizomes creeping to long-creeping, 3–5 mm diam., often slightly glaucous; rhizome scales ovate-lanceolate to ovate-deltate, long-attenuate, entire, glabrous or comose, overlapping-cordate, 6–7 mm long; fronds 30–75 cm long, distant; stipes 1⁄3–2⁄5 the frond length, glabrous, stramineous, rarely brown; blades

Distribution. Epiphytic in oak and pine-oak forests; 1700– 3100 m. Mexico; Guat, Hond. Selected Specimens Examined. Ags (Corres Z. 16106, GUADA, IEB). Chih (Correll & Gentry 22998, NY). Chis (Breedlove 22596, DS). Coah (Barkley et al. 7398, TEX). Col (Goldsmith 38, US). DF (Rusby 369, NY). Dgo (Breedlove 18737, NY). Gro (Rzedowski 26303, NY). Gto (Rzedowski 50559, NY). Hgo (Moore 5301, US). Jal (McVaugh 19605, NY). Me´x (Rzedowski 30912, NY). Mich (Pringle 3328, NY). Mor (Lyonnet 892, MEXU, NY). Nay (Rose 2205, NY). NL (Hinton et al. 22197, NY). Oax (Mickel 3705, NY). Pue (Purpus 4038, NY). Qro (Rzedowski 48648, IEB). Sin (Breedlove 18357, NY). SLP (Pennell 17862, US). Tam (Trejo Avila s.n., 19 Oct 1985, UNL). Ver (Bourgeau 1038, NY).

The characters used in Oaxaca (Mickel & Beitel, 1988) and Chiapas (Smith, 1981) treatments to distinguish P. subpetiolatum from P. adelphum (glabrous rhizome scales, free veins, and densely hairy adaxial costae) are not consistent and are found in varying combinations throughout the range of P. subpetiolatum. Some specimens from Chiapas have more cuneate pinna bases, and some specimens (also from Chiapas) have subconform terminal pinnae rather than strictly pinnatifid apices. Even in the type collection of P. adelphum both pinnatifid apices and subconform terminal pinnae occur. This species varies in having pinnae subentire to coarsely toothed or crenate and subauriculate; some specimens have very narrow pinnae.

53. POLYPODIUM THYSSANOLEPIS Polypodium thyssanolepis A. Braun ex Klotzsch, Linnaea 20: 392. 1847. Figs. 234K–N, 242D, E. Goniophlebium thyssanolepis (A. Braun ex Klotzsch) T. Moore, Index Fil. 396. 1862. Marginaria thyssanolepis (A. Braun ex Klotzsch) Farw., Amer. Midl.

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Naturalist 12: 299. 1931. Syntypes. Colombia. Moritz 22 (B!; isosyntype P), Otto 896 (B!; isosyntype P). Polypodium incanum Sw. var. fimbriatum M. Martens & Galeotti, Me´m. Foug. Mexique 36. 1842. Lectotype (chosen by Morton, Contr. U.S. Natl. Herb. 38: 257. 1973). Mexico. Jalisco: “Barranca de Guadalajara,” Galeotti 6438 (BR!; isolectotypes BR!, P?). Polypodium rhagadiolepis Fe´e, Me´m. Foug. 5: 237. 1852. Goniophlebium rhagadiolepis (Fe´e) Fe´e, Me´m. Foug. 7: 62, t. 19, f. 3. 1857. Type. “Insula Cuba et in Mexico.” Mexico. [Chiapas:] Yerba Buena, Linden s.n. (P!). Polypodium purpusii Christ, Bull. Herb. Boissier, se´r. 2, 7: 416. 1907. Type. Mexico. 1904, Purpus s.n. (P!; isotype US!, frag. NY!).

dathodes on the blades and a black central portion of the rhizome scales. Specimens range from large (e.g., Hallberg 1773, NY, from Oaxaca, 50 ⫻ 18 cm) to very small in the northern part of the range (e.g., Reveal 2730, ARIZ, from Durango, 3.5 ⫻ 1.5 cm, only 1 or 2 pinna pairs).

Rhizomes slender, creeping, 3–5 mm diam.; rhizome scales linear-lanceolate, appressed, imbricate, bicolorous, light brown with central black sclerotic streak, 3–4 ⫻ 0.8–1 mm, margins fringed; fronds (3.5–)10–40 (–50) cm long, distant; stipes ca. 1⁄2–2⁄3 the frond length, densely scaly with round (0.3–0.5 mm diam.) to lanceolate (1.5–2 mm long) scales; blades ovate to deltatelanceolate, pinnatifid, 3–10(–18) cm wide, with apices pinnatifid; pinnae (1–)3–15 pairs, 5–7 mm wide, entire, obtuse; nectaries on acroscopic surcurrent pinna bases; adaxial surfaces glabrous to sparsely scaly with acicular fimbriate scales; abaxial surfaces densely scaly, the scales mixed, round (0.5 mm diam.) to lanceolate (1–1.5 mm long), dark brown-centered, margins pale and ciliate-denticulate; veins partially netted, obscure, hydathodes prominent with white deposits; sori round to slightly oblong, in a single row on each side of the costae, mostly hidden by the scales; sporangia glabrous; 2n⫽74 (Sin, Peru), 2n⫽ca. 148 (Jam).

Polypodium triseriale Sw., J. Bot. (Schrader) 1800(2): 26. 1801. Type. s. coll. s.n. (S). Figs. 239G, H, J.

Distribution. Epipetric or terrestrial on rocky slopes in mesic to dryish areas, open woods or clearings; 850–3000 m. Sw USA; Mexico; Guat, Hond, Nic, CR, Pan; Jam, Hisp; Col, Ven, Ec, Peru, Bol. Selected Specimens Examined. Ags (McVaugh 18388, MICH). Chih (Pringle 977, NY). Chis (Breedlove 26837, NY). Coah (Johnston et al. 11506C, LL). Col (Jones 506, US). DF (Rusby 372, NY). Dgo (Reveal 2730, NY). Gro (Mayfield et al. 1012, UC). Gto (Rzedowski 40583, IEB). Hgo (Dunn & Dunn 19019, NY). Jal (Rzedowski 26214, NY). Me´x (Rzedowski 24308, NY). Mich (Leavenworth 326, NY). Mor (Lyonnet 857, MEXU, NY). Nay (Mexia 768, NY). NL (Lavin 4861, NY). Oax (Mickel 3901, NY). Pue (Nicolas s.n., 3 Nov 1909, NY). Qro (Rzedowski 46626, IEB). Sin (Breedlove 18502, NY). SLP (Pringle 3388, NY). Ver (Bourgeau 2700, NY). Zac (McVaugh 17762, NY).

Polypodium thyssanolepis is distinct from members of the P. squamatum complex (de la Sota, 1966) by the presence of hy-

54. POLYPODIUM TRISERIALE

Goniophlebium haenkei C. Presl, Tent. Pterid. 186, pl. 7. 1836. Polypodium haenkei (C. Presl) Liebm., Mexic. Bregn. 204 (reprint 52). 1842. Type. Mexico. Haenke s.n. (PRC).

Rhizomes short-creeping, 7–10 mm diam.; rhizome scales clathrate with narrow, pale margins, central portion light brown with thickened dark brown walls, ovate-acuminate, 4–5 ⫻ 2–3 mm; fronds 36–90 cm long, 1–2 cm distant; stipes 1⁄4–2⁄5 the frond length, stramineous to castaneous, glabrous; blades 1-pinnate, broadly oblong, 16–50 cm wide, terminal pinnae conform; pinnae 3–14 pairs, 1.2–3.2 cm wide, proximal ones with cuneate bases, distal ones broadly adnate, linear-oblong, acuminate, glabrous except for rare to scattered brown trichomidia 0.1–0.3 mm long, margins entire; veins netted, with 3–4 rows of areoles between costae and pinna margins; sori round, in 2–3 rows between costae and pinna margins; sporangia glabrous; 2n⫽74 (Oax), 148 (Jam).

Distribution. Epiphytic, widespread in wet forests; 100–1500 m. Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Jam, PR, L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol, Parag. Selected Specimens Examined. Camp (Ucan & Andrews 5245, UADY). Chis (Martı´nez S. 11706, NY; Purpus 6757, NY). Gro (Lorea 4797, IEB). Oax (Hallberg 1624, NY; Mickel 5940, NY). Pue (Sa´nchez M. 644, US). Tab (Cowan 3108, NY; Zamudio R. 1007, IEB). Ver (Rzedowski 19080, NY). Unverified, Doubtful, or Mistaken Reports. Mich (Arse`ne 6690, MEXU, cited by Dı´az-Barriga & Palacios-Rios, 1992, but not verified). QR (Cabrera & de Cabrera 4205, CIQR, MEXU, cited by Palacios-Rios, 2002c, but not verified).

POLYSTICHUM

Polypodium triseriale is distinct by its venation and sori in 2–3 rows on each side of the midvein.

55. POLYPODIUM VILLAGRANII Polypodium villagranii Copel., Univ. Calif. Publ. Bot. 19: 292. 1941. Type. Mexico. Hidalgo: Santa Ana, Copeland herb. 143 (MICH!, photo at UC; isotypes NY!, UC!, US! Figs. 241E–H, J. Rhizomes creeping, 1.5–2 mm diam.; rhizome scales concolorous tan, lanceolate, entire to minutely denticulate, 4–6 ⫻ 1 mm; fronds (9–)13–28 cm long, 3–15 mm distant; stipes 1⁄8–1⁄4 the frond length, castaneous to atropurpureous, densely clothed with patent lanceolate scales 4–6 mm long like those on rhizomes, other scales, like those on blades; blades pinnatisect, narrow to linearoblong, (1.5–)2–4 cm wide; pinnae 28–44(–86) pairs, linear, 8– 20 mm long, sterile 2–3.5 mm wide, fertile ca. 2 mm wide; abaxial blade surfaces with dense linear-lanceolate to acicular scales with round fimbriate bases and long-attenuate apices, orangetan but with reddish brown point of attachment, 3–4 mm long, some scales lanceolate, to 1.5 mm long; scales broader overlying sori; adaxial blade surfaces with scales with round fimbriate bases, reddish brown at point of attachment, acicular ca. 2 mm long, glabrescent with age; venation obscure; sori round, medial; sporangia glabrous.

Distribution. Epiphyte on oaks and alder in wet montane forests and dry oak forests; 1000–1300 m. Mexico. Selected Specimens Examined. Gro (Faxson et al. 17M740, TEX). Hgo (Barkley 17M027, F, TEX, US; Moore 5084, UC, US; Sharp 441765, US). Oax (Mickel 7124, NY). Pue (Sa´nchez Mejorada 656, US). Qro (Rzedowski 46800, IEB). SLP (Rzedowski 10972, ENCB; White 19A, US). Ver (Nee 22495, XAL).

Polypodium villagranii closely resembles P. furfuraceum, but is distinct in having the adaxial blade scales subentire, with bases slightly toothed (vs. stellate), and rhizome scales lanceolate and entire (vs. finely toothed). Unplaced Names Goniophlebium serratum Fe´e, Me´m. Foug. 8: 95. 1857, non Polypodium serratum Aubl., 1775. Type. Mexico. [Veracruz:] Ori-

511

zaba, Schaffner 495 (P!). Though treated as a synonym of Polypodium echinolepis (Smith, 1981), the type has glabrous blades and looks somewhat like P. fraternum. However, it lacks a rhizome and thus the diagnostically critical rhizome scales. Polypodium arthropodium Fe´e, Me´m. Foug. 8: 88. 1857. Syntypes. Mexico. [Veracruz:] Cordoba, Schaffner 185 (RB), and Orizaba, Schaffner 186 (RB). From the type description this name may apply to P. longepinnulatum, but that species has not been found in the Valley of Mexico. If it proves to be the same, P. arthropodium has priority.

1 0 2 . P O L Y ST I C H U M Polystichum Roth, Tent. Fl. Germ. 3: 31, 1799, emend. Schott, Gen. Fil., t. 9. 1834. Lectotype (chosen by Schott, loc. cit.): Polystichum lonchitis (L.) Roth [⬅ Polypodium lonchitis L.]. See Pichi Sermolli (Webbia 9: 436. 1953) for more on typification. Plecosorus Fe´e, Me´m. Foug. 5: 150, t. 13A. 1852. Lectotype (first chosen by J. Smith?, Hist. Fil. 11875): Plecosorus speciosissimus (Kunze) T. Moore [⬅ Cheilanthes speciosissimus Kunze] ⬅Polystichum speciosissimum (Kunze) Copel.

Terrestrial or epipetric; rhizomes ascending to erect, short, compact, usually stout, heavily scaly, with many old leaf bases forming stout caudices; fronds monomorphic, medium-sized to large, mostly 50–150 cm long, crown-forming; stipes usually stramineous to tan, usually densely scaly, at least at their bases, scales lanceate to broadly ovate, subentire to erose or denticulate, sometimes also ciliate with long, thin hair-like processes; blades narrowly to broadly deltate, 1–2-pinnate, rarely 3-pinnate, anadromous, some species proliferous with rachis bud near blade apices or in pinna axils; rachises and costae grooved adaxially, the grooves confluent from one axis to the next; segments thin to commonly subcoriaceous or coriaceous, usually with marginal spinulose teeth, glabrous to scaly abaxially, especially on veins and axes, lacking acicular hairs; veins free, forking; sori abaxial, round, typically with peltate indusia, or indusia absent; spores bilateral, with a winged or strongly folded, sometime echinate and/or fenestrate perispore; x⫽41. Polystichum is a largely temperate genus of nearly 200 species, with most of the tropical species at higher elevations, although a few occur at middle elevations in wet forests. The genus is difficult to circumscribe and distinguish clearly from Cyrtomium and Phanerophlebia, and a broad view of Polystichum could include both segregates. Recent phylogenetic work by Little and Barrington (2003) indicates that Phanerophlebia is sister to Polystichum s.l. (including the segregrate genera Cyrtomium and Cyrtomidictyum in Asia), but that Polystichum s.str. can be made monophyletic by the recognition of both of these Asian segregates. Polystichum is usually distinguished by its peltate indusia, 2-pinnate blades, and free veins, but many exindusiate and 1pinnate species are known, even among the Mexican taxa. Of the Mexican species, about half are well defined, mostly allopatric with respect to congeners in Mexico, and rather restricted or rare in their distribution. These include nearly all of

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the exindusiate species (P. mickelii, P. orbiculatum, P. platyphyllum, P. schizophyllum, and P. speciosissimum), as well as the 1pinnate ones (P. acrostichoides, P. muenchii, P. munitum). The remaining ten species are more difficult to distinguish from each other, and very likely they have incompletely diverged or have come into contact again after being allopatric previously. Mostly these appear to be diploids, but there is evidence that some populations are tetraploid, as judged by the large spores of some herbarium specimens and a few chromosome counts. The central and perhaps most variable species seems to be P. hartwegii, which is also the most widely distributed and common of the Mexican species. The species of Polystichum appear difficult to define, in part, because of probable hybridization. In tropical regions, this is not very well documented, except for the studies of Barrington (1985a, 1985b), but in temperate regions, hybridization is well studied in northwestern North America, Europe, and Japan. Extensive work needs to be done on the morphological diversity and ecology of individual populations for a better understanding of the species and their dynamics in tropical America. It is also not likely that a fuller understanding of the systematics can be achieved until modern cytotaxonomic and isozymic studies have been done. Additional species may well be recognized once proper studies can be made, and we allude to some interesting unnamed collections at the end of this treatment; it would, however, be premature to describe them now because of the general taxonomic confusion in the genus and also the paucity of collections.

The genus Rumohra, sometimes considered to be related to Polystichum and similar in its peltate indusia, is known to us from a single Mexican collection of Rumohra adiantiformis (G. Forst) Ching: Jalisco, Los Guayabos, “alterado, en una huerta, lugar hu´medo,” 1500 m, Pilar Mones U. 18218 (UC). It is unclear from the label whether this species, often cultivated around the world, has become established and is reproducing on its own or not. It is generally considered to be a circumaustral, polymorphic species, and is also considered native to much of South America (e.g., Venezuela, Peru, Brazil, Bolivia, Chile, Argentina), S. Africa, Madagascar, Australia, and New Zealand. It is also apparently indigenous to the Greater Antilles, but is unknown from Mesoamerica. From Polystichum, Rumohra differs in its adaxially flattened (vs. sulcate) costae, long-creeping dorsiventral rhizomes (vs. short-creeping to suberect, radial), and apically rounded to acute (vs. aristate) ultimate segments. References Barrington, D. S. 1985a. The morphology and origin of a new Polystichum hybrid from Costa Rica. Syst. Bot. 10: 199–204; Barrington, D. S. 1985b. Hybridisation in Costa Rican Polystichum. Proc. Royal Soc. Edinburgh 86B: 335–340; Barrington, D. S. 1990. Hybridization and allopolyploidy in Central American Polystichum: cytological and isozyme documentation. Ann. Missouri Bot. Gard. 77: 297–305; Barrington, D. S. 1992. Climate and the disjunct distribution of Polystichum alfaroi (Christ) Barr., comb. nov. in Mesoamerica. Rhodora 94: 327– 339; Little, D. P. & D. S. Barrington. 2003. Major evolutionary events in the origin and diversification of the fern genus Polystichum (Dryopteridaceae). Amer. J. Bot. 90: 508–514; Mickel, J. T. 1997. A review of the West Indian species of Polystichum. Pp. 119–143. In: R. J. Johns (ed.), Holttum memorial volume. Royal Botanic Gardens, Kew; Wagner, D. 1979. Systematics of Polystichum in western North America north of Mexico. Pteridologia 1: 1–64.

Key to the Mexican Species of Polystichum 1. Indusia absent. 2. Bud in axil of a distal pinna; stipe base scales concolorous, 5–10 ⫻1–3 mm, with long-ciliate margins. ................................ 14. P. platyphyllum. 2. Buds absent; stipe base scales concolorous or bicolorous, 8–40 ⫻ 2–6 mm, margins sparingly toothed, erose-denticulate, or ciliate. 3. Pinnules incised nearly or completely to costules, the ultimate segments bead-like, lacking spinulose tips or margins, with whitish false indusia; proximal several pinna pairs somewhat reduced. ............................................................................... 17. P. speciosissimum. 3. Pinnules not or only slightly incised, not cut to the costules, the ultimate segments (pinnules) rhombic or oblong, spinulose at tips and sometimes at margins, lacking whitish false indusia; proximal pinna pair(s) not or only slightly reduced. 4. Blades ovate-attenuate; pinnae 6–20 cm long; sori discrete at maturity; pinnules spinulose at tips and sometimes at margins, margins not strongly revolute; low to middle elevations (450–2500 m); s. Mexico. .............................................. 8. P. mickelii. 4. Blades linear-lanceolate; pinnae 1.5–4 cm long; sori confluent at maturity; pinnules sparingly or not at all spinulose, margins strongly revolute; very high elevations (3600–4200 m); Chiapas. ...................................................................... 12. P. orbiculatum. 1. Indusia present, caducous in several species. 5. Blades 1-pinnate, pinnae dentate or serrate at margins. 6. Sori confined to distal pinna pairs, the fertile pinnae contracted relative to sterile ones; sori confluent at maturity; Nuevo Leo´n. ................................................................................................................................................................................................... 1. P. acrostichoides. 6. Sori not confined to distal pinna pairs, the fertile pinnae about the same size and shape as sterile ones; sori discrete at maturity. 7. Pinnae acute or rounded; rachises sparsely scaly; Chiapas. ....................................................................................................... 9. P. muenchii. 7. Pinnae acuminate; rachises densely scaly; Guadalupe Isl. ....................................................................................................... 10. P. munitum. 5. Blades 1-pinnate-pinnatifid, 2-pinnate, or more divided. 8. Blades fully 3-pinnate, at least in the proximal portion, the larger pinnulets lobed, at least acroscopically. ................... 16. P. schizophyllum. 8. Blades 2-pinnate, the larger pinnules sometimes lobed or each with a single nearly free acroscopic segment at the pinnule bases.

POLYSTICHUM ACROSTICHOIDES

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9. Stipe bases with numerous ovate-lanceolate scales 1–3 cm ⫻ 2–6(–8) mm, these black and lustrous, or black with very narrow, light brown, ciliate margins; rachises with a few small blackish scales; stipes mostly ca. 1⁄4–1⁄2 the blade length; segments decidedly spinulose; laminae coriaceous. ........................................................................................................................ 3. P. distans. 9. Stipe base scales dark to light brown (some black and lustrous in P. erythrosorum), ovate-lanceolate to linear-lanceolate; rachises without blackish scales; stipes mostly more than one-half the blade length; segments strongly to weakly spinulose. 10. Stipe and rachis scales strongly spinulose, teeth to 0.4 mm long; pinnules auriculate but otherwise entire, margins spinulose, especially near tips; stipe and rachis scales very dense, brown, concolorous; 2600–2800 m. .............. 6. P. furfuraceum. 10. Stipe and rachis scales denticulate to subentire; pinnules toothed, pinnatisect, or entire; stipe and rachis scales moderately dense to sparse, brown, concolorous or bicolorous with lighter margins; 900–3100 m. 11. Indusia reddish brown; pinnules entire or acroscopically auriculate; some stipe base scales black and lustrous, 10–20 ⫻ 4–6 mm; 2300–3000 m. ............................................................................................................................ 4. P. erythrosorum. 11. Indusia whitish to tan or brown; pinnules entire to pinnatisect; stipe base scales dark to light brown, 5–20 ⫻ 1– 4 mm; middle to high elevations. 12. Indusia mostly 0.8–1.5 mm diam., persistent. 13. Plants small, pinnae less than 8(–10) cm long; proximal pinnae often strongly deflexed; pinnae fewer than 20 pairs per frond; stipe base scales linear-lanceolate, mostly bicolorous, 5–10 ⫻ 1–2 mm, with spinulose margins. .............................................................................................................................................. 5. P. fournieri. 13. Plants larger, with pinnae mostly greater than 10 cm long; proximal pinnae not usually strongly deflexed; pinnae usually more than 20 pairs per frond; stipe base scales linear-lanceolate to ovatelanceolate, concolorous to bicolorous, 8–20 ⫻ 1–5 mm, margins spinulose or not. 14. Pinnules deeply incised or toothed, each with a nearly free acroscopic lobe; indusia tan to brown. ...................................................................................................................................................................... 7. P. hartwegii.

14. Pinnules merely dentate or auriculate, without free lobes; indusia whitish or tan. ................... 15. P. rachichlaena. 12. Indusia 0.5–0.8(–1) mm diam., often not persisting, or hidden by mature sporangia. 15. Stipe base scales and rhizome scales 5–9 ⫻ 1–3 mm, with erose-denticulate margins; 2000–3100 m. ............................................................................................................................................................................ 18. P. turrialbae.

15. Stipe base scales and rhizome scales 7–22 ⫻ 1–6 mm with ciliate margins; 900–2700 m. 16. Stipe base scales strongly marginate, the margins irregular, with rigid to flaccid cilia throughout scale length; fertile pinnules angled ca. 70–80⬚ to costae; 1000–1400 m; Chiapas. ............................ 2. P. alfaroi. 16. Stipe base scales not strongly marginate, the margins entire or ciliate; fertile pinnules mostly 80– 90⬚ to costae; 700–2800 m; widespread in Mexico. 17. Several pairs (2–8) of proximal pinnae reduced to 0.3–0.8 times length of longest pinnae; stipe base scales sharply bicolorous, with greatly thickened bases. .......................................... 13. P. ordinatum. 17. Several pairs of proximal pinnae not reduced or basal pinnae only slightly reduced; stipe base scales not sharply bicolorous, lacking thickened bases or the bases sometimes slightly thickened. 18. Stipe base scales bicolorous, with a dark brown central stripe and lighter brown margins, linear-lanceolate (mostly 1–2 mm wide), ciliate. .................................................................... 7. P. hartwegii. 18. Stipe base scales light brown, concolorous, broadly ovate (to 6 mm), entire. .............. 11. P. muricatum.

1. POLYSTICHUM ACROSTICHOIDES Polystichum acrostichoides (Michx.) Schott, Gen. Fil., t. 9. 1834. Fig. 247A. Nephrodium acrostichoides Michx., Fl. Bor.-Amer. 2: 267. 1803. Type. “Pensylvania, Carolina, Tennessee, Carol. Marit.” Michaux s.n. (P-MICH; see Morton, Amer. Fern J. 57: 172. 1967).

Caudices ca. 2 cm diam.; fronds mostly 30–70 cm long; stipes 8–30 cm long, 2–3 mm diam., mostly ca. 1⁄4–2⁄3 the blade length; stipe bases with numerous concolorous, tan, ovate to lanceolate scales 5–15 ⫻ 1–4 mm, with narrow teeth along margins; blades hemidimorphic, on a given frond the distal pinnae (8–25 pairs)

fertile, constricted, proximal pinnae sterile, broader and longer, chartaceous, blades to ca. 50 ⫻ 12 cm, broadest near the bases, attenuate at the pinnatifid apices; rachises lacking buds, moderately scaly with ovate to hair-like toothed scales to 5 mm long; pinnae undivided, very gradually shortened distally, ca. 22–35 pairs, mostly 3–6 ⫻ 0.8–1 cm (excluding basal auricle), lanceate with a pronounced acroscopic, narrowly deltate auricle 3–8 mm long, otherwise serrate-spinulose along both margins, basiscopic bases cuneate, tips acute and spinulose; indument abaxially along costae and veins of scattered hair-like, twisted scales; sori indusiate, completely confluent at maturity, indusia tan to brown with darker brown centers, ca. 1–1.2 mm diam., subpersistent; 2n⫽82 (USA).

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POLYSTICHUM

Distribution. Terrestrial in pine-oak woods, in pine duff, among rocks; 1500–1750 m. Se Canada, e USA (west to Minn, Iowa, Kan, and e Tex); Mexico; naturalized in Europe. Specimens Examined. NL (Dorr 2596, ANSM, MEXU, UC, UNL; Favela 164, 167, ENCB, s.n., IEB; Lyonnet 430800003, 441200004, IEB, MEXU; Pringle 2604, VT not seen, photo US; Rodriguez Lo´pez 47, UNL, 197, IEB, UNL). Tam (Gonza´lez M. 17489, MEXU).

This is closely related to, and can be confused in Mexico only with, P. munitum. It differs from that especially in the hemidimorphic blades, with the distal pinnae fertile and the proximal pinnae sterile. The stipe and rachis scales are also much lighter in color.

2. POLYSTICHUM ALFAROI Polystichum alfaroi (Christ) Barrington, Rhodora 94: 328. 1992. Figs. 248G, H, J. Polystichum aculeatum (L.) Roth var. alfaroi Christ, Bull. Herb. Boissier, se´r. 2, 4: 963. 1904 [“alfarii”]. Type. Costa Rica. Alajuelito, 1300 m, Alfaro 16471 (P–photo US!; isotype US).

Caudices 3–6 cm diam.; fronds mostly 110–150 cm long; stipes 40–50 cm long, 4–6 mm diam., ca. 1⁄2–2⁄3 the blade length; stipe bases with ovate scales 22 ⫻ 6 mm, entire to denticulate, some scales bicolorous, tan with a broad, dark brown medial stripe, scales concolorous and light brown on distal portions of stipes, margins entire to denticulate; blades subcoriaceous, 60–90+ ⫻ 25–35 cm, broadest at the bases or nearly so, attenuate at apices; rachises lacking buds, with hair-like scales to 5 mm long; pinnae 1–2-pinnate, ca. 30–35 pairs, to 20 ⫻ 2.5–3 cm, pinnules crenate with a deltate, acroscopic auricle, or with one or more pairs of free or nearly free lobes, margins of ultimate segments and segment tips spinulose; indument abaxially along costae and costules of fibrillose scales; sori indusiate, not confluent at maturity, indusia tan, 0.6–8 mm diam.; 2n⫽82 (CR).

Distribution. Terrestrial in moist montane forests; ca. 1000– 1400 m. Mexico; Guat, Salv (also Standley 21567, NY, not previously cited by Barrington in Davidse et al., 1995), CR, Pan. The low elevation approximates the elevation of Finca Irlanda, where four of the five collections from Chiapas have been made. Specimens Examined. Chis (Breedlove 56956, CAS; Purpus 7231, 7232, UC, US, 7243, GH, MO, NY, US, 7243A, BM).

Specimens of this species were cited by Smith (1981) under “Unnamed collections,” not matching any other species from Chiapas. It is one of the more distinctive, but rare, polystichums in Mexico, and has been recollected only once in more than 75 years. All the collections are from the Sierra Madre. Distinguishing characters, especially from P. hartwegii, include the rather large ovate stipe base scales, some scales tan and to 6 mm wide, the 2-pinnate-pinnatifid to nearly 3-pinnate blades with ascending pinnules, the thick blade texture with shiny adaxial surfaces, and the moderately sized tan indusia. Specimens of P. hartwegii from the slopes of volcanoes in western Mexico and Isla Socorro seem particularly close to P. alfaroi in blade size and cutting, and differ primarily in the narrower rhizome scales with darker colored margins. Barrington (in Davidse et al., 1995) cited Purpus 7231, 7232, and 7243A, all at US, as representing the hybrid P. alfaroi ⫻ platyphyllum, but we see no evidence of hybridity in these collections. Earlier, in elevating this taxon to species rank, Barrington (1992) cited these collections as typical P. alfaroi.

3. POLYSTICHUM DISTANS Polystichum distans E. Fourn., Mexic. Pl. 1: 91. 1872. Type. Mexico. [Veracruz:] Orizaba, Schaffner 217a (not found at P or K). Figs. 250G, H, J, K. Polystichum pallidum E. Fourn., Mexic. Pl. 1: 91. 1872, hom. illeg., non Gardner, 1842, nec Todaro, 1866. Type. Mexico. Oaxaca: Ghiesbreght s.n. (P; photo US!).

Caudices 3–8 cm diam., infrequently collected because of their massive size; fronds mostly 50–150 cm long; stipes 10–30(–40) cm long, 3–6 mm diam., ca. 1⁄4 to nearly equaling the blade length;

POLYSTICHUM ERYTHROSORUM

stipe bases with dense, ovate-lanceolate scales mostly 10–30 ⫻ 2– 6(–8) mm, scales mostly blackish, lustrous, with very narrow brownish margins, or scales dark brown, margins denticulate to setose and with long, flaccid cilia, distally the stipes sometimes also copiously scaly; blades subcoriaceous, 25–65(–85) ⫻ 9–30(– 35) cm, broadest at the bases or proximal 1–3 pairs of pinnae slightly reduced, attenuate at apices; rachises lacking buds, with lanceolate and hair-like scales mostly 3–6 mm long, denticulate or ciliate, a few scales often blackish or with a black central stripe; pinnae 1–2-pinnate, mostly 25–40 pairs, mostly 5–15(–18) ⫻ (1– )1.5–2(–2.5) cm, pinnules entire to crenate with a small acroscopic auricle, less often with larger pinnules shallowly to deeply pinnatifid at the bases, infrequently each with a Ⳳ free obovate lobe, both margins and tips spinulose and often slightly revolute; indument abaxially along costae, costules, and veins of lanceolate to fibrillose scales, the larger (broader) ones denticulate or spinulose; sori indusiate to exindusiate, often confluent at maturity, indusia tan to brownish, usually 0.3–0.8 mm diam., peltate, often fugacious; 2n⫽82 (Chis.).

Distribution. Terrestrial in wet pine-oak forests, disturbed woods, moist ravines; 1300–2900 m. Mexico; Guat, Hond, Salv. Selected Specimens Examined. Chis (Breedlove 53865, CAS). Gro (Lorea 4693, FCME, IEB). Hgo (Gimate L. 958, ARIZ, ASU, CAS, IEB, NY; *Pringle 13638, ARIZ, ASU, CAS, UC; *Rzedowski 32683, IEB, MEXU; *Sa´nchez M. 226, MEXU). Jal (Judziewicz & Cochrane 4968, UC). Me´x (Rzedowski 30905, ENCB, IEB, NY). Mich (Escobedo 408, IEB). Mor (Pimentel B. 20, CAS). Oax (Barrington 1731, UC). Pue (*Bashor 1350, UC; Ventura A. 471, ENCB, IEB, MEXU). Qro (*Servı´n O. 1013, IEB, UC). Tam (Lof 787, UC). Tlax (West Q-22, US). Ver (Bartholomew 3359, CAS, UC; *Cochrane 8574, CAS, MEXU, UC, XAL; *Zamora P-1034, UC).

The differences between this species and P. rachichlaena are few and possibly not absolute, and it may be that the two should be considered variants of a single, polymorphic species. In general, P. distans has blacker, glossier, or darker-centered scales that run up the stipes for some distance. Polystichum distans also has more dissected pinnules that tend to be less rounded at tips, harsher blade texture, and generally smaller indusia (or indusia absent). All these characteristics vary considerably, especially in and around the Valle de Me´xico and in the western Sierra Madre.

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Detailed morphometric, cytological, and isozymic work is needed to establish whether the differences are constant or not. Several collections from Veracruz (e.g., LaFrankie 1076, TEX; Narave F. 660, ENCB; Cochrane 8574, CAS, MEXU, UC; Zamora P-1034, UC) and Puebla are remarkable for their abundantly scaly stipes and rachises, very large dark brown to black scales to 3 ⫻ 0.8 cm, scales prominently spinulose at tips, 2pinnate-pinnatifid to subtripinnate blades, and exindusiate sori. However, some collections more typical of P. distans, lacking the abundant large stipe scales, are also apparently exindusiate (e.g., Pringle 13638, from Hidalgo). Exindusiate collections seen are noted with an asterisk (*) in the citations. Mickel 3728 (UC), considered a hybrid between P. hartwegii and P. ordinatum by Mickel and Beitel (1988: 313), is typical P. distans; the spores and sporangia are well formed and appear normal. Some specimens of this species (e.g., Bartholomew 3359, UC; Ferna´ndez N. 509, IEB) have very large spores, and one suspects that they are polyploids. However, the only reported count for this species is diploid. Most collections from Hidalgo differ in their narrower, fewer, and less blackish stipe base scales; they also grow at lower elevations than typical for the species. The differences between these collections and specimens of P. hartwegii, often found at lower elevations than P. distans, from the same area are subtle and not altogether clear. Even more puzzling is the sole collection from Tamaulipas, from 1400 m; it has brown stipe base scales, and might with equal justification be called P. hartwegii. A specimen from Veracruz (Ventura A. 5037, XAL) is peculiar in having numerous (ca. 35) gall-like structures (viral or bacterial in origin?) produced from the receptacle. This number at ENCB lacks such galls (Barrington, in litt.). These are 2–5 mm diam., round in outline, flattened, solid (without noticeable cavities or insect larvae within), and about 1 mm thick. Remarkably, there is a single perfectly normal indusium borne in the center of each gall, and some abortive sporangia produced beneath the galls. The developmental processes that would allow normal indusia to be produced, while suppressing sporangial development cry out for explanation. A specimen of P. hartwegii from Morelos (Lyonnet 540900012, CAS) has some similar gall-like structures.

4. POLYSTICHUM ERYTHROSORUM Polystichum erythrosorum A. R. Sm., Proc. Calif. Acad. Sci., ser. 4, 40: 225. 1975. Type. Mexico. Chiapas: Cerro Mozotal, Breedlove 22709 (DS!). Figs. 249D–G. Caudices 2–6 cm diam.; fronds mostly (60–)110–150 cm long; stipes 30–65 cm long, (3–)5–9 mm diam., ca. 1⁄2–2⁄3 the blade length; stipe bases with ovate-lanceolate scales, larger scales 10– 20 ⫻ 4–6 mm, bicolorous, with a broad, dark brown or black medial band and narrow tan margins, margins ciliate, beneath these with smaller decidedly ciliate scurfy scales; blades subcoriaceous, (34–)50–70 ⫻ (14–)20–40 cm, broadest at the bases or nearly so, attenuate at apices; rachises lacking buds, with ovatelanceolate and hair-like scales mostly 2–9 mm long, a few scales

516

POLYSTICHUM

blackish; pinnae pinnate, mostly 20–30 pairs, (7–)11–26 ⫻ (1.6– )2.5–4 cm, pinnules crenate with an oblong, acroscopic auricle, or larger pinnules sometimes pinnatifid and with a Ⳳ free obovate lobe at bases, both margins and tips spinulose; indument abaxially along costae and costules of fibrillose scales, also with ovate-lanceolate scales along the costae, these denticulate or spinulose; sori indusiate, not confluent at maturity, indusia reddish brown, 0.5–0.8 mm diam.; 2n⫽82 (Chis).

Caudices 1–2 cm diam.; fronds mostly (25–)35–65 cm long; stipes 9–26(–36) cm long, 1–3 mm diam., ca. 2⁄3 to nearly equaling the blade length; stipe bases with lanceate scales, 5–10 ⫻ 1–2 mm, concolorous or bicolorous, stramineous to tan or orange-brown, entire to denticulate or short-ciliate, sometimes with a darker brown medial stripe, scales concolorous and pale brown distally; blades chartaceous, 15–36(–45) ⫻ 6–15(–20) cm, broadest at bases or nearly so, attenuate at apices; rachises lacking buds, with tan, often twisted, hair-like scales 1–3 mm long, these often ciliate or denticulate, especially at the broadened bases; pinnae pinnate, ca. (15–)20–25 pairs, 3–10 ⫻ (0.8–)1–2(–2.5) cm, pinnules crenate or less often shallowly pinnatifid, spinulose on both sides, the basal acroscopic segments often more deeply pinnatifid; indument abaxially along costae, costules, and veins of fibrillose scales; sori indusiate, not confluent at maturity, indusia whitish to tan, (0.6–)1–1.5 mm diam., often overlapping in young fronds; 2n⫽82, 164.

Distribution. Terrestrial in wet forests; 2300–3000 m. Mexico; Guat, Hond. Selected Specimens Examined. Chis (Breedlove 68902, DS, 22047, DS, MEXU, 53590, 53595, CAS; Mu¨nch 113, US). Oax (Barrington 1754, UC; Breedlove 65928, CAS; Mickel 4195, UC, 5280, NY, UC, 5281, UC). Unverified, Doubtful, or Mistaken Reports. Me´x (Tejero-Dı´ez 3012, IZTA, cited by Tejero-Dı´ez & Arreguı´n-Sa´nchez, 2004, but the specimen is P. rachichlaena).

Polystichum erythrosorum is distinguished from P. rachichlaena by having reddish brown indusia 0.5–0.8 mm diam. (rather than pale brown and 1–1.5 mm diam.), stipe base scales bicolorous with lustrous, dark brown to often black central band and thin, pale brown, ciliate margins (vs. scales concolorous with distantly spinulose margins and twisted tips), and twisted hair-like scales 1–1.5 mm on the veins of the abaxial laminar surfaces (vs. straight hair-like scales 0.5–1 mm with reddish brown cross walls). One can often find some blackish scales along the rachis, and in this character P. erythrosorum can be confused with P. distans, which generally has more dissected and more strongly spinulose pinnules, more coriaceous blades, and smaller indusia (or exindusiate). The small reddish brown indusia and relatively broad, lustrous, black-centered bicolorous scales will separate P. erythrosorum from P. fournieri.

Distribution. Terrestrial in wet pine-oak forests, cloud forests; (1200–)1700–3000 m; Mexico; Guat, Salv, CR. Cited also from Honduras and Panama by Barrington (in Davidse et al., 1995). Selected Specimens Examined. Chis (Breedlove 34694, DS, NY, 41745, DS, MEXU). Gro (Lorea 1948, FCME, IEB, US, 2273, FCME, IEB, NY; Mexia 9066, UC, US). Me´x (Tejero Dı´ez 2227, IZTA). Mor (Lyonnet 767, MEXU, NY, US). Oax Herna´ndez G. 314, MEXU, NY; Pringle 5603, ARIZ, CAS, MEXU, NY, UC, US). SLP (Virlet d’Aost 46, P–photo US). Ver (Mu¨ller 129, NY).

Polystichum fournieri is similar to P. hartwegii but is smaller in size (fewer than 25 pinna pairs, fewer than 20 pinnules per pinna, pinnae 3–10 cm long), with smaller denticulate-margined rhizome scales mostly 5–10 ⫻ 1–2 mm, proximal pinnae deflexed, and larger whitish indusia mostly 1–1.5 mm. See P. erythrosorum for differences with that species.

5. POLYSTICHUM FOURNIERI Polystichum fournieri A. R. Sm., Amer. Fern J. 70: 27. 1980. Figs. 249M–P. Polystichum muelleri Mett. ex E. Fourn., Mexic. Pl. 1: 91. 1872, hom. illeg., non Schumacher, 1803). Lectotype (chosen by Smith, 1981: 197). Mexico. Chiapas: Linden s.n. (P!–photo US!).

6. POLYSTICHUM FURFURACEUM Polystichum furfuraceum A. R. Sm., Proc. Calif. Acad. Sci., ser. 4, 40: 226, f. 6A–D. 1975. Type. Mexico. Chiapas: Zontehuitz, Mu¨nch 113 (DS!; isotypes DS!, US!). Figs. 248C–F.

POLYSTICHUM HARTWEGII

Caudices not known, probably massive; fronds ca. 100 cm long; stipes to ca. 30 cm long, 3–5 mm diam., ca. 1⁄2 the blade length, nearly obscured by very dense ferrugineous to brown, concolorous scales; stipe bases with broadly ovate to narrowly lanceolate scales, scales concolorous or slightly darkened towards scale bases, ferrugineous to brown, the largest to 18 ⫻ 9 mm, margins prominently setiform, teeth to 0.4 mm; blades subcoriaceous to coriaceous, to 70 ⫻ 40 cm, broadest at the bases or proximal pinnae slightly shortened and deflexed, attenuate at apices; rachises lacking buds, densely and persistently scaly like the stipes, scales lanceate, to ca. 10 ⫻ 2 mm, setiform along margins; pinnae pinnate, approximate to somewhat imbricate, to 45 pairs, to 20 ⫻ 2 cm, pinnules Ⳳ entire, each with a small, acroscopic auricle, basiscopic margins entire or spinulose toward pinnule tips; indument abaxially on costae, costules, and veins of narrowly lanceolate to fibrillose scales throughout; sori indusiate, indusia brownish, 0.4–0.7 mm diam.

Distribution. Terrestrial in cloud forests; 2600–2800 m. Mexico; Guat. Specimens Examined. Chis (Breedlove 31807, DS). Gro (Lorea 2293, 2379, FCME).

This species has close affinities only to P. erythrosorum which has larger, more reddish indusia and less scaly stipes, rachises, and costae. The densely scaly stipes and rachises, setiformmargined scales, and unlobed but crowded pinnules will distinguish P. furfuraceum from other Mexican polystichums.

7. POLYSTICHUM HARTWEGII Polystichum hartwegii (Klotzsch) Hieron., Hedwigia 46: 355. 1907 [“hartwigii”]. Figs. 249A–C, 250P–S. Aspidium hartwegii Klotzsch, Linnaea 20: 366. 1847. Type. Guatemala. “In montibus Las Nubes, prope Urbam” [near Guatemala City], Hartweg 631 (B!; isotypes BM!, K!, P!–2 sheets). Polystichum grande Fe´e, Me´m. Foug. 8: 98. 1857. Type. Mexico. Veracruz: Huatusco, Schaffner 217 (not found at P; probable isotype K labeled “Orizaba”). Polystichum drepanoides E. Fourn., Mexic. Pl. 1: 92. 1872. Type. Mexico. Chiapas: “inter San Bartolo et Pueblo-Novo,” Linden s.n. (P! photos US!). Aspidium trejoi Christ, Bull. Herb. Boissier, se´r. 2, 5: 255. 1905. Polystichum

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trejoi (Christ) Christ, Bull. Herb. Boissier, se´r. 2, 5: 728. 1905. Lectotype (chosen by Smith, 1981: 197). Mexico. Chiapas: San Pablo, Mu¨nch 167 (not found at P; isolectotype DS!).

Caudices mostly 2–4(–6) cm diam.; fronds mostly (60–)80– 140 cm long; stipes 20–45 cm long, 3–7 mm diam., ca. 1⁄3–1⁄2 the blade length; stipe bases with lanceate scales 8–15(–20) ⫻ 1–3(–5) mm, entire to denticulate or sparsely ciliate, concolorous and brown to bicolorous and with a broad, dark brown or blackish medial band and tan margins; blades chartaceous to subcoriaceous, 35–90 ⫻ (15–)25–55 cm, broadest at the bases or nearly so, attenuate at apices; rachises lacking buds, with brownish to tan, lanceolate and mostly hair-like scales; pinnae pinnate or often pinnate-pinnatifid, mostly 25–30 pairs, 10–25(–30) ⫻ 2–3.5(–5) cm, pinnules often pinnatifid or with a Ⳳ free obovate lobe at bases, both margins and tips spinulose, margins not revolute; indument abaxially along costae and costules of fibrillose scales; sori indusiate, confluent at maturity or not, indusia tan to brown (whitish when young), mostly 0.8–1.5 mm diam.; 2n⫽82 (Chis).

Distribution. Terrestrial in wet forests; (700–)1200–2500 m. Mexico; Guat, Hond, Salv, CR, Pan; Col?, Ven, Ec, Bol. Also cited from Nicaragua by Barrington (in Davidse et al., 1995). Selected Specimens Examined. Chis (Purpus 6727, BM, NY, UC, US). Col (Sanders 10477, MEXU, NY, TEX). Gro (Miller & Campos 2925, MEXU, UC). Hgo (Ramı´rez & Riba 661, XAL). Jal (Mexia 1333, CAS, UC, US). Me´x (Dunn & Dunn 18963, NY). Mich (King & Soderstrom 5075, UC, US). Mor (Lyonnet 540900012, CAS, bearing gall-like structures; see discussion under P. distans). Nay (Yatskievych 86-258, MO, UC). NL (Hinton 22654, TEX). Oax (Barrington 1742, UC). Pue (Riba et al. 275, MEXU, NY). Rev (Felger 15839, ENCB, MEXU, UC, US). Sin (Correll & Correll 28867, MEXU). Ver (Nee & Taylor 25949, NY, UC).

This species and P. rachichlaena are perhaps the most common polystichums in Mexico. The deeply incised pinnules, weakly bicolorous scales 8–15(–20) mm long with ciliate margins, slightly reduced proximal pinna pair, and relatively large indusia mostly 0.8–1.5 mm diam. separate P. hartwegii from P. rachichlaena, P. erythrosorum, P. distans, and P. ordinatum. Polystichum turrialbae differs in having smaller indusia and pale brown, ovate rhizome scales with erose-denticulate margins. See P. fournieri for discussion of differences with that species. Polystichum drepanoides has been distinguished (e.g., by

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POLYSTICHUM

Smith, 1981) primarily by the larger and broader pinnae (to 30 ⫻ 3–5 cm) and entire indusia. We no longer consider these differences taxonomically significant and choose to synonymize this name under P. hartwegii. Many specimens, especially those from western Mexico, are slightly dimorphic, with the fertile pinnae narrower and with smaller pinnules than sterile pinnae. In general, in P. hartwegii and perhaps in other Mexican Polystichums, the fertile blades are more dissected than sterile blades on a given plant, but there is considerable variation in blade size and dissection. Plants from Colima and the Revillagigedo Islands (Isla Socorro, as well as some from the states of Sinaloa, Nayarit, and Jalisco) are the most aberrant of all and are placed here tentatively. They grow at the lowest elevation for the species (700 m on Socorro). Some specimens are nearly 3-pinnate. They have dark brown, concolorous stipe base scales but are otherwise similar to P. distans; they also often have rather glandular indusia. Vasquez V. 540 (NY, UC), from Colima, and McVaugh & Koelz 1004 (IEB, NY), from Jalisco, have irregularly shaped spores that may indicate hybridity. Specimens from Guerrero (e.g., Martı´nez S. 5068, ENCB, XAL; Rzedowski & McVaugh 30, MEXU, NY) tend to have rather sharply bicolorous scales (dark brown medial stripe, tan margins) and very large pinnae, to 28 ⫻ 5 cm; they have often been named P. drepanoides (e.g., by Lorea-Herna´ndez & Vela´zquez M., 1998). Two specimens from Oaxaca, Mickel 3728, 4678 (NY), are intermediate between P. hartwegii and P. ordinatum, and may represent hybrids. Ortega & Calzada 778 (XAL) has irregularly formed spores but looks reasonably typical for P. hartwegii. Polystichum hartwegii is recorded from Nuevo Leo´n by Hinton and Hinton (1995), but the specimen cited (Hinton 22654, TEX) needs verification.

scales, especially at bases of costules; sori exindusiate, not confluent at maturity.

Distribution. Terrestrial in wet forest; (450–)700–2500 m. Mexico; Guat, Hond. Barrington (in Davidse et al., 1995) also cited a collection from El Salvador. Selected Specimens Examined. Chis (Breedlove 34385, DS, 55089, CAS, ENCB, 69015, CAS). Oax (Campos V. 4095, MEXU; Croat 39769, UC; Mickel 1565, US; Torres C. 11636, MEXU). Ver (Riba 529, MEXU; Ventura A. 3431, ENCB, MEXU, 4812, ENCB, MEXU, NY).

This exindusiate species has distinctive, bicolorous scales with a dark brown central stripe and erose-denticulate margins. The only other exindusiate species in Mexico are P. platyphyllum, P. speciosissimum, P. orbiculatum, and some specimens of P. distans, none of which is closely related. The closest relatives of P. mickelii may be in South America, perhaps species of the group of P. montevidense (Spreng.) Rosenst. A peculiarity of P. mickelii is that the stomates on the abaxial surfaces of the blades appear darker green and contrast readily with the surrounding epidermal cells.

8. POLYSTICHUM MICKELII Polystichum mickelii A. R. Sm., Amer. Fern J. 70: 25. 1980. Type. Mexico. Oaxaca: Distrito Mixe, NE slope of Cerro Zempoalte´petl, trail from La Candelaria to Zacatepec, Mickel 4836 (NY!). Figs. 249H, J–L, 250L–O. Caudices 1.5–3 cm diam.; fronds mostly 60–110 cm long; stipes 17–50 cm long, 3–5 mm diam., ca. 1⁄2–2⁄3 (to nearly equaling) the blade length; stipe bases with ovate scales 8–18 ⫻ 2–6 mm, erose-denticulate towards tips, mixed with hair-like, denticulate scales, larger scales bicolorous, tan with a narrow, dark brown medial stripe, scales concolorous and light brown distally; blades coriaceous, 30–60 ⫻ 12–35 cm, broadest at the bases or nearly so, attenuate at apices; rachises lacking buds, with hair-like scales 1–2 mm long, these most numerous at pinna bases; pinnae pinnate, ca. 25–30 pairs, to 20 ⫻ 4 cm, pinnules Ⳳ entire with a small, deltate, acroscopic auricle, or on larger fronds with a Ⳳ free elliptic lobe at pinnule bases and beyond that shallowly lobed to coarsely dentate, basiscopic margins entire to minutely spinulose, crenulate, or shallowly lobed with lobes strongly ascending; indument abaxially on costae and costules of fibrillose

9. POLYSTICHUM MUENCHII Polystichum muenchii (Christ) C. Chr., Index Filic. 585. 1906. Figs. 247C–E. Aspidium muenchii Christ, Bull. Herb. Boissier, se´r. 2, 1: 1120. 1901 [“mu¨nchii”]. Type. Mexico. Chiapas: La Mesilla, near San Cristo´bal Las Casas, Mu¨nch 24 (P!–photo US!; isotypes DS!, US!).

Caudices ca. 1–2 cm diam.; fronds 20–45 cm long; stipes 3–15 cm long, 1–2 mm diam., ca. 1⁄4–1⁄2 the blade length; stipe bases with numerous dull brown to tan, concolorous, fibrillose scales 3–8 ⫻ 0.5–1 mm, entire; blades chartaceous, to ca. 30 ⫻ 3–5 cm, broadest at the bases, attenuate at the pinnatifid apices; rachises lacking buds, sparsely scaly with hair-like scales 1–2 mm long; pinnae undivided, all about the same size except for a few reduced distal ones, ca. 10–22 pairs, 1.5–2.5 ⫻ 0.8–1.5 cm, inequilaterally oblong each with a deltate, acroscopic auricle, otherwise obscurely to coarsely crenulate to dentate along margins, basiscopic base excised, tips acute and spinulose; indument abaxially along costae of a few fibrillose scales to glabrous (or glabrescent),

POLYSTICHUM MURICATUM

blade surfaces glabrous; sori indusiate, not confluent at maturity, indusia brown, ca. 1 mm diam., subpersistent.

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a pronounced acroscopic, deltate auricle 3–5 mm long, otherwise serrate-spinulose along both margins, basiscopic bases rounded to cuneate, tips acute and spinulose; indument abaxially along costae and veins of numerous lanceolate scales with contorted bases; sori indusiate, confluent or discrete at maturity, indusia tan to brown with darker brown centers, ca. 1 mm diam., subpersistent; 2n⫽82 (USA).

Distribution. Habitat unknown, possibly epipetric on limestone; ca. 2500 m. Mexico; Guat. Specimens Examined. Chis (Mu¨nch 14, DS, P-photo US).

As indicated by Smith (1981) and Stolze (1981), this species is most closely related to P. echinatum (J. F. Gmel.) C. Chr., from the Greater Antilles (Mickel, 1997), and differs from that in the pinnae being less numerous and less crowded, the non-spinulose pinna margins, and the stipe bases and rhizome apex scales mostly filiform (some broader scales in P. echinatum). The only other 1-pinnate species in Mexico are P. munitum and P. acrostichoides, both from northern Mexico. These two species are outliers of populations of their species in western and eastern United States and Canada, respectively, and both have much longer, more linear-lanceate pinnae, more broadly deltate blades, and broader rhizome apices and stipe scales than P. echinatum.

10. POLYSTICHUM MUNITUM Polystichum munitum (Kaulf.) C. Presl, Tent. Pterid. 83. 1836. Fig. 247B. Aspidium munitum Kaulf., Enum. Filic. 236. 1824. Type. U.S.A. California: Vicinity of the Presidio, San Francisco, 1816, Chamisso s.n. (LE?; isotype B). Polystichum munitum (Kaulf.) C. Presl subsp. solitarium Maxon, Fern Bull. 11: 39. 1903. Polystichum solitarium (Maxon) Underw. ex Maxon, Contr. U.S. Natl. Herb. 10: 493. 1908. Type. Mexico. Guadalupe Island: Anthony s.n. (GH!; isotypes DS, NY, US).

Caudices ca. 3 cm diam.; fronds mostly 60–90 cm long; stipes to ca. 30 cm long, 3–4 mm diam., ca. 1⁄3–1⁄2 the blade length; stipe bases and also distal part of stipes with numerous concolorous, brown to red-brown, ovate-lanceolate scales 10–20 ⫻ 2– 5 mm, scales entire to sparingly denticulate or rarely ciliate along margins; blades monomorphic, chartaceous, to ca. 60 ⫻ 15 cm, broadest at the bases, attenuate at the pinnatifid apices; rachises lacking buds, densely scaly with ovate-lanceolate to narrowly lanceolate, denticulate scales mostly 3–7 ⫻ 0.4–0.6 mm; pinnae undivided, very gradually shortened distally, to ca. 50 pairs, mostly 6–10 ⫻ 0.8–1 cm (excluding basal auricle), lanceate with

Distribution. Terrestrial in wooded areas; low elevations. Canada, w USA (Alaska to s Calif, east to Mont and n Idaho, disjunct in S Dak); Mexico; naturalized in Europe. Specimens Examined. Guad (Anthony s.n., NY; Palmer 102, GH).

This differs from typical P. munitum from western North America in the very densely scaly rachis and narrow pinnae, prompting Maxon to regard it as a species in 1908. However, similar specimens can be found from southern California and the Channel Islands, especially Santa Cruz Island. See P. acrostichoides for differences with that species, the only other similar one in Mexico.

11. POLYSTICHUM MURICATUM Polystichum muricatum (L.) Fe´e, Me´m. Foug. 5: 278. 1852. Figs. 248A, B. Polypodium muricatum L., Sp. Pl. 2: 1093. 1753. Type. Petiver, Pterigraphia Amer., pl. 1, f. 6, 1712, which is a small copy of Plumier’s Traite´ Foug. Ame´r., pl. 39, 1705, illustrating a plant from Hispaniola.

Caudices ca. 2 cm diam. or less; fronds mostly (30–)50–75 cm long; stipes 15–25 cm long, ca. 3 mm diam., ca. 1⁄2–2⁄3 the blade length; stipe bases with concolorous, tan to brown, ovate scales to 15 ⫻ 6 mm, these entire, lacking a distinct medial stripe but sometimes with a broad, darkened central band, scales of the distal part of stipes light brown; blades herbaceous to chartaceous, ca. 35–50 ⫻ 12–25 cm, broadest at the bases or proximal pair of pinnae only slightly shorter than the next pair, attenuate at apices; rachises lacking buds, with tan hair-like eciliate scales to 6 mm long; pinnae pinnate to pinnate-pinnatifid, ca. 20–25 pairs, 8–12 ⫻ 1.8–2.3 cm, pinnules crenate, each with a deltate, acroscopic auricle, or some pinnules with a Ⳳ free obovate lobe at base and beyond that shallowly lobed to coarsely dentate, both

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margins and tips spinulose; indument abaxially along costae and costules of fibrillose scales; sori indusiate, not confluent at maturity, indusia tan, relatively small, 0.2–0.4 mm diam.; 2n⫽82 (Chis).

Distribution. Terrestrial in montane rain forests; 1350–1850 m. Mexico; Guat, Hond, Salv, Nic, CR, Pan; Jam, Hisp; Ven. Barrington (in Davidse et al., 1995: 223) also cited a specimen from El Salvador (Villacorta et al. 906, MO!) and included Galapagos in the range. In our opinion, the Salvadorean specimen is doubtfully this species.

Nephrodium polyphyllum C. Presl, Reliq. Haenk. 1(1): 37. 1825. Polystichum polyphyllum (C. Presl) C. Presl, Suppl. Tent. Pterid. 83. 1836. Type. Peru. Haenke s.n. (PR or PRC; photo BM or PR).

Caudices mostly (1–)2–5 cm diam.; fronds to ca. 45 cm long; stipes (8–)12–22 cm long, 1–3 mm diam., 1⁄3 to nearly equaling the blade length; stipe bases with concolorous, brownish to tan or orangish, ovate-to lanceolate scales, larger ones to 15 ⫻ 3(–5) mm, ciliate and also denticulate, the scales similar to but paler than those of rhizome apices, distally with similar but fewer scales, but still rather densely scaly; blades subcoriaceous to coriaceous, 15–30 ⫻ (2–)3–6 cm, broadest at bases or nearly so, very gradually reduced at apices; rachises moderately to often densely scaly, scales lanceolate, ciliate and denticulate, to ca. 5 mm long, lacking buds; pinnae pinnate, ca. 25–40 pairs, often strongly ascending and imbricate, mostly 1.5–3(–4) ⫻ 1–1.5 cm, pinnules strongly revolute, rounded to oblong or obovate, non-auriculate, not aristate along margins and only faintly so at tips; indument abaxially along costae, costules, and veins of scattered, hair-like to lanceolate scales; sori exindusiate, often confluent at maturity; 2n⫽164 (CR).

Specimens Examined. Chis (Breedlove 21808, DS, 24927, 28923, 31316, 32514, DS, MEXU, 32708, DS; Purpus 10155, UC). Oax (Maya J. 1103, CHAPA, NY; Wendt et al. 5170, CHAPA, MEXU, NY). Unverified, Doubtful, or Mistaken Reports. Rev (Mason 1630, 1677, and Barkelew 227, cited in the range by Levin & Moran, 1989: 22, but these specimens are here regarded as aberrant P. hartwegii).

Polystichum muricatum seems to be found at lower elevations than most other Polystichums in southern Mexico except P. platyphyllum and P. alfaroi. It is distinguished by its broad, often tan, concolorous, entire scales at the stipe bases, 2-pinnatepinnatifid blades, fibrillose rachis scales, thin blade texture, and relatively small indusia. The species with which it is most often confused in Mexico include P. hartwegii (which is generally larger, thicker-textured, and with narrower, often bicolorous stipe base scales), P. ordinatum (which has narrower blades, several pairs of reduced proximal pinnae, and narrower stipe base scales with greatly thickened bases), and P. fournieri (which is generally found at higher elevations and has narrower, usually bicolorous stipe base scales and much larger indusia).

12. POLYSTICHUM ORBICULATUM

Distribution. Terrestrial, flat meadows and steep slopes with pine, juniper, and Buddleia; 3600–4200 m. Mexico; Guat, CR; Col, Ven, Ec, Peru, Bol. Specimens Examined. Chis (Breedlove 24306, DS, MEXU, NY, 24319, 26705, DS, MEXU, 29377, DS, MEXU, NY; MacDougall s.n., US; Matuda 2352, MEXU, NY, US).

This high-elevation species is extremely polymorphic, especially in South America, and is known to hybridize with others in Costa Rica. There is the likelihood that the species in the sense recognized here and in other recent floras (Mesoamerica, Peru) is really a species complex. Tryon and Stolze (1991) recognized two varieties, but the sole variety in Mexico and Central America is var. orbiculatum. It differs from P. speciosissimum, the closest relative in Mexico, by the less dissected and narrower blades, lack of modified whitish segment margins, and by the proximal pinnae not or only very slightly reduced.

Polystichum orbiculatum (Desv.) J. Re´my in Gay, Hist. Fis. Pol. Chile, Bot. (Fl. Chilena) 6: 515. 1853. Figs. 248K–N.

13. POLYSTICHUM ORDINATUM

Aspidium orbiculatum Desv., Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck. Gesammten Naturk. 5: 321. 1811. Type. “America australi.” Collector not known (P, Herb. Desv.; photos GH, US).

Polystichum ordinatum (Kunze) Liebm., Mexic. Bregn. 276 (reprint 124). 1849. Figs. 249Q–T.

POLYSTICHUM PLATYPHYLLUM Aspidium ordinatum Kunze, Linnaea 18: 347. 1844. Type. Mexico. Leibold s.n. [Leib. Verz. No. 39] (LZ, destroyed; presumed isotypes B!, K!, P!–6 sheets, NY!; frag. US!). Polystichum bicolor A. R. Sm., Proc. Calif. Acad. Sci., ser. 4, 40: 223, f. 6E– G. 1975. Type. Mexico. Chiapas: Mpio. Tenejapa, Colonia ‘Ach’lum, Ton 1987 (DS!; isotype MEXU!).

Caudices ca. 2–4 cm diam.; fronds mostly 45–150+ cm long; stipes 10–35(–50) cm long, 2–8(–12) mm diam., ca. 1⁄4 to nearly equaling the blade length; stipe bases with lanceate, ascending scales 7–12 ⫻ 1–2(–3) mm, scales entire to sparsely ciliate, thickened at bases, usually discordantly bicolorous, with a narrow, dark brown, thickened central band and lighter tan margins, scales of the distal part of stipes concolorous and light brown; blades thin-chartaceous, ca. 30–60(–90) ⫻ 10–28 cm, proximal 2–5(–8) pairs of pinnae reduced to ca. 0.3–0.8 the length of the longest pinnae, attenuate at apices; rachises lacking buds, with tan hair-like eciliate scales to 5 mm long; pinnae pinnatepinnatisect, ca. 20–25(–36) pairs, 5–20 ⫻ 1.5–2.5 cm, pinnules serrately incised to pinnatifid, some pinnules with a Ⳳ free obovate lobe at base, both margins and tips spinulose; indument abaxially along costae and costules of fibrillose scales; sori indusiate, not confluent at maturity, indusia tan to brownish, relatively small, apparently reniform or reniform-spatulate, fugacious, 0.2–0.5 mm diam.

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closest relative appears to be P. hartwegii, and some specimens are difficult to place in one or the other species. The stipe base scales are sometimes abraded with age, and show clearly the stoutness of the scale at the point of attachment, leaving the stipes almost muricate. As stated earlier (Smith, 1981), the name P. ordinatum has been too widely applied, and we are still uncertain that we have applied it correctly to the taxon circumscribed herein. The type and most collections cited here have several pairs of shortened proximal pinnae, the lowermost about half or less the length of the longest ones. Other Mexican polystichums may have the proximal pinnae slightly reduced, but generally only a single pair, and not shortened to the extent found in P. ordinatum. Polystichum ordinatum is also quite variable, and divisible into two elements: 1) very large-fronded specimens with small, seemingly peltate indusia and thick-chartaceous to subcoriaceous blade texture; and 2) small-fronded plants with reniform indusia and thin-textured blades. The taxonomic status of these two variants needs clarification. The large-fronded variant appears to be common in southern Veracruz in the vicinity of Xalapa, but elsewhere it is rather uncommon. Mickel 4122 (UC) is a mixture of P. erythrosorum and P. ordinatum, and shows the value of mixed collections. Such gatherings call attention to “genus communities,” wherein the differences between two closely related species are maintained.

14. POLYSTICHUM PLATYPHYLLUM Polystichum platyphyllum (Willd.) C. Presl, Suppl. Tent. Pterid. 84. 1836. Figs. 250A–F. Aspidium platyphyllum Willd., Sp. Pl., ed. 4, 5(1): 255. 1810. Phegopteris platyphylla (Willd.) Mett., Abh. Senckenberg. Naturf. Ges. 2: 296. 1858. Type. Venezuela. Caracas, Bredemeyer s.n. (B-Willd. 19789).

Distribution. Terrestrial in wet forest; 1200–2800 m. Mexico; Guat. Selected Specimens Examined. Chis (Matuda 2971, K, MEXU, NY, US; Breedlove 34686, DS, MEXU). Gro (Lorea 1225, FCME, IEB; Rodrı´guez B. 108, ENCB, MEXU). Hgo (Luna et al. 798, FCME). Oax (Mickel 5311, NY; Tenorio L. 18568, MEXU). Pue (Arreguı´n 540, ARIZ, IEB, XAL; Rzedowski 29999, IEB, NY, XAL). Ver (Barrington 2038, VT, XAL; Ventura A. 5083, IEB, MEXU, NY, XAL).

Polystichum ordinatum differs from P. hartwegii, P. fournieri, and P. muricatum, all close relatives, in its lanceolate, usually rather decidedly bicolorous scales with thickened bases, reduced proximal pinnae, and small indusia mostly 0.2–0.5 mm diam. In some specimens seen, the indusia are lost early in development, and the indusia that are seen appear to be reniform or spatulate-reniform, rather than truly peltate as found in other Mexican indusiate species. This is especially evident in a series of specimens collected from near Acajete, in Edo. Veracruz. The

Caudices 1–3 cm diam.; fronds mostly (32–)50–80(–110) cm long; stipes (10–)25–50 cm long, 2–3 mm diam., half to nearly equaling the blade length; stipe bases with scales concolorous, tan to brown, ovate to lanceolate, larger ones 5–10 ⫻ 1–3 mm, margins with long, flaccid, twisted cilia, also with hair-like, denticulate scales 2–4 ⫻ 0.5–0.8 mm, distally with scattered hair-like scales; blades chartaceous, (20–)35–60(–70) ⫻ (8–)20–30 cm, broadest at the bases or nearly so, very gradually attenuate at apices and generally with many (to12 pairs) simple pinnae similar to pinnules of proximal pinnae; rachises with dense to scattered, twisted, Ⳳ entire, hair-like, eciliate scales 1–7 mm long, nearly always bearing a small scaly bud(s) at the base of a distal pinna; pinnae pinnate, ca. 25–35 pairs, (5–)10–16 ⫻ (1.5–)3–4.5 cm, pinnules planar, oblong-rhomboid, each with a small, deltate, acroscopic auricle, then crenate or crenately lobed along the acroscopic margin, basiscopic margin Ⳳ entire to crenulate, apices rounded with aristate tips; indument abaxially along costae, costules, and veins of scattered, hair-like scales; sori exindusiate, not confluent at maturity; 2n⫽82 (Chis), 164 (Parag).

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did (for other suspected examples of false labeling, see, e.g., Yatskievych, G. 1996. A revision of the fern genus Phanerophlebia (Dryopteridaceae), Ann. Missouri Bot. Gard. 83: 191, 1996; also Reineck s.n. (UC), Thelypteris puberula (Baker) C. V. Morton var. puberula, labeled from Brazil, Rio Grande do Sul, a specimen probably collected in Mexico).

15. POLYSTICHUM RACHICHLAENA Polystichum rachichlaena Fe´e, Me´m. Foug. 8: 100. 1857. Type. Mexico. Popocate´petl, 3000⫹ m, Schaffner 290 (RB, not found at P; probable isotype K!). Figs. 246A–D. Distribution. Terrestrial in wet montane forests; 750–1600 m. Mexico; Hond, Nic, CR, Pan; Jam; Col, Ven, Ec, Peru, Braz, Bol, Parag, nw Arg. Selected Specimens Examined. Chis (Breedlove 24048, DS, MEXU, 50779, CAS; Martı´nez S. 17177, XAL; Palacios-Rios 2842, UC, XAL). Oax (Herna´ndez G. 1539, CHAPA; Mickel 6166, NY, UC, 7279, NY). Ver (Purpus 6200, MO, NY, UC). Unverified, Doubtful, or Mistaken Reports. Gro (Campos 1663, FCME, cited by Lorea-Herna´ndez & Vela´zquez M., 1998, but not verified).

This exindusiate taxon bears a small bud (occasionally more than one bud) in the axil of a distal pinna, which distinguishes it from all other Mexican congeners. Its closest relatives are P. polystichiforme (Fe´e) Maxon from the Greater Antilles, and a host of South American exindusiate species, most numerous in southern Brazil. Polystichum platyphyllum is the most wide-ranging species of the genus in Mexico and is probably a species complex over its range, as evidenced by diploid and tetraploid chromosome number reports. Mu¨nch 96 (DS), from Chiapas, is problematic. Nothing like it has since been collected in Chiapas, Mexico, Mesoamerica, or the Antilles, as far as we know. The blade on the right side of this sheet resembles most closely Polystichum sellowianum C. Presl, from Rio de Janeiro, Brazil, particularly Brade 8579 and Brade 20726 (both UC). It is also very similar to Glaziou 4667 (UC), labeled P. lanosum Fe´e, also from Rio de Janeiro (see Smith, 1981: 199); the two may be synonymous, with P. sellowianum having priority. This taxon differs from P. platyphyllum in lacking a rachis bud (all specimens seen) and in the very distinctive, large, rhombic-ovate pinnules having a deltate auricle at the acroscopic base. The separate blade fragment (three pinnae) on the left side of Mu¨nch 96 seems different from the plant on the right, and could be P. platyphyllum; this left-hand specimen has decidedly more denticulate costal and rachis scales than the plant on the right. If we are correct in suspecting that the plant on the right is conspecific with P. sellowianum (and P. lanosum), and the specimen is correctly labeled as to origin, then it would be a most remarkable disjunct distribution. It seems more likely, however, that it is mislabeled, a circumstance that is known to have happened with other Mexican/Brazilian ferns, especially in specimens emanating from Paris, as this one likely

Polystichum confluens E. Fourn., Mexic. Pl. 1: 92. 1872. Syntypes. Mexico. San Andres, Bottieri 1443, Herb. Van Heurck (P!); San Rafael, Guillemin s.n. (P!–photo US!).

Caudices 2–5 cm diam.; fronds mostly 50–150 cm long; stipes 20–45 cm long, 2–8 mm diam., ca. 1⁄3–2⁄3 the blade length; stipe bases with ovate-lanceolate scales mostly 10–20 ⫻ 1.5–3 mm, scales concolorous, dark red-brown or sometimes blackish, Ⳳ entire or sparsely ciliate, beneath these with smaller ciliate scurfy tan scales; blades subcoriaceous, (30–)50–100 ⫻ 10–30 cm, broadest at the bases or nearly so, attenuate at apices; rachises lacking buds, with lanceolate and hair-like scales mostly 2–5 mm long, a few scales sometimes blackish; pinnae pinnate, mostly 25–35 pairs, mostly 6–16 ⫻ 1.2–3 cm, pinnules entire to crenate with a small acroscopic auricle, less often with larger pinnules shallowly pinnatifid at the bases, usually lacking a Ⳳ free obovate lobe, both margins and tips spinulose; indument abaxially along costae and costules of fibrillose scales, also with ovate-lanceolate scales along the costae, these denticulate or spinulose; sori indusiate, often confluent at maturity, indusia whitish to tan, usually 1–1.5 mm diam., peltate.

Distribution. Terrestrial in wet forests; 1400–3200 m. Mexico; Guat. Selected Specimens Examined. Chis (Breedlove 41092, CAS). DF (Rzedowski 32250, CHAPA, IEB). Gro (Lorea 1921, NY). Gto (Ventura A. & Lo´pez P. 7796, IEB, UC). Jal (McVaugh 12888, MEXU, US). Me´x (Hinton 7134, ARIZ, MEXU, NY, US). Mich (Rzedowski 48333, 48376, IEB, UC). Mor (Lyonnet 1409, CAS, MEXU, US). Nay (Te´llez 11131, MEXU– or P. distans?). Oax (Mickel 6021, NY). Pue (Antonio s.n.,

POLYSTICHUM SPECIOSISSIMUM

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MEXU). Qro (Carranza 2404, IEB, UC). Sin (Breedlove 17111, CAS, NY).

See P. erythrosorum and P. distans for discussions of differences with those species. The identity of the sole collection from Sinaloa (as well as several of the collections from the Sierra Madre Occidental) is problematic, but it seems closest to P. rachichlaena; however, it is in this region that P. rachichlaena and P. distans are most difficult to distinguish. Polystichum rachichlaena hybridizes with P. speciosissimum to form plants with intermediate morphology. Spores of these hybrids are strongly malformed. Hybrids have been seen from the following localities: DF (Lyonnet 2686, MEXU). Me´x (Lugo 130, Ramı´rez E. 4, CAS, MEXU). Mor (Sa´nchez S. 582, US; Lyonnet 82, MEXU, US). sin loc. (Sa´nchez s.n., MEXU).

16. POLYSTICHUM SCHIZOPHYLLUM Polystichum schizophyllum Lorea-Hern. & A. R. Sm., Amer. Fern J. 89: 184. 1999. Type. Mexico. Guerrero: Mpio. Malinaltepec, 4 km S of Paraje Montero, Lorea 4574 (XAL; isotype UC!). Fig. 246E. Caudices erect, massive; fronds clumped, 1–1.4 m long; stipes 30–55 cm long, stramineous to dark red-brown, sparsely to densely scaly; stipe bases with scales of two types, some bicolorous with a central, lustrous dark brown to blackish band (the distal part sometimes blackish) and wide, translucent, light brown to brown margins, 6–20 ⫻ 0.8–4.5 mm, lanceolate to ovatelanceolate, entire to erose or fibrillose toward the tips, others light brown, concolorous, dull, 2–9 ⫻ 0.1–1.5 mm, capillary to lanceolate; blades 70–92 ⫻ 28–42 cm, deltate, 3-pinnate, rachises lacking buds, sparsely to conspicuously scaly, scales lanceolate to capillary, pinnae 2-pinnate-pinnatifid, 32–36 pairs, proximal pinnae 13–25 ⫻ 2.5–8 cm, about the same size as, or only slightly shorter than the third or fourth pinna pair, strongly ascending toward blade apices, gradually tapering to pinnatifid tips, free pinnules 20–25 pairs in largest pinnae, largest pinnules pinnatepinnatifid, each with 2–10(–16) free segments, penultimate blade segments (pinnulets) inequilateral at bases, basiscopically excavate, acroscopically the lobes larger and more spreading or slightly auriculate, margins denticulate-spinulose, not revolute; indument abaxially of moderate to sparse linear-lanceolate scales along costae and costules, also with sparse hair-like scales on veins, laminae glabrous adaxially; sori indusiate, 1–6 per ultimate segment (pinnulet), indusia (0.6–)1(–1.7) mm diam., circular or nearly so, entire, tan.

Distribution. Terrestrial in montane forests; 2000–2250 m. Mexico. Specimens Examined. Gro (Lorea 4538, XAL, UC).

This is the most divided of any Mexican Polystichum and is the only one that is fully 3-pinnate, with 2–10(–16) free ultimate segments on the larger pinnules. The presence of black-tinged scales mixed with fibrillose scales along the stipes and rachises in P. schizophyllum suggests a relationship to P. distans. However, the latter species has 2-pinnate fronds, more clearly bicolorous scales with lustrous black center and narrow light brown margins, and generally smaller indusia. It is likely that P. schizophyllum is most closely related to P. hartwegii, which also differs in its 2-pinnate fronds, but with a tendency, in some specimens, to have a nearly free acroscopic lobe on the pinnules. Polystichum schizophyllum is known from a small area in the Sierra Madre in southern Guerrero, and probably thrives in places where patches of mixed oak forest occur.

17. POLYSTICHUM SPECIOSISSIMUM Polystichum speciosissimum (A. Braun ex Kunze) Copel., Gen. Fil. 109. 1947. Figs. 247F–H. Cheilanthes speciosissima A. Braun ex Kunze, Analecta Pteridogr. 35, t. 23. 1837. Plecosorus speciosissimus (A. Braun ex Kunze) T. Moore, Index Fil. xl. 1857. Type. “E regno mexicano retulit de Karwinsky” (LZ, destroyed; isotype B!). Cheilanthes speciosissima A. Braun ex Kunze var. elata Liebm., Mexic. Bregn. 258 (reprint 106). 1849. Type. Mexico. Oaxaca: “Cerro de Sempoaltepec,” 9000’, Liebmann s.n. [Pl. Mex. 2479] (C!).

Caudices massive, to 30 cm long, 5–10(–15) cm diam.; fronds mostly 75–120 cm long; stipes 10–35 cm long, 5–10 mm diam., ca. 1⁄5–1⁄2 the blade length; stipe bases and also distal portions with very dense concolorous reddish brown to orangish tan scales (often obscuring stipes) 15–40 ⫻ 2–5 mm, margins with distant teeth, scales inrolled at tips, also with an underlayer of smaller fimbriate scales; blades coriaceous, 35–85 ⫻ 11–22 cm, narrowed at bases (to ca. 15 proximal pairs somewhat shortened with the lowest pairs ca. half the length of the longest pinnae), attenuate at apices; rachises lacking buds, densely scaly with narrow entire to denticulate scales 5–10 mm long and smaller fim-

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briate scales; pinnae pinnate-pinnatifid to 2-pinnate, ca. 25–50 pairs, 5–13 ⫻ 1–3 cm, the ultimate divisions with strongly revolute margins, bead-like; indument abaxially along costae of dense scales similar to rachis scales, veins with numerous hair-like scales, blade surfaces obscured by the revolute margins abaxially; sori exindusiate, seemingly confluent at maturity, almost completely covered by the whitish, strongly revolute, dentate segment margins; 2n⫽82 (CR).

Distribution. Terrestrial in alpine meadows, open grassy (sometimes grazed) areas, and pine forests at high elevations; (1600–)2150–4000 m. Mexico; Guat, CR, Pan. Selected Specimens Examined. Chis (Matuda 2318, F, MEXU, MICH, US). DF (Lyonnet 2040, IEB, MEXU, US). Gro (Hinton 14310, ARIZ, US). Gto (Rzedowski 47867, IEB). Hgo (Medina C. 1318, ASU, IEB, MEXU). Jal (McVaugh 11644, MEXU). Me´x (Balls B4181, UC). Mich (Zamudio & Pe´rez C. 7101, IEB, UC). Mor (Martı´nez 23, CAS, DS). Oax (Mickel 4577, NY, UC). Pue (Beaman 1977, MEXU, UC). Tlax (Vela G. 115, MEXU). Ver (Calzada & Horvitz 2287, UC, XAL).

18. POLYSTICHUM TURRIALBAE Polystichum turrialbae Christ, Bull. Herb. Boissier, se´r. 2, 6: 163. 1906. Type. Costa Rica. Volca´n Turrialba, 1905, Werckle´ s.n. (P). Figs. 247J–L. Polystichum smithii Mickel & Beitel, Mem. New York Bot. Gard. 46: 315. 1988. Type. Mexico. Oaxaca: Distrito Centro, top of Cerro San Felipe, 18 km N of Rte 175 on lumber road, Mickel 7056 (NY!).

Caudices 1–2 cm diam.; fronds mostly 40–70 cm long; stipes 15–30 cm long, 1.5–3 mm diam., often purplish brown (at least proximally), ca. 1⁄2 to nearly equaling the blade length; stipe bases with pale brown concolorous (sometimes darkened toward the center and base but not decidedly and sharply bicolorous), ovatelanceate scales ca. 5–9 ⫻ 1–3 mm, scales erose-denticulate and with short cilia along margins, rather few distally; blades chartaceous, 25–40 ⫻ 10–18 cm, broadest at the bases or nearly so, attenuate at apices; rachises lacking buds, with tan, ovate to lanceolate or hair-like scales 1–3 mm long, these denticulate or ciliate, especially at the broadened bases; pinnae pinnate, ca. 20– 25 pairs, 3–9 ⫻ 1.2–2.2 cm, pinnules crenate or shallowly incised, the basal acroscopic segments often more deeply pinnatifid, both margins spinulose, at least distally; indument abaxially along costae, costules, and veins of sparse to moderate lanceolate to fibrillose scales, the larger ones denticulate; sori indusiate, not confluent at maturity, indusia tan, ca. 0.5–0.8 mm diam., not overlapping in young fronds; 2n⫽82 (Oax).

Unverified, Doubtful, or Mistaken Reports. Qro (Ameneyro s.n., ENCB, cited by Arreguı´n et al., 2001, but not verified). SLP (reported by Mickel & Beitel, 1988, and Palacios-Rios, 1992, but not verified).

Polystichum speciosissimum is easily distinguished from all other species in Mexico by the strongly revolute, bead-like segments, thus appearing to lack spinulose margins; the margins appear erose to dentate with magnification. See P. rachichlaena for discussion of hybrids with that species. Hinton 26214 (TEX), from Oaxaca, 3300 m, appears to be a hybrid between P. speciosissimum and perhaps P. distans. It is less dissected than typical for P. speciosissimum and bears, along the stipes and proximal part of the rachises, some lanceolate scales that have a broad black central stripe and narrow tan margins; it also has the proximal pinnae only very slightly shortened relative to the longest lateral pair. Mickel 7066 (NY), also from Oaxaca, may also represent a hybrid involving P. speciosissimum, but the specimen is juvenile and sterile. Barrington (1990) reported hybrids in Costa Rica between P. speciosissimum and two additional species, P. concinnum Lellinger ex Barrington and P. talamancanum Barrington.

Distribution. Terrestrial in wet forest at high elevations; (2000– )2500–3100 m. Mexico; Salv (Monro et al. 1970, 2010, MO), CR, Pan; Bol. Selected Specimens Examined. DF (Lyonnet 1447, 3064, MEXU; Rzedowski 19492, NY). Hgo (Sa´nchez Mejorada 712, CAS, MEXU). Me´x (Lo´pez 603, ENCB, IEB; Lyonnet 761, MEXU, NY, US; Rzedowski 34237, ENCB; Schaffner s.n., MEXU, NY, US). Mich (Rzedowski 45801, 48510, IEB). Oax (Smith 514, UC).

This species has previously been called P. smithii, but Barrington (in litt., 1998) pointed out that the type of the name P. turrialbae (not treated by Barrington in Davidse et al., 1995) appears to be conspecific; we have not seen the type of P. tur-

POLYTAENIUM

rialbae, but specimens so identified in herbaria appear very similar to P. smithii. A collection of P. turrialbae from “in montibus San Rafael, prope San Luis Potosı´” matches perfectly Lyonnet 761 from San Rafael (Edo. Me´xico); one presumes that this locality is not in the state of San Luis Potosı´. Polystichum turrialbae in Mexico seems to be most common in and around the Valle de Me´xico and in Oaxaca. It closely resembles P. fournieri in the size and cutting of the blades and deflexed proximal pinnae, but that species differs primarily in the larger, whitish indusia and bicolorous, narrower, and distantly denticulate rhizome and stipe base scales. In P. turrialbae, the larger scales have cells that flare outward toward the margins and are decidedly more irregular in size and orientation (the margins thus erose-denticulate). Both species are found at relatively high elevations. Mickel and Beitel (1988: 315) compared P. turrialbae (as P. smithii) to P. mickelii in its distinctive stipe and rachis scales, but P. mickelii seems more distantly related by its considerably larger and broader stipe base scales, larger fronds, and exindusiate sori. The sole chromosome report for this species was made from Smith 514, originally reported as P. muricatum (Smith & Mickel, 1977). Hybrids and Problematic Specimens Here we list and discuss a number of problematic specimens, both those that are putative hybrids (parentage uncertain) and those that cannot be placed satisfactorily in any species recognized in this treatment at this time. Breedlove 22605 (DS, MEXU), Chis. Smith (1981) identified this as P. fournieri, with a query. It is not that, but probably a hybrid of uncertain parentage, perhaps involving P. distans. The spores are malformed and very irregular in size and shape. Mickel 4674 (NY, UC), Oax. This was originally cited as P. hartwegii by Mickel and Beitel (1988), but Barrington annotated it in 1993 as Polystichum sp. nov., aff. P. fournieri. We agree that it is not P. hartwegii and cannot match it with any known species. It differs from P. fournieri in having smaller indusia; perhaps its closest affinity is with P. turrialbae. Mickel 5310 (NY, UC), Oax. This is peculiar in the exindusiate sori and very reduced proximal pinnae, lowermost ca. 5 pairs gradually shortened, the most basal ones ca. 1 cm long; fronds ca. 45 cm long; blades ca. 35 ⫻ 15 cm; larger pinnules pinnatifid, ca. 10 ⫻ 5 mm; stipe base scales lanceate, to ca. 8 ⫻ 1.5 mm, brownish, entire, not particularly distinctive. We suspect that this specimen represents an undescribed species but are uncertain as to its affinities. Mickel and Beitel (1988) cited it as P. ordinatum, no doubt because of the reduced proximal pinnae, but it does not appear very closely related to that. Mohr s.n., 1857 (US), Ver. Although the blade is incomplete, this appears to have several pairs of reduced proximal pinnae. It might be P. ordinatum, but the indusia are very large, to 1.8 mm diam. The pinnules are relatively undivided, and so approach those of P. rachichlaena, but we do not otherwise know that species from Edo. Veracruz.

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Pringle 5588 (MEXU, US), Ver. This appears to match Mohr s.n., discussed above, with very large indusia and reduced proximal pinnae. Tenorio 18554 (MEXU–3 sheets), Oax. Most likely this represents an undescribed species, probably confined to high elevations (2800 m). The stipe base scales are most similar to those of P. alfaroi, but concolorous and light orange-brown without a darkened middle band. Pinnules are Ⳳ entire and spreading (thus differing from P. alfaroi), with a deltate basal auricle, and coriaceous with revolute margins, similar to some specimens of P. distans (which has completely different stipe base scales). We are unable to suggest its nearest ally. Unplaced Names Polystichum aculeatum (L.) Roth α incisum Fe´e, Me´m. Foug. 8: 100. 1857. Type. Mexico. “Cordoba des Mexicains,” Schaffner 250 (not located). Polystichum aculeatum (L.) Roth ß gracile Fe´e, Me´m. Foug. 8: 100. 1857. Type. Mexico. “Guatimalpan et San Agostin,” Schaffner 315 (not located). Polystichum aculeatum (L.) Roth γ distans Fe´e, Me´m. Foug. 8: 100. 1857. Type. Mexico, “Totutla des Mexicains,” Schaffner 116 (not located).

1 0 3. P O L Y T A E N I U M Polytaenium Desv., Me´m. Soc. Linn. Paris 6: 218. 1827. Antrophyum subg. Polytaenium (Desv.) Benedict, Bull. Torrey Bot. Club 34: 447. 1907. Type: Polytaenium lanceolatum (Sw.) Desv. [⬅ Hemionitis lineata Sw.] ⬅ Polytaenium lineatum (Sw.) J. Sm. Epiphytic; rhizomes short-creeping, with decidedly clathrate scales; roots densely tomentose with abundant root hairs; fronds small; stipes short to absent; blades simple, linear to elliptic or oblanceolate, entire, glabrous; veins anastomosing to form 2– several series of areoles, without included veinlets; epidermis with linear idioblasts; sori linear, in 2–6(–8) lines parallel to the midvein or in shorter, irregular oblique lines, either on the surface or more commonly immersed in the lamina; indusia absent; paraphyses absent; spores yellow-hyaline or rarely green, tetrahedralglobose; x⫽60. Polytaenium comprises 8–10 neotropical species, with four species occurring in Mexico. The genus is restricted to tropical and montane rain forest habitats throughout its range. Often, it has been subsumed under Antrophyum s.l., which in the restricted sense comprises about 20 species in Africa, south-central and southeastern Asia, and islands of the western Pacific (Polynesia). According to phylogenetic reconstruction done by Crane (1997), Antrophyum is sister to a predominantly New World clade comprising Polytaenium, Vittaria s.str., Ananthacorus, and Scoliosorus. Polytaenium and Antrophyum have historically been

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distinguished by the presence of a distinct, or at least proximally distinct, midrib in the former genus and the absence of a midrib in the latter. Species of Polytaenium lack paraphyses, and those of Antrophyum bear paraphyses. Polytaenium is distinguished from Vittaria by having more than one sorus on each side of the midrib, or by having continuous sori along the reticulate veins.

References Benedict, R. C. 1907. The genus Antrophyum–I. Synopsis of the subgenera, and the American species. Bull. Torrey Bot. Club 34: 445–458; Benedict, R. C. 1911. The genera of the fern tribe Vittarieae: their external morphology, venation, and relationships. Bull. Torrey Bot. Club 38: 153–190; Crane, E. H. 1997 [1998]. A revised circumscription of the genera of the fern family Vittariaceae. Syst. Bot. 22: 509–517; Tryon, R. 1964. Taxonomic fern notes IV. Some American vittarioid ferns. Rhodora 66: 110–117.

Key to the Mexican Species of Polytaenium 1. Sporangia (sori) in (2–)4–6 lines parallel to the midrib; spores tetrahedral. ............................................................................................ 4. P. lineatum. 1. Sporangia (sori) in irregular lines at an angle to the midrib; spores bilateral or tetrahedral. 2. Blades coriaceous, oblanceolate to obovate, 2–3 cm wide at broadest point; sori in grooves. .......................................................... 1. P. cajenense. 2. Blades chartaceous, narrowly to linear-elliptic, 1–2 cm wide at broadest point; sori surficial or in very shallow grooves. 3. Spores green; veins free and Ⳳ oblique or perpendicular to the blade margins; sori surficial. ........................................... 2. P. chlorosporum. 3. Spores pale yellowish, hyaline; veins forming marginal areoles or, if free, curving and running Ⳳ parallel to blade margins; sori usually in very shallow grooves, sometimes surficial. ......................................................................................................................... 3. P. feei.

1. POLYTAENIUM CAJENENSE Polytaenium cajenense (Desv.) Benedict, Bull. Torrey Bot. Club 38: 169. 1911. Figs. 251K–M. Hemionitis cajenensis Desv., Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck. Gesammten Naturk. 5: 311. 1811. Antrophyum cajenense (Desv.) Spreng., Syst. Veg. 4: 67. 1827. Type. French Guiana. Cayenne, Martin s.n. (PDesv.!; isotype B-Willd. 19560, microfiche UC!). Antrophyum lacantunense Rovirosa, Pteridogr. Sur Me´xico 240, t. 38A, f. 10– 11. 1909. Type. Mexico. Chiapas: fluvium Lacanja´ e tribu Lacandonorum, Martı´nez s.n. (Herb. Rovirosa 1097, holotype not found; isotype GH).

Rhizomes short-creeping; rhizome scales linear-lanceolate, long-attenuate at apices, 3–4 ⫻ 0.3–0.5 mm; fronds 8–20 cm long, clumped; stipes alate, midribs prominent; blades coriaceous, oblanceolate, 2–3 cm wide at broadest point (2⁄3 up frond length), 1.5–2 mm wide at base, apices acute; veins obscure, anastomosing in a dense network, veins near blade margins forming areoles; sporangia (sori) in many anastomosing lines 1–10 mm long, sunken in grooves; spores pale yellowish, hyaline.

Selected Specimens Examined. Chis (Breedlove 34060, DS, 57378, CAS, ENCB; Martı´nez S. 10096, 16904, 25330, 25448, MEXU). Oax (Herna´ndez G. 1933, CHAPU, MEXU; Mickel 6466, 6467, NY, UC). Ver (Valdivia 1826, XAL).

Polytaenium cajenense is distinguished by its coriaceous, oblanceolate blades with sori in distinct grooves. The South American species, Polytaenium brasilianum (Desv.) Benedict (syn. ⫽ Antrophyum subsessile Kunze, a superfluous renaming), is very similar to, and possibly conspecific with, P. cajenense, according to Tryon and Stolze (1989: 87); we agree.

2. POLYTAENIUM CHLOROSPORUM Polytaenium chlorosporum (Mickel & Beitel) E. H. Crane, Syst. Bot. 22: 516. 1997 [1998]. Figs. 251G, H, J. Antrophyum chlorosporum Mickel & Beitel, Mem. New York Bot. Gard. 46: 41–42. 1988. Type. Mexico. Oaxaca: Distrito Choapan, ridge between Yetzelalag and Lovani, from summit to river-crossing at bottom of grade toward Lovani, Hallberg 1546 (NY! isotype UC!).

Rhizomes erect to short-creeping; rhizome scales lanceolate, 3– 4 mm long; fronds 12–20 ⫻ 1.3–2.2 cm wide, clumped; stipes absent; blades papyraceous, narrowly oblanceolate to almost elliptic, bases long-attenuate, decurrent, 2 mm wide, apices acuminate; veins netted, oblique, those near blade margins free, midribs prominent; sporangia (sori) following the entire vein pattern, surficial, not impressed in grooves; spores green.

Distribution. Epiphytic in dense wet forests of Atlantic slope, lowland and lower montane rain forests; 100–600 m. Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Cuba, Jam; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol.

POLYTAENIUM FEEI

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Rhizomes short-creeping; rhizome scales clathrate, lanceattenuate, 2–3 ⫻ 0.5–0.8 mm; fronds 7–23 cm long, clumped; stipes absent; blades papyraceous, linear-oblanceolate to linearelliptic, 9–12(–18) mm wide at widest point (1.5–2 mm wide at bases), apices acuminate, laminae glabrous; veins forming 3–4 rows of oblique areoles between midribs and blade margins, veins near blade margins also areolate; sporangia (sori) surficial or in very shallow grooves, following most of the netted vein pattern, at least in distal 2⁄3 of blades; spores pale yellowish, hyaline; 2n⫽ca. 120 (PR).

Distribution. Epiphytic in montane rain forests; 1100–1800 m. Mexico; Nic, CR; w Col. Specimens Examined. Chis (Breedlove 32212, DS, 35203, DS, MEXU, 38946, DS, NY, 49719, CAS). Oax (only the type collection).

The green spores, thin blade texture, and free marginal veins distinguish this species from its Mexican congeners. It seems quite peculiar that P. chlorosporum, and no other species in the genus (or family, for that matter) produces chlorophyllous spores, but we have been unable to document this condition in any other neotropical Vittariaceae. Perhaps the green-spored state is a function of certain environmental conditions, but in the absence of an explanation we maintain the species as distinct, based not only on this spore character, but also the free veins that are oblique or nearly perpendicular to the blade margins. Collections from Chiapas were previously cited by Smith (1981: 34) as Antrophyum guayanense Hieron., a species we now believe to be confined to Trinidad and South America (Col, Ven, Sur, Fr Gui, Peru, Braz, Bol). Polytaenium guayanense (Hieron.) Alston is perhaps a close ally of P. chlorosporum, but it lacks green spores; another close relative may be Polytaenium anetioides (Christ) Benedict, from Costa Rica, which lacks prominent blade midribs. Polytaenium cajenense differs from P. chlorosporum in having pale yellowish spores, sunken veins and sori, oblanceolate, coriaceous fronds, and marginal areoles.

3. POLYTAENIUM FEEI Polytaenium feei (W. Schaffn. ex Fe´e) Maxon, Pteridophyt. Porto Rico 405. 1926. Figs. 251A–C. Antrophyum feei W. Schaffn. ex Fe´e, Me´m. Foug. 7: 42, t. 22, f. 1. 1857. Type. Mexico. [Veracruz:] Huatusco, Schaffner 133 (P!; isotype K!). Hemionitis lanceolata L., Sp. Pl. 2: 1077. 1753. Antrophyum lanceolatum (L.) Kaulf., Enum. Filic. 198. 1824. Polytaenium lanceolatum (L.) Benedict, Bull. Torrey Bot. Club 38: 169. 1911, hom. illeg., non (Sw.) Desv., 1827. Lectotype (designated by Proctor, 1985: 258). Plumier, Traite´ Foug. Ame´r., pl. 127, f. C, 1705 (cited as f. 6 by Linnaeus, in error), based on a specimen from St. Vincent. Antrophyum stenophyllum Rovirosa, Pteridogr. Sur Me´xico 242. 1910, hom. illeg., non Baker, 1898. Type. Mexico. Chiapas: Cerro de Moyos, Rı´o Lacanja´ region, Martı´nez s.n. (Herb. Rovirosa 1104, not seen, not found at K, MEXU, P, PH).

Distribution. Epiphytic in wet forests on Atlantic slope, lower montane rain forests; 150–1400 m. Mexico; Guat, Bel, Hond, Nic, CR, Pan; Gr & L Ant; Ven, Trin, Sur, n Braz. This is accorded a much wider distribution in South America (Col, Ec, Peru, Bol) by Moran (in Davidse et al., 1995), but we doubt its occurrence beyond northern South America, and it is not cited from Peru by Tryon and Stolze (1989). Selected Specimens Examined. Chis (Breedlove 53509, CAS, ENCB; Matuda 18389 DS, MEXU, US; Purpus 6758, UC, US). Oax (Mickel 5727, 6465, 7206, NY, UC). Ver (Copeland herb. 159, MICH, UC; Purpus 2230, UC; Ventura A. 12429, ENCB; Wendt et al. 4014, NY).

This species is distinguished from P. cajenense by the elliptic blades, thinner blade texture, nearly surficial sori, and long axis of the areoles more or less parallel to the midribs. Like P. chlorosporum, P. feei has veins often or mostly free along the blade margins, but these tend to curve and extend for quite a long distance parallel to the blade margins, whereas in P. chlorosporum the free veins are relatively short and oblique or perpendicular to the blade margins. The veins are generally more visible and sunken abaxially in P. feei than in P. chlorosporum. Polytaenium feei often goes under the name of Antrophyum lanceolatum; however, the epithet lanceolatum cannot be used in Polytaenium for this species because of the prior existence of P. lanceolatum (Sw.) Desv., a synonym of P. lineatum. Material of P. feei from the West Indies is often somewhat larger and has thicker texture; this variation is probably not significant taxonomically.

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4. POLYTAENIUM LINEATUM Polytaenium lineatum (Sw.) J. Sm., J. Bot. (Hooker) 4: 68. 1841. Figs. 251D–F. Hemionitis lineata Sw., Prodr. 129. 1788. Vittaria lanceolata Sw., Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck. Gesammten Naturk. 2: 133. 1799, non Vittaria lineata (L.) Sm., 1793. Antrophyum lineatum (Sw.) Kaulf., Enum. Filic. 199. 1824. Polytaenium lanceolatum (Sw.) Desv., Me´m. Soc. Linn. Paris 6: 218. 1827. Type. Jamaica. Swartz s.n. (presumably S; isotype B-Willd. 20033!, photo GH!, microfiche UC!).

Rhizomes short-creeping; rhizome scales lanceolate, 3–4 ⫻ 0.6– 0.8 mm; fronds 6–30(–43) cm long, clumped; stipes alate, lightcolored, no darker than surrounding tissue; blades coriaceous or subcoriaceous, narrowly elliptic, 0.5–1(–1.5) mm wide, midribs stramineous to light green, not very evident; veins obscure, netted, those along blade margins also forming areoles; sporangia (sori) sunken in (1–)2–4 Ⳳ straight parallel rows on each side of the midvein; spores pale yellowish, hyaline; 2n⫽120 (Jam.).

ing to pendent, dichotomously branched several times, terete, angular or flattened, glabrous; true leaves absent but stems with minute, awl-shaped, scale-like enations; sporangia mostly 3lobed, subtended by 2-lobed enations on distal branches; spores monomorphic, bilateral; gametophytes subterranean, fleshy, cylindrical; x⫽52. Psilotum is generally considered to have two pantropical species, growing mostly on the trunks of either living or fallen trees or at the bases of trees, but occasionally among rocks. Bierhorst (1977) proposed that Psilotum be considered a fern, and molecular evidence (Hasebe, M. et al., 1995; Pryer, K. M., et al. 2001) has shown that the family Psilotaceae is most closly allied to the Ophioglossaceae. References Bierhorst, D. W. 1977. The systematic position of Psilotum and Tmesipteris. Brittonia 29: 3–13; Mu¨ller, K. 1856. Monografische Kritik der Lycopodiaceen Gattung Psilotum. Bot. Zeitung (Berlin) 1856: 217–227; 233–243, t. 7, f. 1–11; Wagner, W. H., Jr. 1977. Systematic implications of the Psilotaceae. Brittonia 29: 54–63.

Key to the Mexican Species of Psilotum 1. Aerial stems and branches flattened; main aerial stems ca. 2 mm diam. ........................................................... 1. P. complanatum. 1. Aerial stems and branches terete to angled but not flattened; main aerial stems 1–1.5 mm diam. ................... 2. P. nudum.

1. PSILOTUM COMPLANATUM Psilotum complanatum Sw., J. Bot. (Schrader) 1800(2): 110. 1801. Type. Jamaica. Swartz s.n. (S; isotype B-Willd. 19437!). Figs. 252D, E. Distribution. Epiphytic in wet forests of Atlantic and Pacific slopes, montane rain forests, montane cloud forests; 800–1600(– 2000) m. Mexico; Guat, CR, Pan; Gr Ant; Col, Ven, Trin, Ec, Peru, s Braz, Bol, nw Arg. Selected Specimens Examined. Chis (Breedlove 22152, 31524, 32213, DS, 56497, CAS; Mu¨nch 175, DS, P). Gro (Lorea 3039, US). Oax (Hallberg 1491, NY; Mickel 6190, NY, UC). Ver (Mu¨ller 434 NY; Purpus 254, US).

The parallel soral lines, usually in 4–6 rows, distinguish P. lineatum from congeners in Mexico.

1 0 4 . PS I L O T UM Psilotum Sw., J. Bot. (Schrader) 1800(2): 8, 109. 1801. Lectotype (first chosen by Brongniart, Dict. Class. Hist. Nat. 9: 558. 1825): Psilotum triquetrum Sw. [⫽ Psilotum nudum (L.) P. Beauv.]. Epiphytic or terrestrial; rhizomes short-creeping, much branched, with brown hairs; roots absent; aerial stems erect, arch-

Stems erect or arching, 10–50 cm long; main aerial stems flattened, ca. 2 mm wide, terete or angled at bases, 2–7 times dichotomously branched; ultimate branches ca. 1 mm diam.; enations 1–2 mm long; 2n⫽104 (various South Pacific islands), 208 (Malaysia).

PTERIDIUM

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Pan; Bah, Gr & L Ant; Col, Ven, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol, Parag, Arg; Old World tropics, Australia, New Zealand. Selected Specimens Examined. Chih (Pringle 2972, MO). Chis (Martı´nez 16991, MEXU). Col (Orcutt 6317, MEXU). DF (Balls B4259, BM, K, UC). Gro (Moreno G. & M. Luna 817, FCME). Hgo (Herna´ndez M. & Herna´ndez V. 4522, MEXU). Jal (Palmer 688, BM, GH, K, MO, NY). Me´x (Hinton 3923, BM, ENCB, K, MEXU, MO). Mor (Sandoval 99, MEXU, UAMIZ). Pue (Riba et al. s.n., 25 Aug 1967, GH). QR (Lot & Novelo 819, GH, MEXU). Rev (Barkelew 218, GH, MO, NY, UC, US). Son (Gentry 2110, ARIZ, MO, UC). Tab (Ventura A. 20438, CHAPA, ENCB, IEB, MEXU). Tam (Dressler 2021, MO). Ver (Purpus 1988, BM, F, NY, UC). Unverified, Doubtful, or Mistaken Reports. Yuc (Ciau 10, UADY, cited by Palacios-Rios, 2002b, but not verified).

Distribution. Epiphytic in wet montane forests on tree fern trunks, or terrestrial among rocks, or on basaltic rocks; (600–) 1100–2300 m. Mexico; Guat, Hond, Salv; Cuba, Jam, Hisp; Col, Ven, Peru; Malaysia, Oceania. Selected Specimens Examined. Chih (Pringle 450, BM, K, MEXU). Chis (Mu¨nch 102, DS). DF (Rzedowski 22147, ENCB, TEX). Gro (Lorea 4282, FCME). Hgo (Gonza´lez G. 58, CHAPA, ENCB). Jal (Pringle 328, MEXU). Mor (Riba 1680, UAMIZ). Nay (McVaugh 15240, MICH). Oax (Mickel 5624, NY). Pue (Rzedowski 23413, ENCB). Son (Rondeau 90–48, ARIZ). Tam (Gonza´lez-Medrano 1431, MEXU). Ver (Purpus 1989, UC).

2. PSILOTUM NUDUM Psilotum nudum (L.) P. Beauv., Prodr. Aethe´ogam. 106, 112. 1805. Figs. 252A–C. Lycopodium nudum L., Sp. Pl. 2: 1100. 1753. Psilotum triquetrum Sw., J. Bot. (Schrader) 1800(2): 109. 1801. Type. “In Indiis.” LINN 1257.1. For additional synonymy, see Proctor (Fl. L. Antill. 2: 16, 18. 1977).

Stems erect or arching, mostly 20–60 cm long; main aerial stems 1–1.5 mm diam., terete or angled, 3–7 times dichotomously branched; ultimate branches ca. 0.8 mm diam.; enations 1–1.5 mm long; 2n⫽104 (NZ), 156, 208 (India).

Distribution. Epiphytic in wet montane forests, evergreen rain forests, also among rocks, lower montane rain forests; 180– 800(–1500) m. Se USA; Mexico; Guat, Bel, Hond, Nic, CR,

The slender, terete to angled aerial stems readily distinguish this species from P. complanatum.

105. PTERIDIUM Pteridium Gled. ex Scop., Fl. Carniol. 169. 1760, nom. cons. Type: Pteridium aquilinum (L.) Kuhn [⬅ Pteris aquilina L.]. Terrestrial; rhizomes long-creeping, subterranean, with hairs; fronds medium-sized to very large; stipes long, glabrous to shorthairy, the bases not strongly distinct from the rhizomes, with branch buds near the bases; blades broadly deltate, 2–4-pinnate, the basal pinnae often exaggerated basiscopically, each basal pinna nearly equal to distal part of blade in size, with nectaries at the base of first and sometimes distal pinna pairs; laminae chartaceous to coriaceous, sparsely to densely hairy abaxially with straight to lax hairs, often with scattered minute 2-celled hairs adaxially; veins free, forking, ends fusing at soriferous margins; sori marginal with differentiated recurved marginal false indusia and sometimes very slight inner, extrorse indusia; spores tetrahedral-globose, brown; x⫽52. Pteridium has been treated as a single widespread species with several regional varieties (Jacob & Peck in FNA Ed. Comm., 1993; R. Tryon, 1941) or as several distinct species (Moran in Davidse et al., 1995; Mickel & Beitel, 1988). Three species, one including three varieties, occur in Mexico. The plants are generally aggressive, invading disturbed areas as weeds in pastures, cultivated fields, and roadsides. The genus is closely allied to Paesia and Hypolepis. Pteridium is distinguished by its hairy rhizomes well beneath the soil surface, coriaceous blades, vestigial inner indusia, and tetrahedral spores. Throughout much of Mexico, Pteridium arachnoideum, P. caudatum, and P. aquilinum var. feei are separated by correlated characters, except for a few individual plants that combine character states of two species. Since these character states remain consistent even when two or more taxa grow sympatrically (Mickel & Beitel, 1988), it seems desirable to treat them at species rank. This view is supported by Thomson’s (2000) conclusions based on morphological and molecular data. Taxa here treated as varieties of Pteridium aquilinum (vars. feei, latiusculum,

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PTERIDIUM

and pubescens) appear closely allied by their long, lax hairs, and have more or less allopatric distributions. There is wide variation in degrees and type of pubescence between and sometimes within taxa. Collections that combine the characters of two varieties are treated as hybrids. In the few fertile specimens, the spores appear to be well formed and the characters seem to be so variable. Field studies and modern biosystematic studies are needed in Pteridium.

References Perring, F. H. & B. G. Gardener (eds.). 1976. The biology of bracken. J. Linn. Soc., Bot. 73: 1–302 (papers by several authors); Thomson, J. A. 2000. Morphological and genomic diversity in the genus Pteridium (Dennstaedtiaceae). Ann. Bot. (Oxford) 85 (Suppl. B): 77–99; Thomson, J. A. & M. E. AlonsoAmelot. 2002. Clarification of the taxonomic status and relationships of Pteridium caudatum (Dennstaedtiaceae) in Central and South America. J. Linn. Soc., Bot. 140: 237–248; Tryon, R. M. 1941. Revision of the genus Pteridium. Rhodora 43: 1–31; 37–70 (reprinted in Contr. Gray Herb. 134: 1–31, 37–70.

Key to the Mexican Species of Pteridium 1. Free lobes, or rarely long-decurrent segment bases, present on axes between segments; laminar surfaces abaxially with dense white hairs 0.5–0.8 mm on veins and minute, arachnoid, white hairs covering the laminar surfaces between veins; minute glandular hairs absent on adaxial laminar surfaces near segment tips; segment midveins sulcate abaxially bearing twisted, clear hairs with dark end walls. ............................................................................................................................................................................................ 2. P. arachnoideum. 1. Free lobes absent on axes between segments; laminar surfaces abaxially with rare to dense, white hairs 0.5–0.8 mm, minute arachnoid hairs absent, laminar surfaces between veins glabrous; minute glandular hairs absent or scattered on adaxial blade surfaces near segment tips; segment midribs slightly flattened to terete, bearing either scattered, straight, clear brown glandular hairs 1–1.5 mm or scattered, minute, brown, glandular hairs less than 0.1 mm. 2. Fronds generally small, 1 m or less; segment midribs terete, bearing scattered, straight, clear brown hairs 1–1.5 mm; minute glandular hairs absent on adaxial laminar surfaces near segment tips. .............................................................................................. 1. P. aquilinum. 2. Fronds generally large, 1.5–4 m long; segment midribs flattened, bearing scattered, minute, brown, clavate hairs less than 0.1 mm; minute glandular hairs 0.1 mm scattered on adaxial blade surfaces near segment tips. ......................................................... 3. P. caudatum.

1. PTERIDIUM AQUILINUM Pteridium aquilinum (L.) Kuhn in v. d. Decken, Reisen OstAfrika 3(3): 11. 1879. Pteris aquilina L., Sp. Pl. 2: 1075. 1753. Type. Filix femina Fuchs, Hist. 596, misprinted 569. 1542. Figure of a plant from Europe. There is no specimen in the Linnaeus herbarium.

Fronds 0.5–1 m long; rachises slightly to strongly pubescent; blades lacking free lobes between segments; penultimate axes pubescent abaxially with white hairs, scattered, clear brown, straight hairs 1–1.5 mm long, adaxially with short, white hairs, red-tipped, glandular hairs confined to medial grooves; costae terete abaxially; laminar surfaces abaxially between costae and segment margins pubescent to glabrous on veins, adaxially with scattered to rare hairs only along reflexed margins, hairs 0.5–0.8 mm long, white; false indusia 0.2–0.8 mm wide with erose margins, with scattered to numerous long cilia, or glabrous. Distribution. Terrestrial in open disturbed areas; 0–3300 m. North America, Mexico, Central America, Europe, Asia, Hawaii. Distinct by the penultimate axes with lax white hairs, often also with scattered straight clear brown hairs.

Key to the Mexican Varieties of Pteridium aquilinum 1. Abaxial laminar surfaces glabrous, the hairs limited to the midveins and margins (but not the indusia). ..................................................................................... 1b. var. latiusculum.

1. Abaxial laminar surfaces with hairs on the veins as well as on the midveins and margins. 2. Abaxial hairs straight, not covering the laminar surfaces; sterile indusia 0.25 mm or less wide; Baja California. ................................................................ 1c. var. pubescens. 2. Abaxial hairs tortuous to curved, often obscuring the laminar surfaces; sterile indusia 0.4 mm or more wide; most of Mexico other than Baja California and the northern tier of states. .............................................. 1a. var. feei.

1a. Pteridium aquilinum var. feei (W. Schaffn. ex Fe´e) Maxon ex Yunck., Publ. Field Mus. Nat. Hist., Bot. Ser. 17: 308. 1938. Figs. 253A–C. Pteris feei W. Schaffn. ex Fe´e, Me´m. Foug. 8: 73. 1857. Pteridium feei (W. Schaffn. ex Fe´e) Faull, Contr. Arnold Arbor. 11: 87. 1938. Syntypes. Mexico. [Veracruz:] Huatusco, Schaffner 138, 141 (both P!); Popocate´petl, Schaffner 286 (P?).

Adaxial laminar surfaces with scattered hairs, primarily on costae and near segment margins; terminal cell of costal hairs pale, inconspicuous; abaxial costae with lax pale hairs, also with frequent clear brown stiff hairs; segments with abundant, long, tangled hairs 0.6–1 mm long, often obscuring the laminar surfaces, also evident from curved adaxial margins and margins of indusia; indusia 0.4 mm or more wide.

PTERIDIUM AQUILINUM

Distribution. Open, disturbed areas at higher elevations; 1800– 3050 m. Mexico; Guat, Hond, Salv. Selected Specimens Examined. Ags (McVaugh 18453, MEXU, MICH). Chis (Ghiesbreght 224, NY). Coah (Riskind et al. 1719, LL). DF (Ventura A. 1428, NY). Dgo (Fisher s.n., 13 Aug 1937, NY). Gro (Hamilton & Rowell 3218, NY). Gto (Galva´n 2498, IEB). Hgo (Rose 9204, NY). Jal (Puga & Carvajal H. 10172, ENCB). Me´x (Dunn & Dunn 18881, NY). Mich (King & Soderstrom 4742, NY). Mor (Lyonnet 2417, US). Nay (Norris & Taranto 14536, MICH). NL (Rodrı´guez L. 95, NY). Oax (Mickel 1392, NY). Pue (Ventura A. 1260, NY). Qro (Zamudio 2674, IEB). Sin (Gonza´lez Ortega 4079, MEXU). SLP (Mickel 557, NY). Ver (Nee & Taylor 26764, NY). Unverified, Doubtful, or Mistaken Reports. Tab (reported by Magan˜a, 1992, but figured plant appears to be P. caudatum).

Pteridium aquilinum var. feei is distinguished by lamina abaxially with tortuous or curved hairs, indusia broad and ciliate, laminar hairs 1–1.5 mm long, straight, clear, brown. 1b. Pteridium aquilinum var. latiusculum (Desv.) Underw. ex Heller, Cat. N. Amer. Pl., ed. 3, 17. 1909. Fig. 253F. Pteris latiuscula Desv., Me´m. Soc. Linn. Paris 6: 303. 1827. Type. Canada. Newfoundland and St. Pierre, s. coll. s.n. (P-Desv., photos GH, US).

Adaxial laminar surfaces with scattered hairs, primarily on costae and near segment margins; abaxial costae with lax pale hairs, axes with occasional brown hairs; segments with hairs on costae and on curved adaxial margins but glabrous between costae and margins; indusial margins glabrous; 2n⫽104 (USA).

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Distribution. Dry to moist pine-oak forests; 2000–2650 m. Canada, e USA, w to e slope of Rocky Mts; Mexico; Europe, Asia. Selected Specimens Examined. Chih (Knobloch 78, MEXU; Palmer 288, NY). Coah (Wendt & Adamcewicz 501, TEX). Dgo (Gonza´lez 1530, MEXU; Maysilles 7748, NY). NL (Hinton 22135, TEX; Mueller & Mueller 366, TEX, MEXU). Sin (Dunn et al. 21801, NY; Johnston et al. 11441, NY). Son (Phillips 418, US).

Most reports of var. pubescens from Mexico are var. latiusculum. Var. latiusculum is distinct from var. pubescens in having glabrous abaxial laminar surfaces except for hairs on the costae and segment margins (but not on indusia). Var. pubescens is distinct from var. latiusculum in its abaxial surfaces densely pubescent between margins and costae vs. only on costae, and the indusia ciliate to pubescent vs. glabrous. It varies from glabrous to sparsely hairy on the laminar tissue abaxially. Var. pubescens, with its straight hairs scattered on the abaxial surfaces, is limited to the Baja California peninsula and extends northward along the coast to Alaska, extending inland across Canada to northern Michigan and widespread in the western United States. Var. latiusculum, besides occurring in eastern North America, is found in the eastern Rockies and also in Arizona and New Mexico. As Tryon (1941) pointed out, in areas of sympatry var. latiusculum and var. pubescens intergrade. Jacobs and Peck (in FNA Ed. Comm., 1993) claimed that in the United States, all Pteridium west of the Great Plains is var. pubescens, and var. latiusculum occurs east of the Great Plains, but not in Mexico. 1c. Pteridium aquilinum var. pubescens Underw., Native Ferns, ed. 6, 91. 1900. Figs. 253D, E. Pteris aquilina L. var. pubescens (Underw.) Clute, Fern Bull. 15: 124. 1907. Type. No type designated by Underwood. Neotype (chosen here). U.S.A. California: San Bernardino Mts, Aug 1890, Parish s.n. (NY!).

Adaxial laminar surfaces with hairs more or less evenly distributed on costae, margins, and laminar tissue, terminal cell of costal hairs dark reddish brown; costae with lax pale hairs 0.3– 0.5 mm long, clear brown stiff hairs sparse to absent; segments with scattered erect to curved hairs on costae and laminar surfaces abaxially (also on veins) and also evident on curved adaxial margins and indusial margins; indusia 0.25 mm or less wide.

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PTERIDIUM

Distribution. In dry to moist woods and open areas in partial to full sun, forming abundant colonies; 200–1000 m. Canada, USA; Mexico. Specimens Examined. BCN (Boyd 2612, UC; Gallegos s.n., in 1923, MEXU, US; Moran 26634, ASU, 30473, UC; Wiggins 16673, MEXU, US, 21481, DS). BCS (Aguirre Claveran 96, ENCB).

Reports of var. pubescens from other Mexican states are generally var. latiusculum.

Distribution. Open, disturbed areas; 100–2450 m. Mexico; Guat, Hond, Salv, Nic, CR, Pan; Gr Ant, St. Lucia; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Galapagos, Peru, Braz, Bol, Parag, Arg, Uru. Selected Specimens Examined. Chis (Breedlove 21816, NY). Gro (Hinton 14069, NY). Hgo (Pe´rez S. 623, ENCB, MEXU). Jal (McVaugh & Koelz 1159, MEXU). Me´x (Hinton 7371, NY). Oax (Mickel 4742, 7146, NY). Pue (Clausen 7466, NY). Tam (Lof 759, UC). Ver (Purpus 6191, NY).

2. PTERIDIUM ARACHNOIDEUM

Unverified, Doubtful, or Mistaken Reports. Mor (reported by Riba et al., 1996, but cited specimens misidentified, or doubtful). Tab (reported by Magan˜a, 1992, but figured plant is P. caudatum).

Pteridium arachnoideum (Kaulf.) Maxon, J. Wash. Acad. Sci. 14: 89. 1924. Figs. 253G, H, J.

Pteridium arachnoideum is distinguished most readily by free lobes between the segments near the tips of the pinnae.

Pteris arachnoidea Kaulf., Enum. Filic. 190. 1824. Pteris aquilina L. var. arachnoidea (Kaulf.) D. C. Eaton, Proc. Amer. Acad. Arts, ser. 2, 8: 203. 1861. Pteridium aquilinum (L.) Kuhn var. arachnoideum (Kaulf.) Brade, Z. Deutsch. Ver. Wiss. Kunst. Sa˜o Paulo 1: 56. 1920. Pteridium caudatum (L.) Maxon subsp. arachnoideum (Kaulf.) Lellinger, Amer. Fern J. 93: 150. 2003. Type. Brazil. Chamisso s.n. (LE!, photo GH). For additional synonymy, see R. Tryon (1941).

Fronds (1–)1.5–3 m long; blades with free lobes or rarely longdecurrent segment bases between ultimate segments; rachises glabrous to slightly pubescent; penultimate axes pubescent abaxially with hairs scattered to dense, curly, clear, 0.5 mm, with dark end walls, adaxially with hairs white, especially in medial grooves; costae and veins usually sulcate abaxially; laminar surfaces densely hairy abaxially on veins with hairs 0.5–0.8 mm, twisted, white, and hairs minute, arachnoid, white, covering the laminar surfaces between veins, adaxially with scattered, white hairs mostly along margins, minute glandular hairs absent; false indusia 0.1– 0.3 mm wide with erose margins with rare to scattered cilia; 2n⫽104 (Trin), 208 (Galapagos).

3. PTERIDIUM CAUDATUM Pteridium caudatum (L.) Maxon, Proc. U.S. Natl. Mus. 23: 631. 1901. Figs. 253K–M. Pteris caudata L., Sp. Pl. 2: 1075. 1753. Pteris aquilina L. var. caudata (L.) Link, Hort. Berol. 2: 33. 1833. Pteridium aquilinum (L.) Kuhn var. caudatum (L.) Sadeb., Jahrb. Hamburg Wiss. Anst. 14(Beih. 3): 5. 1896. Pteridium aquilinum subsp. caudatum (L.) Bonap., Notes Pterid. 1: 62. 1915. Lectotype (chosen by R. Tryon, 1941: 55). Probably a plant from West Indies, s. coll. s.n. (LINN 1246.15). Pteris aquilina L. var. mexicana Fe´e, Me´m. Foug. 7: 114. 1857. Syntypes. Mexico. [Veracruz:] Orizaba, Schaffner 136, 468; Huatusco, Schaffner 139 (P?, not found). Pteris caudata L. var. mexicana Fe´e, Me´m. Foug. 9: 8. 1857, nom. nud. Based on Mexico. [Veracruz:] Huatusco, Schaffner 140, Jalapa, Galeotti 6401 (BR!–3 sheets; specimen at P! is P. arachnoideum). For additional synonymy, see R. Tryon (1941).

Fronds 1.5–4 m long; blades lacking free lobes between ultimate segments; rachises glabrous to slightly pubescent; penultimate axes abaxially almost glabrous with hairs rare, curled, clear, with brown septa, others scattered, minute, less than 0.1 mm long, brown, clavate, adaxially with hairs rare, curled, clear, with brown septa only in medial grooves; costae flattened to slightly sulcate abaxially; laminar surfaces abaxially glabrous or with hairs scattered, 0.5 mm long, white, on the veins only, adaxially glabrous except for scattered, marginal, white hairs 0.5 mm long,

PTERIS

and minute, glandular, brown hairs less than 0.1 mm long, especially near segment tips; false indusia 0.3–0.5 mm wide with margins erose, eciliate; 2n⫽104 (Galapagos), 208 (see below).

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of stomates and indusia. One parent is thought to be P. arachnoideum and the other P. aquilinum var. pubescens, P. aquilinum var. pseudocaudatum, or some other taxon sharing genomic elements with those two varieties.

106. PTERIS Pteris L., Sp. Pl. 2: 1073. 1753. Lectotype (first chosen by J. Sm.?, Hist. Fil. 295. 1875): Pteris longifolia L. Litobrochia C. Presl, Suppl. Tent. Pterid. 148. 1836. Type: Litobrochia ampla (Kunze) C. Presl [⬅ Pteris ampla Kunze] ⫽ Pteris grandifolia L.

Distribution. Open, disturbed areas; 0–2500 m. USA (Fla); Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Berm, Bah, Gr & L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Galapagos, Peru. Selected Specimens Examined. BCS (Anthony 400, MEXU, NY, UC). Camp (Martı´nez S. 2934, MEXU). Chis (Ghiesbreght 237, NY). Dgo (McVaugh 11570, MICH). Gro (Hinton 14083, NY). Hgo (Fisher s.n., 14 Aug 1937, NY). Jal (Detling 883, ENCB). Mich (Rzedowski 46099, IEB). Mor (Sa´nchez 398, MEXU). Nay (McVaugh 18813, NY). NL (Pennell 16911, NY). Oax (Mickel 4176, NY). Pue (Turra 34, ENCB). QR (Cowan 3024, NY). Qro (Rzedowski 43861, ENCB). Rev (Felger 15873, UC). Sin (Pitelka 285, UC). SLP (Mickel 557, NY). Tab (Cantu´ 134, MEXU). Tam (Stanford et al. 2220, NY). Ver (Orcutt 3392, NY). Yuc (Ucan & Flores 988, MEXU).

Pteridium caudatum is distinguished by the lack of free lobes between ultimate segments, axes generally glabrous or only sparsely hairy, and indusia narrow, erose, and non-ciliate. This species varies from densely hairy to less commonly subglabrous abaxially. Glabrous or subglabrous specimens occur throughout the species’ range in Mexico. Thomson (2000) and Thomson and Alonso-Amelot (2002) reported that P. caudatum throughout Central and South America is a tetraploid, based on DNA studies and relative cell sizes

Terrestrial, less commonly epipetric; rhizomes short-creeping to erect, compact, with scales at apices; fronds medium-sized to very large, clumped, monomorphic; stipes usually glabrous; blades pinnate to several times pinnate, often with basal pinnae exaggerated basiscopically, blades often pentagonal to pedate; laminae chartaceous to coriaceous, minor axes mostly glabrous, generally with slender, antrorse awns on the adaxial axes at base of pinnae (rachis awns), pinnules, and ultimate segments (pinna awns); veins free or netted, without included veinlets; sori marginal, protected by differentiated recurved margins (false indusia), mostly continuous except at sinuses and segment apices; paraphyses usually present, mixed with sporangia; spores each generally with an equatorial ridge, perispore lacking; x⫽29. Pteris is a pantropical genus of about 250 species, of which only about 60 are American. It is distinguished by its generally large fronds, marginal sori, and spores with an equatorial flange. Awns, which occur adaxially on the blade axes of many species of Pteris, are most conspicuous in the more laminated parts of the blades. Molecular studies (Cranfill, unpubl.) suggest that the genus is polyphyletic. In particular, P. longifoliia and P. vittata are not closely related to the rest of Pteris, and probably not congeneric with the remaining Mexican species. References Scamman, E. 1961. The genus Pteris of Costa Rica. Rhodora 63: 194–205; Tryon, R. M. 1964b. Pteris. In: The Ferns of Peru. Contr. Gray Herb. 194: 188–210.

Key to the Mexican Species of Pteris 1. Blades 1-pinnate, or if more divided (2–3-pinnate) the suprabasal pinnae not pinnatifid; rhizome scales concolorous; blade awns absent (scales bicolorous and very small awns present in P. chiapensis). 2. Veins anastomosing; sterile blade margins entire or serrate. 3. Veins irregularly anastomosing towards the margins (outer 1⁄2 of ultimate segments); basal pinnae simple; rachis awns absent adaxially; margins entire; pinnae 25–45 ⫻ 3–4.5 cm; rhizome scales concolorous. ..................................................... 6. P. grandifolia. 3. Veins totally anastomosing; basal pinnae pinnate-pinnatifid or more divided; rachis awns present on rachises above basal pinnae; margins at sterile tips sharply serrate; pinnae 10–25 ⫻ 1.2–1.5 cm; rhizome scales bicolorous. ..................................3. P. chiapensis. 2. Veins free; sterile blade margins serrulate. 4. Basal pinnae with exaggerated basiscopic pinnules; blades deltate or ovate, broadest at bases; stipes glabrous, 1⁄2 or more of the frond length; pinna bases cuneate. ................................................................................................................................................. 4. P. cretica. 4. Basal pinnae not divided; blades tapering toward bases; stipes less than 1⁄2 the frond length, densely scaly; pinnae with cordate bases, each often with basiscopic auricle. 5. Pinnae (28–)32–70 pairs; pinnae articulate; indusia erose; terminal pinnae not strongly exaggerated. ........................... 7. P. longifolia. 5. Pinnae 11–22(–35) pairs; pinnae not articulate; indusia entire; terminal pinnae strongly exaggerated. ............................... 18. P. vittata.

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PTERIS

1. Blades more than 1-pinnate, the suprabasal pinnae pinnatifid (or more divided); rhizome scales usually bicolorous (concolorous in three species); pinna awns generally present (absent in P. pulchra). 6. Veins entirely free; rhizome scales concolorous or bicolorous to tricolorous; paraphyses present or absent. 7. Basal pinnae pinnate-pinnatifid beyond the basal pinnules; abaxial leaf surfaces with dense to scattered hairs 0.5–1 mm long. 8. Axes smooth abaxially; segments rounded or toothed at tips, not mucronate; blades membranaceous-chartaceous; paraphyses absent. ......................................................................................................................................................................... 9. P. muricella. 8. Axes with prickles abaxially; segments mucronate at tips, the midribs prolonged into segment tips; blades somewhat coriaceous; paraphyses abundant. ............................................................................................................................................... 8. P. muricata. 7. Basal pinnae only pinnatifid beyond the basal pinnules; abaxial leaf surfaces with scattered hairs 0.1 mm long. 9. Sterile veins ending before segment margins; paraphyses absent; indusia 0.8–1 mm wide; some veins arising from costae of pinnae; pinna awns blunt, 0.1 mm long. ........................................................................................................... 11. P. paucinervata. 9. Sterile veins extending to and fusing with segment margins; paraphyses present; indusia 0.3–0.5 mm wide; all veins arising from costules of segments, or some arising from costae of pinnae; pinna awns acuminate, 0.5–5 mm long. 10. All veins arising from costules of segments; indusia 0.3 mm wide; rhizome scales tricolorous, central portion lustrous, golden brown at bases, dark brown at tips, outer portion brown, erose. .............................................. 16. P. quadriaurita. 10. Some veins arising from costae of pinnae; indusia 0.3–0.5 mm wide; rhizome scales concolorous, golden brown. ................................................................................................................................................................................................... 15. P. pungens.

6. Veins netted, at least along the pinnule costules; rhizome scales bicolorous; paraphyses present. 11. Veins free except for costular areoles; sterile margins entire; indusia entire. ................................................................................. 2. P. biaurita. 11. Veins netted with several rows of areoles; sterile margins (at least segment tips) serrate or serrulate; indusia entire or erosefimbriate. 12. Pinna awns absent; blades strictly bipinnate. ............................................................................................................................ 14. P. pulchra. 12. Pinna awns present, 0.3–1 mm long; blades bipinnate to tripinnate-pinnatifid. 13. Costal areoles 2–5 between adjacent costules; indusia entire; pinna bases cuneate. ..................................................... 1. P. altissima. 13. Costal areole 1 between adjacent costules; indusia entire or erose-fimbriate; pinna bases cuneate or truncate. 14. Pinna bases truncate, sessile. 15. Apices of ultimate segments serrate, acute to acuminate; blades abaxially with terete, erect, golden brown hairs 0.5–0.8 mm long and flaccid or appressed hairs 0.1–0.3 mm long; sinuses broadly U-shaped to rounded; 800–2250 m; common; widespread in s. Mexico. ........................................................................ 12. P. podophylla. 15. Apices of ultimate segments entire or crenulate, obtuse to acute; blades abaxially subglabrous with scattered minute hairs 0.1–0.3 mm on veins; sinuses slender, V-shaped; 180 m; rare; Ver. ....................... 17. P. tripartita. 14. Pinna bases narrowly cuneate, petiolulate. 16. Blades abaxially glabrous; 0–500 m elevation. ................................................................................................ 13. P. propinqua. 16. Blades abaxially with flaccid or appressed hairs 0.3–0.8 mm long; 300–2700 m. 17. Indusia entire; rhizome scales 3–4 mm long; rachis awns absent or reduced to minute papillae. ..... 10. P. orizabae. 17. Indusia erose-fimbriate; rhizome scales 1–2 mm long; rachis awns present, 1 mm long. ........................... 5. P. erosa.

1. PTERIS ALTISSIMA Pteris altissima Poir. in Lamarck, Encycl. 5: 722. 1804. Type. Puerto Rico. Ledru s.n. (P-Lam.!). Figs. 255A, B. Pteris kunzeana J. Agardh, Recens. Spec. Pter. 62. 1839. Litobrochia kunzeana (J. Agardh) Fe´e, Me´m. Foug. 5: 135. 1852. Syntypes. French Guiana. Leprieur s.n. (G-Delessert); “ad Cayennam,” Richard s.n. (P); Peru, Matthews s.n. (K!, frag. NY), Poeppig [225] (K!; isosyntype B, frag. NY!). Litobrochia grandis Fe´e, Me´m. Foug. 8: 75. 1857. Syntypes. Mexico. [Veracruz:] Barranca de San Francisco, (Mirador), Schaffner 144 [1854] (P); Cuba. Morelet s.n. (P); Saint Dominique [Dominican Republic]. de Tussac s.n. (P). Pteris protea Liebm., Mexic. Bregn. 228 (reprint 76). 1849. Syntypes. Mexico. Veracruz: Colipa, Liebmann s.n. [Pl. Mex. 2728, Fl. Mex. 758, 759], Pital to Rı´o Nautla, [Pl. Mex. 2729, Fl. Mex. 760], between Colipa and Misantla [Pl. Mex. 2730, Fl. Mex. 557] (all C!; isosyntypes BM, K).

Rhizomes stout, erect; rhizome scales 1–2 ⫻ 0.5–0.8 mm, bicolorous, centers black, heavily sclerified, margins wide, brown, fringed; fronds 0.8–2.5 m tall; stipes ca. 1⁄2 the frond length, stramineous, glabrous; blades bipinnate to deeply pinnatepinnatifid, slightly pedate, pinna axils smooth; ultimate segments 3–6 cm long, bases cuneate, apices acute to acuminate, sterile margins serrate, the teeth slightly mucronate, sinuses symmetrical, adaxial surfaces glabrous, abaxial surfaces with scattered brown hairs 0.5 mm long along midribs; rachis awns absent or reduced to a minute bump; pinna awns 0.3–1 mm long; veins anastomosing with 2–5 areoles between costules, usually one row of areoles along costules, extending to base of sinuses; indusia 0.5 mm wide, whitened, entire; sporangia mixed with paraphyses 0.1–0.3 mm long; spores reddish brown; 2n⫽166 (Jam, Ver).

PTERIS CHIAPENSIS

Distribution. Wet montane forests; 50–1400 m. Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, s Braz, Bol. Selected Specimens Examined. Chis (Croat 40188, MEXU, MO; Purpus 6730, B, UC). Hgo (Copeland herb. 76a, UC; Estrada M. 182, ENCB). Oax (Martı´nez C. 412 MEXU, MICH, NY, UC; Mexia 9188, B, GH, MICH, NY, UC). Pue (Sa´nchez Mejorada & Cha´vez 638, ENCB, MEXU, US). SLP (King 4420, UC). Tab (Cowan et al. 3534, CSAT, ENCB, MEXU, UC). Ver (Conzatti 607, MEXU).

Pteris altissima is distinguished among the bi- or tripinnatifid awn-bearing taxa by the netted venation with 2–5 areoles between adjacent costules, parallel to costae. Smith (1981) reported intermediates between this species and Pteris orizabae from Chiapas, although none have been found elsewhere in Mexico.

2. PTERIS BIAURITA Pteris biaurita L., Sp. Pl. 2: 1076. 1753. Lectotype (chosen by Proctor, 1977: 145). Plumier, Descr. Pl. Ame´r., pl. 14, 1693 (Traite´ Foug. Ame´r., pl. 15. 1705), illustrating a plant from Martinique or Hispaniola. LINN 1246.19 was added after 1755, according to Jackson (1912). Figs. 254J, K. Rhizomes suberect, moderately stout; rhizome scales 2–2.5 ⫻ 1–1.2 mm, bicolorous, centers black, heavily sclerified, margins wide, brown, fringed; fronds 60–100 cm long; stipes 1⁄3–1⁄2 the frond length, stramineous, glabrous with occasional trichomidia; blades pinnate-pinnatifid, with one exaggerated basiscopic pinnule per basal pinna; pinnae sessile; segments to 17–20 ⫻ 5 cm, ultimate ones entire, obtuse, adaxial surfaces glabrous, abaxial surfaces glabrous except for occasional trichomidia 0.1 mm long; pinna awns lanceolate 0.1–0.3 mm; veins generally free except for single costal areoles between adjacent costules; indusia 0.5– 0.8 mm wide, entire; sporangia mixed with paraphyses 0.3–0.5 mm long; spores reddish brown; n⫽2n⫽58 (Jam), 87 (several Old World).

535

Distribution. Along river banks in wet lowland forests, lower montane rain forests, tropical deciduous forests, montane cloud forests, in ravines; 350–800 m. Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, s Braz, Bol; Old World tropics. Selected Specimens Examined. Chis (Breedlove 22573, 32837, DS). Gro (Lorea 2426, FCME; Vela´zquez 343, FCME). Nay (Ortega 46, US). Oax (Campos V. & Reyes 1423, MEXU; Conzatti 586, 1281, MEXU; Mickel 5166, ENCB, NY, UC; Morton & Makrinius 2383, MICH, US). Ver (Calzada 7534, XAL).

Pteris biaurita closely resembles P. quadriaurita in blade form, but is distinguished by the costal areoles.

3. PTERIS CHIAPENSIS Pteris chiapensis A. R. Sm., Proc. Calif. Acad. Sci., ser. 4, 40: 226. 1975. Type. Mexico. Chiapas: 26–28 km N of Ocozocoautla, Breedlove 22492 (DS!; isotype NY!). Figs. 259F–H, J. Rhizomes erect, ca. 3 cm diam.; rhizome scales bicolorous; fronds over 1 m long (complete specimen not seen), probably with stipes about equaling blades in length; stipes ca. 8 mm diam. at base, stramineous, glabrous; blades deltate, ca. 60 cm long, 2– 3-pinnate at bases, distally 2-pinnate, ultimately 1-pinnate; proximal pinnae largest, ca. 50 ⫻ 35 cm; ultimate segments 10–25 ⫻ 1.2–1.5 cm, narrowly cuneate and stalked to 3 cm at bases, entire, apices attenuate; awns of rachises and pinnae present, 0.5 mm long; venation netted, areoles in 2–3 series between midveins and margins; leaf tissue chartaceous, abaxially with scattered erect inconspicuous hairs ca. 0.1 mm; sori continuous along margins except at serrate apices.

536

PTERIS

Distribution. Terrestrial in montane rain forest; 700 m. Mexico. Specimens Examined. Chis (Breedlove 31237, DS). Gro (Hamilton & Rowell 3243, US; Hinton 14208, US; Lorea s.n., 3 Jul 1980, ENCB, FCME; Rzedowski 23584, ENCB, NY).

Pteris chiapensis is similar to P. pulchra except the blades are less divided (1-pinnate, basal pinnae pinnatifid or more divided), the pinnae smaller, 15–25 ⫻ 1.2–1.5 cm, and the apices long, sharply serrate (entire or minutely serrate in P. pulchra).

4. PTERIS CRETICA Pteris cretica L., Mant. Pl. 130. 1767. Lectotype (chosen by R. M. Tryon, Contr. Gray Herb. 194: 192. 1964). Crete. s. coll. s.n. (LINN 1246.7, photos A, K, SPF). Fig. 254A. Pteris triphylla M. Martens & Galeotti, Me´m. Foug. Mexique 51, pl. 14, f. 1. 1842, hom. illeg., non J. Agardh, 1839. Pteris trifoliata Fe´e, Me´m. Foug. 8: 114. 1857. Type. Mexico. Veracruz: Xalapa, Galeotti 6393 (BR!; US!, photo ex BR; isotype BR!).

Rhizomes small, ca. 0.5 cm diam., short-creeping; rhizome scales 2–3 ⫻ 0.5 mm, concolorous, dark brown, margins with long, scattered hairs; fronds 15–100 cm long, fertile longer than the sterile, clumped; stipes glabrous, 1⁄2–3⁄4 the frond length; blades deltate or ovate, pinnate to pedate with one pair of exaggerated basal basiscopic pinnules, terminal pinnae conform; pinnae linear, 80–150 ⫻ 7–12 mm, proximal ones sessile or shortpetiolate, distal ones long-decurrent on rachises, pinna bases cuneate, chartaceous to coriaceous, sterile pinna margins serrulate, with attenuate teeth, rarely crenulate, adaxial surfaces glabrous, abaxial surfaces glabrous with occasional brown hairs 1 mm long on midribs; awns absent; veins free; indusia 0.5 mm wide, entire; sporangia mixed with colorless, paraphyses 0.3 mm long; spores orange-tan; 2n⫽58, 87, 116, 174, 232 (all Old World), n⫽2n⫽58 (Taiwan, India, Italy, Japan, Nepal), 116 (Zimbabwe, Ascension Island).

Distribution. Moist, lightly wooded banks in mesic forests, montane rain forests, cloud forests, edge of streams in oak and pine forests; (700–)1400–2800 m. USA (Fla); Mexico; Guat, Hond, Salv, Nic, CR, Pan; Jam; Peru, Braz, Arg; Old World tropics and subtropics. Selected Specimens Examined. Chih (Bye 7510, MEXU). Chis (Breedlove 51936, CAS, ENCB). DF (Lyonnet 1485, MEXU, US). Dgo (Sa´nchez 745, MEXU). Gro (Hinton 14206, ENCB, US). Gto (Ventura & Lo´pez 9184, ENCB, IEB, MEXU). Hgo (Sa´nchez-Mejorada 168, MEXU, US). Jal (McVaugh 26166, MEXU, MICH). Me´x (Matuda 27441, MEXU, UC, US). Mich (Arse`ne 2427, MEXU). Mor Lyonnet 2878, FCME, UC, US). NL (Pringle 1985, MEXU, UC, US). Oax (Rzedowski 21027, MEXU, MICH, NY). Pue (Orcutt 4028, MEXU, US). Qro (Rzedowski 42554, IEB, MEXU, XAL). Sin (Lehto 24378, ARIZ, ENCB). SLP (Rzedowski 10571, ENCB, MEXU). Son (Turner & Martin 79-379, MEXU). Tam (Diggs & Nee 2409, XAL). Ver (Palacios-Rios & Herna´ndez M. 535 UAMIZ).

Tryon (1964) and Smith (1981) considered this species to be native to the Old World and perhaps to Mexico, but introduced elsewhere. Pteris cretica is distinct in its 1-pinnate blades, free veins, and exaggerated basiscopic pinnules.

5. PTERIS EROSA Pteris erosa Mickel & Beitel, Mem. New York Bot. Gard. 46: 322–323. 1988. Type. Mexico. Oaxaca: Distrito Juquila, 7–13 km N of San Gabriel, Mickel 6138 (NY!). Figs. 255C–E. Rhizomes short-creeping to erect, compact; rhizome scales 1–2 0.8 mm, strongly bicolorous, each with a black central streak, heavily sclerotic, margins brown, crisped, erose; fronds 40–90 cm long, clumped; stipes ca. 1⁄2 the frond length, stramineous, glabrous, about as long as the blades; blades pinnate-pinnatifid above the base, the basal pair of pinnae pinnate-pinnatifid; pinnae petiolulate 3–7 mm to sessile, cut 3⁄4 to the costae, lobes falcate, bases cuneate, narrowly decurrent (on acroscopic side), with short, jointed hairs on petiolules, margins entire, sterile tips serrulate, apices with long, undivided portion, adaxial surfaces glabrous, abaxial surfaces glabrous with flaccid or appressed, scattered hairs 0.5–0.8 mm long on veins; rachis awns 1 mm long with several teeth, absent on rachis at basal pinna; pinna awns 0.8–1 mm long; venation netted with one long areole between ⫻

PTERIS LONGIFOLIA

537

adjacent costules; indusia 0.3 mm wide, margins erose-fimbriate with scattered, short, jointed hairs on indusial surfaces; sporangia mixed with paraphyses 0.3 mm long; spores brown.

Distribution. Roadside ditches and margins of wet forests; 0– 1150 m. Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Ec, Peru, Braz. Distribution. Wet montane forests; 900–1800(–2300) m. Mexico; Guat. Selected Specimens Examined. Gro (Rzedowski 18631, ENCB). Jal (McVaugh 23376, ENCB, MEXU, MICH; McVaugh & Koelz 1255, MEXU, MICH). Me´x (Hinton 6049, NY, US; Moreno 161, ENCB, IEB). Mor (Lyonnet 550800005, MEXU). Nay (Gonza´lez & Vela´zquez 861, MEXU, UAN; McVaugh 18982, MEXU). Oax (Campos 1205, MEXU; Mickel 6142, NY, UC). Unverified, Doubtful, or Mistaken Reports. Tab (reported by Magan˜a, 1992, but not verified).

Pteris erosa differs from P. podophylla and P. orizabae by the characters in the key and from P. propinqua J. Agardh in lacking the decurrent pinna bases and having fimbriate indusial margins.

6. PTERIS GRANDIFOLIA Pteris grandifolia L., Sp. Pl. 2: 1073. 1753.

Selected Specimens Examined. Chis (Breedlove 22231, DS, MEXU). Gro (Kruse 611, ENCB). Hgo (Gimate 929, MEXU). Mich (Hinton et al. 16025, UC, US). Oax (Mickel 5806, ENCB, MEXU, NY, UC). Pue (Knobloch 700, US). QR (Aguilar & Corte´s 504, XAL). Qro (Rzedowski 42723, ENCB). SLP (Copeland herb. 84, BM, MEXU, MICH, NY, UC, US). Tab (Cowan et al. 3516, CAS, UC). Ver (Ventura A. 11462, ENCB). Unverified, Doubtful, or Mistaken Reports. Camp (Martı´nez S. et al. 28082, MEXU, XAL, cited by Palacios Rios, 2002a, but not verified.).

This large, strictly 1-pinnate species is free-veined in the central portion of each pinna, but is net-veined in the outer half of the pinnae toward the margins.

7. PTERIS LONGIFOLIA Figs. 254E, F.

Litobrochia grandifolia (L.) J. Sm., J. Bot. (Hooker) 4: 163. 1841. Heterophlebium grandifolium (L.) Fe´e, Me´m. Foug. 5: 140, t. 11A, f. 9–12. 1852. Lectotype (chosen by Proctor, Fl. L. Antill. 2: 143. 1977). Plumier, Traite´ Foug. Ame´r., pl. 105, 1705. LINN 1246.1 was added after 1755, according to Jackson (1912).

Rhizomes ca. 1 cm diam., creeping; rhizome scales dimorphic in width, 5–6 ⫻ 0.1–1 mm, concolorous, thin, tan to gray-brown; fronds 1–5 m tall; stipes ca. .1⁄3 the frond length, stramineous, glabrous; blades 1-pinnate, chartaceous to firm; pinnae entire, 25– 45 cm ⫻ 30–45 mm, bases cuneate, margins entire, hairs in axils elongate and moniliform, both surfaces glabrous; awns absent; veins netted in outer half of pinnae or only near the margins, or only casually anastomosing; indusia 1–1.3 mm wide, entire; sporangia mixed with paraphyses 0.3 mm long; spores tan to brown; 2n⫽116 (Jam, SLP).

Pteris longifolia L., Sp. Pl. 2: 1074. 1753. Lectotype (chosen by Proctor, Fl. L. Antill. 2: 141. 1977). Plumier, Descr. Pl. Ame´r., pl. 18, 1693 (Traite´ Foug. Ame´r., pl. 1705), illustrating a plant from Haiti, near Port de Paix. LINN 1246.2 was added to LINN after 1755 (Jackson, 1912). Figs. 254B, 259D, E. Pteris aequalis C. Presl, Reliq. Haenk. 1(1): 54. 1825. Type. Mexico. Haenke s.n. (PR!).

Rhizomes short-creeping to suberect, densely clothed with scales at apices; rhizome scales dimorphic, 5–7 mm long, grading to hairs 2 mm long, concolorous, yellow to light brown; fronds 20–100 cm long, clumped; stipes less than 1⁄2 the frond length, stramineous with dense pale scales and hairs; blades 1-pinnate, not pedate, tapering proximally, chartaceous to coriaceous; pinnae (28–)32–70 pairs, articulate, auriculate, entire to crenulate, adaxial surfaces glabrous or with scattered hairs on costae, abaxial surfaces, with dense clear hairs 0.8–1 mm long on veins; awns absent; veins free; indusia 0.5–0.8 mm wide, erose; sporangia mixed with paraphyses 0.3 mm long; spores cream to tan; 2n⫽116 (Grand Cayman Island, Jam, Peru).

538

PTERIS

running to sinuses; indusia 0.3–0.5 mm wide, entire; sporangia mixed with paraphyses 0.3 mm long; spores brown.

Distribution. Rocky roadbanks in mesic to dry areas; 50–1900 m. USA (Fla); Mexico; Guat, Bel, Hond; Bah, Gr & L Ant; Ven, Trin. Selected Specimens Examined. Chis (Breedlove 33062, DS, ENCB). Col (Lo´pez F. et al. 627, ENCB, UAMIZ). Gro (Rzedowski 22785, ENCB, MEXU). Me´x (Hinton 3961, MEXU). Mich (Soto 7215, MEXU). Mor (Mendoza 48, UAMIZ). Nay (Maltby 86, US). Oax (Mickel 4012, ENCB, MEXU, NY, UC, US). Pue (Knobloch 706, US). QR (Torres 58, MEXU). Qro (Zamudio Ruı´z 6152, IEB). Sin (Herna´ndez & Herna´ndez 523, ENCB). SLP (Pringle 3359, B, MEXU, UC, US). Tab (Matuda 3535, US). Tres Marı´as Islands (Nelson 4201, US). Ver (Ventura A. 8231, ENCB).

Distribution. Wet montane forests on Atlantic slope, evergreen cloud forests; 2020–2700 m. Mexico; Guat, Nic, CR, Pan; Col, Ven, Ec, Peru, Bol. Selected Specimens Examined. Chis (Breedlove 31979, DS, MEXU, 32359, DS, ENCB; Martı´nez S. et al. 13221, MEXU; Matuda 2065, MEXU, 5433, US; Ventura & Lo´pez 3315, ENCB, MEXU). Oax (Campos V1577, MEXU; Mickel 4546, ENCB, MEXU, NY, UC, US). Ver (Copeland herb. 82, MEXU, MICH, NY, UC, US, XAL; Ventura 15768, ENCB, MEXU).

Unverified, Doubtful, or Mistaken Reports. Camp (Martı´nez S. et at. 28081, MEXU, XAL, cited by Palacios-Rios, 2002a, but not verified). NL (reported by Aguirre Clavera´n, 1983, but cited plants are P. vittata).

This rare species is the only Mexican Pteris to bear prickles on the blade axes.

This 1-pinnate, free-veined, awnless species is abundantly covered with scales and hairs on the stipes and rachises and has erose indusia. It is closely related to the Old World P. vittata, which has escaped from cultivation in many parts of the New World (Morton, Amer. Fern J. 47: 7–14. 1957), but it has not been reported previously in Mexico or Central America (Smith, 1981; Stolze, 1981). Pteris vittata can be distinguished from P. longifolia in having fewer pinnae (11–22(–35) pairs vs. (28–)32– 70 pairs), pinnae not articulate (vs. articulate), indusia entire (vs. erose), and terminal pinnae more strongly developed.

9. PTERIS MURICELLA

8. PTERIS MURICATA Pteris muricata Hook., Sp. Fil. 2: 193, t. 123B. 1858. Type. Colombia. Antioquia: Jervise (K!). Figs. 254C, D. Rhizomes stout, suberect; rhizome scales 2–4 ⫻ 0.5 mm, bicolorous, centers black, sclerotic, margins narrow, brown; fronds 50–200 cm long; stipes ca. 1⁄2 the frond length, stramineous, with scattered scales, hairs 0.5 mm, and conical prickles 0.5–1 mm; blades 3–4-pinnate, pedate-pinnate, basal pinnae pinnatepinnatifid beyond the basal pinnules, firm to coriaceous; pinnae petiolulate to 5 mm; rachis awns short, broad, toothed, 1–1.5 mm long; ultimate segments entire, mucronate, serrate, segments broadly adnate, with scattered 0.5–1 mm hairs on abaxial costae, sinuses asymmetric; pinna awns 0.8–1 mm long; veins free, not

Pteris muricella Fe´e, Me´m. Foug. 8: 73. 1857. Syntypes. Mexico. Veracruz: “pre`s de Co´rdoba et de Huatusco,” 1854, Schaffner 143 (RB); Schaffner s.n. (P!). Figs. 257C, D. Hypolepis pteridiodes Hook., 2 Cent. T. 59. 1861. Pteris pteridioides (Hook.) Ballard, Kew Bull. 1937: 348. 1937. Type. Peak of Fernando Po, 7000', Gustav Mann 348 (K). Pteris mollis Christ, Bull. Herb. Boissier 4: 658. 1896. Type. Costa Rica. “Foreˆts de San Marcos,” Tonduz 7565 (isotypes P, US).

Rhizomes erect, caudices 1.5–2 cm diam.; rhizome scales 5–6 ⫻ 0.8–1 mm, bicolorous, centers dark brown, margins narrow, light brown; fronds ca. 1 m long; stipes ca. 1⁄2 the frond length, light reddish brown at bases, smooth; blades pedate, 3-pinnate to 3pinnate-pinnatifid, pinnate-pinnatifid distally, basal pinnae pinnate-pinnatifid beyond the basal pinnule, chartaceous; ultimate segments obtuse, crenate, adaxial surfaces glabrous, abaxial surfaces with hairs 0.5–0.8 mm long along midribs; rachis awns 1–1.5 mm long; pinna awns 1.5–2 mm long; veins free, not running to bases of sinuses; indusia 0.3–0.5 mm wide, entire; sporangia without paraphyses; spores cream to tan; 2n⫽58 (Chis).

PTERIS PAUCINERVATA

Distribution. Wet montane forests, lower montane rain forest; (800–)1200–2100 m. Mexico; Guat, Salv, Nic, CR, Pan; Ven; tropical Africa and Madagascar. Selected Specimens Examined. Chis (Breedlove 33076, 34408, DS, ENCB, 42015, DS, MEXU; Matuda 233, MEXU, US). Gro (Hinton 14983, US; Lorea 3022, FCME). Oax (Maya J. 1514, CHAPA, MEXU; Wendt et al. 4702, MEXU). Ver (Finck 19, GH, MEXU, MO, NY, US; Purpus 7764, GH, MO, NY, UC).

10. PTERIS ORIZABAE Pteris orizabae M. Martens & Galeotti, Me´m. Foug. Mexique 53, pl. 13. 1842. Figs. 255F, G. Pteris apicalis Liebm., Mexic. Bregn. 230 (reprint 78). 1849, nom. superfl. Litobrochia orizabae (M. Martens & Galeotti) Fe´e, Me´m. Foug. 5: 135. 1852. Type. Mexico. Veracruz: Orizaba, Vaquerı´a de Jacal, Galeotti 6252 [1840] (BR!, photo BM!, photo US!; isotype P!). Litobrochia setifera Fe´e, Me´m. Foug. 5: 138. 1852. Type. Mexico. “Super montem ignivomum S. Martini-Crate`re del Cerro, San. Martin,” Galeotti 6571 (P?, not found; isotypes BR!–4 sheets, K!, frag. NY!). Litobrochia hemipteris Fe´e, Me´m. Foug. 8: 76. 1857. Pteris hemipteris (Fe´e) Hook., Sp. Fil. 2: 224. 1858. Type. Mexico. [Distrito Federal/Edo. Me´xico:] “Valle´e du Mexique,” Schaffner s.n. (P?, not found).

Rhizomes erect, stout; rhizome scales 3–4 ⫻ 0.3–1 mm, bicolorous, centers black, sclerotic, margins light, brown, fimbriate; fronds 1–2 m long; stipes ca. 1⁄2 the frond length, stramineous, glabrous; blades bipinnate-pinnatifid; pinnae petiolulate to 6 mm, bases long-cuneate; ultimate segments acute, margins and mucronate teeth thickened, adaxial surfaces glabrous, margins with occasional hairs, abaxial surfaces pubescent with hairs 0.3– 0.8 mm, hairs usually flaccid or appressed; rachis awns absent or reduced to minute papillae; pinna awns 0.5–0.8 mm long; veins netted, ending before bases of sinuses, one long areole between adjacent costules; indusia 0.8 mm wide, entire, with flange 0.3 mm wide along recurved margins, with scattered hairs or glabrous; sporangia mixed with paraphyses 0.5–0.8 mm long; spores reddish brown; 2n⫽ca. 116 (Chis).

539

Distribution. Margins of wet montane forests, oak forests; 300– 2700 m. Mexico; Guat, Hond, Salv, Nic. Selected Specimens Examined. Chis (Breedlove 34409, DS, ENCB). DF (Lyonnet 3310, MEXU, US). Gro (Lorea 2294, FCME). Hgo (Moore 5280, MEXU, US). Jal (McVaugh 13918, MEXU, MICH, US). Me´x (Arreguı´n 503, ENCB, MEXU). Mich (Santos Martı´nez 2108, IEB, MEXU). Mor (Lyonnet 1361, US). Oax (Mickel 4384, ENCB, NY). Pue (Tenorio L. 15414, MEXU). Qro (Rzedowski 45210, IEB). Ver (Ventura A. 272, ENCB).

Pteris apicalis Liebm. was an avowed substitute name for P. orizabae, with the same type, and therefore superfluous. Pteris apicalis was substituted for P. orizabae because Liebmann protested in a vitriolic manner (it turns out, wrongly) against calling this species “orizabae” as it “is not found at all on Volcan Orizaba” in Veracruz. It is an interesting example of what low esteem Liebmann had for the work of Martens and Galeotti, as shown by his statement: “It is deplorable that the authors should have had so slight regard for science, the truth and themselves, that rebukes of this sort are needed” (translation). Pteris orizabae is closely related to P. podophylla and P. erosa (q.v.) but differs in larger plant size, entire indusial margins, and more easterly distribution. It is also related to P. propinqua, which has cuneate pinna bases that become decurrent basiscopically (vs. only acroscopically). Pteris propinqua also has glabrous abaxial costal surfaces (vs. bearing jointed golden hairs). Stolze (1981) incorrectly synonymized P. orizabae under P. polita Link, the type (frag. NY!) of which has free veins.

11. PTERIS PAUCINERVATA Pteris paucinervata Fe´e, Me´m. Foug. 8: 73. 1857. Type. Mexico. Veracruz: “pre`s de Mirador, a` la Barranca de San Martin,” Schaffner 152 (P!; isotypes B, frag. NY!, GH, RB). Figs. 257A, B. Rhizomes short, erect; rhizome scales 3–4 ⫻ 0.8–1 mm, concolorous, golden brown; fronds 60–150 cm long; stipes ca. 1⁄2 the frond length, reddish brown, glabrous; blades pinnate to bipinnate-pinnatifid, not pedate, basal basiscopic pinnules pinnatifid; ultimate segments entire to crenate, 6–12 mm wide, obtuse, chartaceous to firmly herbaceous, decurrent, adaxial

540

PTERIS

surfaces glabrous, abaxial surfaces with scattered, short hairs 0.1 mm long, sinuses asymmetrical; rachis awns reduced to 1–2 round papillae 0.1 mm long; pinna awns blunt, 0.1 mm long; veins free, not running to sinuses, sterile veins ending before margins, some veins arising from costae of pinnae; indusia 0.8–1 mm wide, entire; sporangia without paraphyses; spores reddish brown.

margins, 1 costal areole between adjacent costules, more than one costal areole at sinuses; indusia 0.8 mm wide, entire, surface with occasional hairs and short flaps; sporangia mixed with paraphyses 0.5–0.8 mm long; spores reddish brown.

Distribution. Margins of wet montane forests, tropical montane forests; 800–2250 m. Mexico; Guat, Salv, Nic, CR, Pan; Gr, Ant; Col, Ven, Sur, Ec, Peru, s Braz, Bol. Distribution. Wet montane forests, montane cloud forests, pineoak-Liquidambar forests; (120–)800–2600 m. Mexico; Guat, Hond, Nic, CR, Pan. Selected Specimens Examined. Chis (Breedlove 33111, DS, MEXU, 46196, CAS, ENCB; Martı´nez S. 19642, MEXU, XAL; Matuda 2949, MEXU, NY, US, 5389, MEXU, 5449, US). Gro (Campos 1660, FCME; Lorea 1171, ENCB, FCME, MEXU). Ver (Finck 185, UC.

Selected Specimens Examined. Chis (Breedlove 32365, DS; Mu¨nch 145, DS). Gro (Lorea 2040, FCME, US; Martı´nez S. 996, IEB, MEXU). Jal (Pichardo 303, CHAPA, IEB, NY). Oax (Martı´nez C. 845, CAS, MEXU, NY; Mickel 1097, ENCB, MEXU, NY, UC). Pue (Riba 84, MEXU). Ver (Lira 204, IEB, MEXU, UAMIZ).

Pteris paucinervata resembles P. pungens in having some veins arising from the pinna costae, but is distinct by the reduced rachis awns and veins ending before the segment margins.

Pteris podophylla is closely related to P. erosa and P. orizabae; the characters in the key distinguish the three. The abaxial surfaces of P. podophylla, with erect, terete, concolorous hairs, are clearly different from surfaces of P. orizabae, which have long, flaccid hairs or are glabrous.

12. PTERIS PODOPHYLLA

13. PTERIS PROPINQUA

Pteris podophylla Sw., J. Bot. (Schrader) 1800(2): 67. 1801. Figs. 255H, J. Lonchitis pedata L., Sp. Pl., ed. 2, 2: 1536. 1763, non Pteris pedata L., 1753. Type. Jamaica. Browne s.n. (LINN 1249.1; Browne, Civ. Nat. Hist. Jamaica, t. 1, f. 1, 2. 1756). Pteris inflexa Copel., Univ. Calif. Publ. Bot. 19: 289. 1941. Type. Mexico. Veracruz: Orizaba, Copeland herb. 79 (US!, frag. NY!; isotypes GH, MEXU, MICH, UC!, US!).

Rhizomes stout, erect, massive; rhizome scales bicolorous, centers black; fronds 70–200(–300) cm tall, clumped; stipes ca. 1⁄2 the frond length, stramineous to castaneous; blades pedate, pinnate-pinnatifid above the basal pinna pair; suprabasal pinnae truncate, sessile; bases of penultimate segments abrupt to cuneate, not decurrent; fertile sinuses broadly U-shaped (nearly round); segments sessile, falcate, serrate, with recurved, sharp-pointed teeth, adaxial surfaces glabrous, abaxial surfaces with long, brown, flaccid or appressed hairs 0.5–0.8 mm long, trichomidia 0.1–0.3 mm long, and erect, terete, golden brown hairs 0.5–0.8 mm long; rachis awns absent or replaced by hairs; pinna awns 1 mm long; veins netted, 1–2 rows of areoles between midribs and

Pteris propinqua J. Agardh, Recens. Spec. Pter. 65. 1839. Lectotype (chosen by R. Tryon, 1964). Jamaica. Bancroft s.n. (K!). Figs. 258E, F. Rhizomes erect to short-creeping; rhizome scales bicolorous; fronds 50–150 cm; stipes ca. 1⁄2 the frond length, glabrous or minutely hairy, gray-brown; blades 30–120 cm wide, tripartite; basal pinnae pinnate-pinnatifid above the basal pinnules, basal basiscopic pinnules not exaggerated, pinnatifid; suprabasal pinnae 4– 8 pairs, 2–4 cm wide, pinnatifid, pedicellate, the pedicels 1–2.5 cm, bordered by a narrow green wing decurrent onto the rachis; ultimate segments slightly falcate, tips acute, serrate; rachises and costae unarmed with dark, tortuous hairs, awns ca. 0.5 mm; blade tissue chartaceous, glabrous; veins netted, one costal areole between adjacent costules; sori continuous along margins, interrupted at segment apices.

PTERIS PUNGENS

Distribution. Secondary forests, shores of lakes; 0–500 m. Mexico; Hond, Nic, CR, Pan; Jam; Col, Ven, Guy, Sur, Fr Gui, Ec, Peru, s Braz, Bol. Specimen Examined. Ver (Palacios-Rios 3221, XAL).

The only specimen cited for Mexico occurs at low elevation typical of P. propinqua, but lacks the basiscopically decurrent pinna bases typical of Pteris propinqua, and the golden hairs of the abaxial pinna costae of P. orizabae.

541

Distribution. Margins of wet lowland forests, pine-oak forests, montane rain forests; 100–1000 m. Mexico; Guat, Nic; Col. Selected Specimens Examined. Chis (Breedlove 33075, DS). Gro (Lorea 2041, FCME, US). Hgo (Copeland herb. 77, MEXU, MICH, P, UC, US; Rzedowski 32476, ENCB). Oax (Hallberg 1630, ENCB, NY, UC, US). Pue (Lo´pez-Forment 142, MEXU; Ventura A. 20211, ENCB). Qro (Rubio 2705, CAS, IEB). SLP (Rzedowski 10967, ENCB). Ver (Chazaro et al. 3671, ENCB).

Pteris pulchra is distinct by its strictly bipinnate blades, netted veins, and lack of awns adaxially.

14. PTERIS PULCHRA 15. PTERIS PUNGENS Pteris pulchra Schltdl. & Cham., Linnaea 5: 614. 1830. Type. Mexico. Veracruz: “Inter Misantlam et Colipam, regionis subcalidae” Schiede & Deppe 787 (B!–4 sheets; isotype NY! frag. ex B). Figs. 254G, H. Pteris arborescens M. Martens & Galeotti, Me´m. Foug. Mexique 54. 1842. Type. Mexico. Oaxaca: Chinantla, Teotalcingo, Galeotti 6375 (BR!; isotypes B!, BR!). Litobrochia mexicana Fe´e, Me´m. Foug. 5: 136. 1852. Pteris mexicana (Fe´e) E. Fourn., Mexic. Pl. 1: 117. 1872. Type. Mexico. Oaxaca: Chinantla, Teotalcingo, Galeotti 6376 (P!; isotypes B!, BR!–6 sheets, K!, US!).

Rhizomes stout, suberect; rhizome scales bicolorous; fronds 1– 2 m long; stipes ca. 1⁄3–1⁄2 the frond length, stramineous, glabrous; blades bipinnate with exaggerated basiscopic pinnules, chartaceous to firm; pinnules lanceolate, 90–150 ⫻ 12–22 mm, bases cuneate, margins entire, apices serrulate, adaxial surfaces glabrous, abaxial surfaces with hairs 0.3 mm long on midveins, otherwise glabrous; awns absent; veins netted throughout; indusia 0.5 mm wide, entire; sporangia mixed with paraphyses 0.1 mm long; spores dark brown.

Pteris pungens Willd., Sp. Pl., ed. 4, 5(1): 387. 1810. Lectotype (chosen by Proctor, Fl. L. Antill. 2: 144. 1977). Plumier, Traite´ Foug. Ame´r., pl. 14, 1705, illustrating a plant from Haiti, along La Grande Rivie`re, near Leogane. Figs. 257E–G. Rhizomes erect, ca. 2–3 cm diam.; rhizome scales concolorous, golden brown; fronds to 1.4(–2) m long; stipes ca. 1⁄2–2⁄3 the frond length, weakly aculeate, glabrous, scaly, stramineous to black; blades 30–60 ⫻ 30–50 cm, pentagonal-oblong, pinnatepinnatifid with terminal pinnae conform, pinnatifid, basal pinnae with basal basiscopic pinnules elongate and deeply pinnatifid; pinnae 20–35 ⫻ 4–8 cm, axils aculeolate, tips abruptly caudate; segments linear-oblong, slightly oblique, straight or subfalcate, those at bases of pinnae evenly reduced (sometimes obsolete on basiscopic side), margins serrate near apices, glabrous adaxially, with sparse hairs 0.1 mm long abaxially; pinna awns 0.8–1 mm long; veins free, 1–2 basal ones arising from costae; indusia 0.3–0.5 mm wide, entire; sporangia with paraphyses; spores reddish brown.

542

PTERIS

long; veins free, running to margins; indusia 0.3 mm wide, entire, paraphyses present; spores dark brown, of varying sizes; 2n⫽58 (Ceylon, Jam), 116 (SLP, Jam), n⫽2n⫽87 (Jam, Nepal).

Distribution. In wet lowland forests, evergreen rain forests; 0– 820 m. Mexico; Guat, Bel, Hond, Nic, Cr, Pan; Gr & L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Bol. Selected Specimens Examined. Chis (Breedlove 34067, DS; Martı´nez S. 11712, 14687, 15452, MEXU; Purpus 6729, B, UC, US; Sohns 1693, US). Oax (Herna´ndez G. 686, 851, CAS, MEXU, NY). Ver (Croat 40059, UC; Riba & Pe´rez 844, XAL).

Distribution. Wet lower montane forests, pine-oakLiquidambar forests, coffee plantations; 450–1400 m. Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Bol, n Arg; Africa, Asia.

Pteris pungens closely resembles P. quadriaurita in blade form but is distinguished by the concolorous rhizome scales, the caudate pinna apices, basal veins sometimes arising from the pinna costae, and decurrent terminal pinnae. A specimen from Chiapas (Breedlove 22661, NY) has malformed spores and is intermediate between Pteris pungens and P. paucinervata. It has the general blade dissection, paraphyses, and narrow indusia of Pteris pungens, and the asymmetric sinuses and short caudate pinnae of P. paucinervata. Some veins end before the margins, whereas others run to the margins.

Selected Specimens Examined. Chis (Breedlove 32167, DS, ENCB). Col (Sanders et al. 10480, MEXU, UC). Gro (Mexia 8895, B, UC, US). Hgo (Rzedowski 32468, ENCB). Jal (Mexia 1692, CAS, UC, US). Me´x (Matuda 30501, MEXU, US). Mich (Ventura A. 4434, IEB). Mor (Lyonnet 3147, MEXU, US). Nay (McVaugh 16526, US). Oax (Mickel 1069, MEXU, NY, US). Qro (Sotero S. 809, IEB). SLP (Pringle 3971, DS, MEXU, UC, US). Tam (Breedlove & Anderson 63689, CAS). Ver (Purpus 2842, NY, UC).

16. PTERIS QUADRIAURITA Pteris quadriaurita Retz., Observ. Bot. 6: 38. 1791. Type. Sri Lanka. Ko¨nig s.n. (LD). Figs. 254L, M. Pteris plumula Desv., Me´m. Soc. Linn. Paris 6: 297. 1827. Type. “Antilles.” s. coll. s.n. (P-Desv., photo GH). Pteris nemoralis Willd. var. major M. Martens & Galeotti, Me´m. Foug. Mexique 53. 1842. Type. Mexico. [Veracruz:] Zacuapan, Galeotti 6291 (BR!–6 sheets). Litobrochia galeottii Fe´e, Me´m. Foug. 8: 75. 1857. Pteris galeottii (Fe´e) Hook., Sp. Fil. 2: 204. 1858. Type. Mexico. Oaxaca: “Teotalingo,” Galeotti 6485 (P).

Rhizome erect or suberect; rhizome scales 3–4 ⫻ 0.5 mm, tricolorous, central portion lustrous, bases golden brown, tips dark brown, outer portion brown, erose; fronds 50–200 cm long; stipes ca. 1⁄2 the frond length, stramineous, glabrous; blades pinnatepinnatifid to bipinnate, sinuses symmetrical, the sinus bases often with one to several teeth, either on adaxial or abaxial surfaces or both; ultimate segments entire, obtuse, adaxial surfaces glabrous, abaxial surfaces with short hairs 0.1 mm long; rachis awns reduced to 1–2 papillae, 0.1 mm long, pinna awns 0.5–5 mm

This is the most common species with pinnate-pinnatifid blades, exaggerated basiscopic pinnules, and free veins. It resembles Pteris biaurita and P. pungens, which see for further discussion. The spores of all the specimens exhibit great heterogeneity, with some spores two to three times larger than others. Pringle 3971 from San Luis Potosı´ and Breedlove 22175, 32167, 33060, 33157 (DS), and 56562 (CAS) from Chiapas, have scaly buds on the adaxial side of the rachises, but in all other respects resemble nonproliferous specimens.

17. PTERIS TRIPARTITA Pteris tripartita Sw., J. Bot. (Schrader) 1800(2): 67. 1801. Figs. 256A–E. Litobrochia tripartita (Sw.) C. Presl, Suppl. Tent. Pterid. 150. 1836. Type. Java. Thunberg s.n. (UPS-Thunb.-24968, 24969).

Rhizomes erect, massive; rhizome scales bicolorous; fronds 0.3– 2.5 m; stipes ca. 2⁄3 the frond length, glabrous, unarmed, stramineous to brown; blades to 50 cm wide, tripartite, pinnatepinnatifid above the basal pinna pair, basal pinnae pinnatifid beyond the first 1–2 basiscopic pinnules, these exaggerated and pinnatifid; suprabasal pinnae 13–33 pairs, 1–5 cm wide, pinnatisect, the bases truncate, sessile; ultimate segments (2–)4–5 mm wide in the middle, erect or slightly falcate, the apices obtuse to

18. PTERIS VITTATA

543

acute, entire or crenulate, or serrulate in sterile segments, chartaceous, subglabrous with sparse minute surface hairs 0.1–0.3 mm; awns often reduced to papillae; veins netted, with one areole between adjacent costules, free near margins; sori continuous along the margins, interrupted at segment apices; indusia entire, paraphyses present; spores reddish brown; 2n⫽116 (CR, Trin), 2n⫽232 (Malaya, Java, N. Guinea, Australia).

Distribution. In calcareous soils; low elevations. Mexico; Hond, Salv, Pan; Gr & L Ant; Ven, Peru, Braz, Arg; Old World. Selected Specimens Examined. NL (Aguirre 574, 575, 576, ENCB; Barkley 14555, US; Clausen 7640, US; Fairey T311M221, UC; Flores F. s.n., ENCB; Knobloch 2013, US; Kruckeberg 4850, NY; Rodriguez L. 82, NY). Tam (Trejo Avila s.n., 19 Oct 1985, UNL).

Distribution. Riverbanks, montane forests; 180 m. Mexico; CR, Pan; Gr & L Ant; Col, Ven, Guy, Sur, Fr Gui, Ec, Peru, Bol; Old World tropics. Specimen Examined. Ver (Sinaca Colin 1506, UAMIZ), possibly naturalized.

This rare species is distinct by its single areole between costules, truncate pinna bases, and the basal segments of the pinnae being auriculate acroscopically, occasionally also basiscopically. The nearest records of this species are from Costa Rica and Jamaica.

18. PTERIS VITTATA Pteris vittata L., Sp. Pl. 2: 1074. 1753. Type. China. Osbeck s.n. (LINN-1246.3). Figs. 259A–C. Rhizomes ascending or erect; rhizome scales yellow, 5–7 mm long; fronds 25–80 cm, clumped; stipes 1⁄8–1⁄6 the frond length, ca. 10 cm long, densely scaly; blades pinnate, gradually reduced proximally, widest above the middle, (6–)12–25 cm wide; pinnae 11–22(–35) pairs, linear, the bases slightly auriculate both acroscopically and basiscopically, not articulate, sterile margins serrulate near apices, terminal pinnae strongly exaggerated; rachises and costae sparsely scaly, stramineous to yellow-brown, lacking awns adaxially, abaxial laminae glabrous between costae and margins; veins free; sori continuous along margins, interrupted at pinna apices; indusia entire; spores pale orange; 2n⫽116 (Sri Lanka, Ghana, Cameroon, Natal, Nepal, W. Himalaya, N. Guinea, Philippines).

Unverified, Doubtful, or Mistaken Reports. QR (Castillo R. & Tun G. 247, UADY, XAL, cited by Palacios-Rios, 2002c, but not verified). Yuc (Palacios-Rios et al. 3688, XAL, cited by Palacios-Rios, 2002b, but not verified).

Pteris vittata is native to eastern Asia and is frequently introduced and naturalized in the New World tropics and subtropics. It can be confused with P. longifolia, which see for distinguishing characters. Excluded Species Pteris tremula R. Br., Prodr. 154. 1810. Type. Australia. Port Jackson, Hunter’s River or Coal River, no specimen cited, “seen alive.” Figs. 258A–D. Rhizomes compact, erect; rhizome scales linear-lanceolate, concolorous, black, 4–5 mm; fronds 45–150 cm, clumped; stipes ca. 1⁄2 the frond length, stramineous to reddish brown, glabrous; blades ovate, (2–)3–4-pinnate, 20–70 cm wide; ultimate segments linear, chartaceous to membranous, glabrous, margins toothed, segments 3–4 mm wide; awns blunt, ca. 0.1 mm long; veins free, sterile ones ending before margins; sori continuous along margins, interrupted at segment apices; spores tan; indusia 0.8–1 mm wide, paraphyses absent; 2n⫽232 (Aus). Distribution. Terrestrial, montane cloud forests; 1600–1900 m. Mexico; Austr, NZ. Specimen Examined. Qro (Dı´az B. & Carranza 6560, IEB).

This species is known from a single collection from Quere´taro. It seems most likely that it represents an escape from cultivation; the species is known in the wild only from Australia and New Zealand. Among the free-veined, 2–3-pinnate species of Pteris, P. tremula resembles P. muricella and P. muricata in having the basal pinnae pinnate-pinnatifid beyond the basal pinnules but is distinct in the lamina being glabrous, membranous-chartaceous,

544

RADIOVITTARIA

the stipes lacking prickles, and the sori running nearly the entire segment length.

1 0 7 . R A DI O V I T T A R I A Radiovittaria (Benedict) E. H. Crane, Syst. Bot. 22: 514. 1997 [1998]. Vittaria Sm. subg. Radiovittaria Benedict, Bull. Torrey Bot. Club 38: 166. 1911. Type: Vittaria remota Fe´e [⬅ Radiovittaria remota (Fe´e) E. H. Crane].

Vittaria stipitata Kunze, Linnaea 9: 77. 1834. Type. Peru. Poeppig 176 (LZ, destroyed; isotypes, authentic specimens, and type fragments, B!, K, P, US!).

Rhizome scales dark brown to blackish, 1–2.5 ⫻ 0.1–0.3 mm, linear, 2(–3) cells wide at the base, uniseriate at tips beyond the base; petioles dark purple-brown, smooth, terete, indurate, not alate; blades mostly 20–60 ⫻ 0.2–0.5(–0.8) cm, linear; midribs not prominent, concolorous with respect to the blades; areoles in 1 row between the midrib and blade margins, veins not readily visible; sori sunken; paraphyses 2–4-celled.

Epiphytic; rhizomes suberect, radial, solenostelic, with strongly clathrate scales at the apex; fronds polystichous, clumped, 1–50(–110) cm long; stipes dark, flattened to terete; blades simple, entire, linear to narrowly elliptic, short-stipitate, mostly less than 50 ⫻ 0.2–1(–1.8) cm, costate; veins obscure, anastomosing and forming submarginal commissural veins parallel to the midrib, without included veinlets; sori of two submarginal continuous lines, one on each side of the midrib, surficial or in shallow to deep grooves; indusia absent; paraphyses reddish, with a funnelform (obconic) apical cell; spores bilateral, fusiform, hyaline to yellowish, smooth to sparsely papillate with fine spherical deposits; gametophytes with gemmae single; x⫽? Radiovittaria comprises about eight neotropical species, and is easily distinguished from Vittaria by the suberect rhizomes with radially arranged fronds, reddish paraphyses with funnelshaped apical cells, usually broader blades, and dark purplish brown lustrous stipes. It is apparently more closely related to Hecistopteris, differing from that in the entire linear blades. In addition, it is closely related to the Old World Haplopteris, a larger genus of about twenty species, according to Crane (1997). Until recently, Radiovittaria was considered part of a more inclusive genus Vittaria, which has been shown by Crane et al. (1995) and Crane (1997) to be polyphyletic, as treated in most taxonomic works. The vittarioid ferns (Vittariaceae, including also the genera Ananthacorus, Anetium, Hecistopteris, Polytaenium, Scoliosorus, and Vittaria, in Mexico) are monophyletic and believed to be most closely related to, and the sister group of Adiantum, which see. Vittariaceae plus Adiantum form the sister group to many other (but not all) genera in the Pteridaceae s.l., as that family is often circumscribed (Gastony & Rollo, 1995; Hasebe et al., 1995; Pryer et al., 1995; Cranfill, unpubl. data). References Benedict, R. C. 1911. The genera of the fern tribe Vittarieae. Their external morphology, venation and relationships. Bull. Torrey Bot. Club 38: 153–190; Crane, E. H. 1997 [1998]. A revised circumscription of the genera of the fern family Vittariaceae. Syst. Bot. 22: 509–517; Crane, E. H., D. R. Farrar & J. F. Wendel. 1995. Convergent simplification leads to a polyphyletic Vittaria. Amer. Fern J. 85: 283–305.

1. RADIOVITTARIA STIPITATA Radiovittaria stipitata (Kunze) E. H. Crane, Syst. Bot. 22: 515. 1997 [1998]. Figs. 260A–C.

Distribution. Epiphytic in montane rain forests; 300–1000 m. Mexico; Guat, Bel, Nic, CR, Pan; Gr Ant; Col, Ven, Fr Gui, Ec, Peru, s Braz, Bol. Specimens Examined. Chis (Breedlove 49966, CAS; Calzada 2823, XAL; Martı´nez S. 17601, 25908, MEXU; Martı´nez S. & Soto 18658, MEXU).

The sole Mexican species differs from other vittarioids by the characters mentioned under the genus.

1 0 8 . SA C C O L O M A Saccoloma Kaulf., Berlin. Jahrb. Pharm. Verbundenen Wiss. 21: 51. 1820. Type: Saccoloma elegans Kaulf. Orthiopteris Copel., Bernice P. Bishop Mus. Bull. 59: 14. 1929. Type: Orthiopteris ferulacea (T. Moore) Copel. [⬅ Davallia ferulacea T. Moore] ⬅ Saccoloma ferulaceum (T. Moore) R. M. Tryon & A. F. Tryon.

Terrestrial; rhizomes stout, compact, woody, erect, with dark, sclerotic scales and some hairs; fronds large, clumped, monomorphic; stipes glabrous, axes grooved, grooves confluent from one axis to the next; blades 1–4-pinnate, herbaceous to coriaceous, glabrous; veins free, simple to branched; sori submarginal, individual on veins, indusia forming outward-facing conical cups; paraphyses absent; spores tetrahedral-globose, tan, parallel ridges anastomosing at corners; x⫽ca. 63? (fide Smith, 1981). Saccoloma is a small pantropical genus of about 12 species, mostly of the southwestern Pacific, with only six species in the neotropics, occurring at low to middle elevations in wet forests. It is distinct in its generally large, coriaceous fronds, rhizome indument of dark, sclerotic scales, submarginal, discrete or sub-

SACCOLOMA INAEQUALE

discrete sori, and spores with parallel ridges. The relationships of Saccoloma are uncertain, but probably not with the dennstaedtioid ferns. Apparently, it is an isolated genus; see Wolf et al. (1999).

545

2. SACCOLOMA INAEQUALE Saccoloma inaequale (Kunze) Mett., Ann. Sci. Nat. Bot., se´r. 4, 15: 80. 1861. Figs. 261C–E.

Reference Tryon, R. M. 1962. Taxonomic fern notes. III. Contr. Gray Herb. 191: 91–107.

Key to the Mexican Species of Saccoloma 1. Fronds 1-pinnate, the pinnae entire to serrulate; sori contiguous along pinna margins. ....................................... 1. S. elegans. 1. Fronds bipinnate to quadripinnate-pinnatifid; sori one per segment lobe. ............................................................. 2. S. inaequale.

1. SACCOLOMA ELEGANS Saccoloma elegans Kaulf., Berlin. Jahrb. Pharm. Verbundenen Wiss. 21: 51. 1820. Figs. 261A, B. Davallia saccoloma Spreng., Syst. Veg., ed. 16, 4: 119. 1827 [based on Saccoloma elegans]. Type. Brazil. Sello s.n. (isotypes B, photo GH, not found, US!).

Rhizomes erect; rhizome scales castaneous, lustrous, linearlanceolate, 4–6 ⫻ ca. 1 mm; fronds 1–2 m long, clumped; stipes 1⁄3–1⁄2 the frond length, to 6 mm diam., stramineous to reddish brown, darker and muricate toward bases; blades oblong, 30–65 cm wide, 1-pinnate; pinnae linear-lanceolate, 18–36 ⫻ 1.8–4 cm, undivided, bases rounded to broadly cuneate, petiolulate (to 7 mm), entire to serrulate, apices acuminate to caudate; laminae papyraceous, glabrous; veins simple or 1-forked, prominent, ending in teeth; sori submarginal, contiguous the length of the pinnae, the inner indusia coalescing laterally.

Distribution. Terrestrial in wet lowland forests; 200 m. Mexico; Guat, Bel, Nic, CR, Pan; Gr Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol. Specimens examined. Oax (Herna´ndez G. 1555, 1643, 1770, 1910, 2058, 2540, NY—all from same area; Va´zquez T. et al. 1548, XAL). Ver (Vitale s.n., 5 Aug 1974, GH).

The large, once-pinnate fronds with contiguous, submarginal sori distinguish this species.

Davallia inaequalis Kunze, Linnaea 9: 87. 1834. Type. Peru. Loreto: Near Yurimaguas, Poeppig, Diar. 2113 (LZ, destroyed; isotype B!, photo GH!, frag. US!). Microlepia galeottii Fe´e, Me´m. Foug. 5: 327. 1852. Type. Mexico. Teotalcingo, Galeotti 6526 (P?, not found). Microlepia caudata Fe´e, Me´m. Foug. 5: 327. 1852. Type. Mexico. Galeotti 6527 (P!). For additional synonymy, see Proctor, Fl. L. Antill. 2: 118. 1977.

Rhizomes erect, to 20 mm diam.; rhizome scales bicolorous, central portion dark brown, lustrous, margins thin, erose, lighter brown, linear-lanceolate, 3–4 ⫻ 1–1.5 mm; fronds mostly 70–120 cm long (to 2 m), clumped; stipes nearly 1⁄2 the frond length, to 1 mm diam., green-yellow to black at bases, grooved, with minute bumps proximally; blades deltate, 40–90 cm wide, bipinnate to quadripinnate-pinnatifid; pinnae 4–7 pairs, alternate, inequilateral, petiolulate (stalks of proximal pinnae 0.5–1.5 cm); segments lanceolate-deltate to obliquely oblong, 1–2 cm long, sessile, distal segments adnate-decurrent, bases cuneate, margins serrate to lobed, slightly incurved, glossy dark green above, slightly lighter below, glabrous with minute papillae, coriaceous; veins free, nearly reaching the segment margins; vertical wing present on costae adaxially; sori submarginal, one per segment lobe, terminal on veins, 1 ⫻ 0.8 mm, usually impressed and surfaces embossed adaxially; indusia glabrous, crenulate, adnate to the blade along three sides forming a conical cup just back from the margin; 2n⫽126 +/– 4 (Chis).

Distribution. Wet montane forests on Atlantic slope; 150– 2000 m. Southern and eastern Mexico; Guat, Bel, Hond, Nic, CR, Pan; Cuba, PR; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol. Selected specimens examined. Chis (Breedlove 32208, NY; Calzada et al. 8809, NY; Ghiesbreght 355, NY). Oax (Herna´ndez G. 961, NY; Mickel 5696, 6606, NY). Pue (Ventura A. 396, NY). Ver (Ventura A. 3272, NY). Unverified, Doubtful, or Mistaken Reports. Gro (Lorea 2108, FCME; Vela´zquez 817, FCME, cited by Lorea-Herna´ndez & Vela´zquez M., 1998, but not verified).

546

SALVINIA

The dark green, coriaceous, highly dissected blades resembles those of Dennstaedtia bipinnata, but Saccoloma inaequale is distinct by its erect, scaly rhizomes and submarginal (vs. marginal) sori.

109. SALVINIA Salvinia Adans., Fam. Pl. 2: 15. 1763. Type: Uncertain, usually said to be Salvinia natans (L.) All. [⬅ Marsilea natans L.]. Aquatic, floating on water; roots absent, their function perhaps taken by highly dissected submersed leaves resembling a cluster of roots; rhizomes horizontal, slender, branched, hairy, the nodes close or distant from each other; leaves not circinate in bud, in groups of three at a node, two floating, green, flattened, sessile or short-stalked, the third leaf submerged, pendent, root-like and functioning as a root, brownish, highly branched, the branches covered with dark septate hairs; floating leaves ca. 5–25 mm long, round to oblong, often cordate at base, entire, flat or somewhat folded upward, with a distinct midrib and reticulate veins, without included veinlets, adaxial surfaces bearing numerous hairs from the blades or from multicellular papillae, in groups of 4, abaxial surfaces with uniseriate, septate, brownish hairs; sporocarps of 2 kinds (heterosporous), in chain-like or cyme-like clusters on the submerged leaves (both kinds on the same submerged leaf), a few, basal megasporocarps and more numerous, distal microsporocarps, both kinds globose, ca. 1.5–2 mm diam., the wall membranaceous (interpreted as a 2-layered indusium), each sporocarp containing numerous sporangia (collectively, the sorus) lacking annuli; sporangia ovoid or pearshaped, with a highly vacuolate layer in which the spores develop; megasporangia to ca. 40 per megasporocarp, on branched or unbranched stalks, each producing 32 spores but only one maturing, this tetrahedral-globose, with a minutely perforate surface; microsporangia 100+ per microsporocarp, on slender branched stalks, each containing 32 or 64 tetrahedral-globose, rugulose microspores, these shed as a unit and embedded in the vacuolate tissue; x⫽9. Salvinia is a pantropical genus of about 11 species with seven species in America. All species occur in stagnant, nutrient-rich, freshwater lakes and pools, and in slow moving rivers, at low to middle elevations. The genus was monographed by Herzog (1935), and the Salvinia auriculata complex was studied by de la Sota (1976), Mitchell (1972), and Mitchell and Thomas (1972). Some species form extensive mats that may clog waterways. Salvinia molesta D. S. Mitch. is an aggressive and noxious weed in some areas, as evidenced by its rampant growth on the artificial Lake Kariba in Zambia and Zimbabwe (Mitchell, 1969, 1972; Moran, 1992). This species, demonstrating hybrid vigor and abortive spores, is postulated by Mitchell (1972) to be a hybrid of horticultural origin in South America which has now spread to Africa, Ceylon, Indonesia, Australia, and the United States. Salvinia is related to Azolla, and the two genera (often treated in separate families) have been shown recently to be most closely related to the third (and only other) family of heterosporous

ferns, the Marsileaceae (Pryer et al., 1995; Rothwell & Stocky, 1994). Salvinia is distinct by its floating habit, round to oblong vegetative leaves with compound hairs, and root-like leaves with sporocarps. Because of the habitat in which the plants are found, both the vegetative and the reproductive structures are highly modified and do not resemble those of other ferns. Fertile material is rarely seen. Fossils of the genus are known from the Upper Cretaceous. References de la Sota, E. R. 1976. Sinopsis de las especies argentinas del ge´nero Salvinia. Bol. Soc. Argent. Bot. 17: 47–50; Forno, I. W. 1983. Native distribution of the Salvinia auriculata complex and keys to species identification. Aquatic Bot. 17: 71–83; Herzog, R. 1935. Ein Beitrag zur Systematik der Gattung Salvinia. Hedwigia 74: 257–284; Mitchell, D. S. 1969. The ecology of vascular hydrophytes on Lake Kariba. Hydrobiologia 34: 448–464; Mitchell, D. S. 1972. The Kariba weed: Salvinia molesta. Brit. Fern Gaz. 10: 251–252; Mitchell, D. S. & P. A. Thomas. 1972. Ecology of waterweeds in the Neotropics, an ecological survey of the aquatic weeds Eichhornia crassipes and Salvinia spp. and their natural enemies in the Neotropics. UNESCO Techn. Pap. Hydrol. 12: 1–50; Moran, R. C. 1992. The story of the molesting Salvinia. Fiddlehead Forum 19: 26–28; Pe´rez-Garcı´a, B., A. Mendoza & R. Riba. 2002. Pteridofitas. Familia Salviniaceae. Flora de Me´xico 6(6): 9–17; Rothwell, G. W. & R. A. Stockey. 1994. The role of Hydropteris pinnata gen. et sp. nov. in reconstructing the cladistics of heterosporous ferns. Amer. J. Bot. 81: 479–492; Weatherby, C. A. 1937. A further note on Salvinia. Amer. Fern J. 27: 98–102.

Key to the Mexican Species of Salvinia 1. Hairs at the tips of the papillae on the adaxial leaf surfaces joined together into a darkened knot at their tips; sporocarps stalked; rhizome stele U-shaped; floating leaves 1.5–2 ⫻ 1.8–2.5 cm, notched at both ends. ....... 1. S. auriculata. 1. Hairs at the tips of the papillae on the adaxial leaf surfaces free at their tips, spreading; sporocarps sessile; rhizome stele circular; floating leaves 0.6–1.5 ⫻ 0.6–2 cm, round (to rarely notched) at apex. ........................... 2. S. minima.

1. SALVINIA AURICULATA Salvinia auriculata Aubl., Hist. Pl. Guiane 2: 969, t. 367. 1775. Type. French Guiana. “In Territorio Caux,” Aublet s.n. (P?). Figs. 262E–H, J, K. Salvinia rotundifolia Willd., Sp. Pl., ed. 4, 5(1): 537. 1810. Type. Brazil. Hoffmannsegg s.n. (B-Willd. 20250!). See C. V. Morton, Contr. U.S. Natl. Herb. 38: 5. 1967, for discussion of the type and previous misinterpretation.

Rhizomes with U-shaped stele in cross section; floating leaves orbicular to orbicular-cordate, wider than long (in the sun), 1.5–2 ⫻ 1.8–2.5 cm, notched on both ends, adaxial side with numerous elongate conical papillae over entire surface in rows between main veins, papillae with 4 (rarely 2) hairs, joined together at their tips into a darkened knot; sporocarps pedunculate on branched axes, oldest on longest peduncles, with all sporocarps hanging at about same level, globose, not markedly apiculate; 2n⫽54 (Trin).

SCHAFFNERIA

547

Distribution. Floating in still or stagnant fresh water or margins of slowly flowing streams; 0–50 m. Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Gr Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol, Chile, Parag, Arg, Uru.

Distribution. Floating in marshes, lakes, ponds and quiet backwaters of streams and rivers; 0–650 m. USA (Ga, Fla); Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Berm, Gr & L Ant; Col, Ven, Fr Gui, Ec, Peru, Braz, Bol, Parag, n Arg, Uru.

Selected Specimens Examined. Camp (Ramı´rez R. 91, UAMIZ). Chis (Breedlove 34932, DS, LL, MICH, MO, NY, 56079, CAS, ENCB). Gro (Boege 638, CAS). Oax (Riba 1701, UAMIZ). Tab (Cowan 2807, CSAT, NY; Magan˜a & Priego 1269, ENCB, XAL). Tam (Palmer 312, CAS, K, NY, US; Rzedowski 24575, DS, ENCB, LL). Ver (Rico-Gray 39, XAL).

Selected Specimens Examined. Camp (Sanders et al. 9743, UC, UCR). Chis (Breedlove 56065, CAS, NY; Morley 714, MO, UC, US). QR (Lot & Novelo 787, MEXU). Tab (Cowan 2805, CSAT, NY; Magan˜a et al. 335, CAS, K, MEXU). Tam (LeSueur 8, TEX). Ver (Faden et al. 76/154, UC; Gutie´rrez 1594, XAL). Yuc (Gaumer 1007, F, GH, NY, US).

Unverified, Doubtful, or Mistaken Reports. Jal (Pocovi 15485, GUADA, cited by Mickel, 1992, but not verified; see comments under S. minima). Mich (reported in the range by Mickel & Beitel, 1988, and Mickel, 1992, but not verified).

Pocovi s.n. (UAMIZ), from Jalisco, seems to be S. minima on the basis of leaf size, but the hairs at the tips of the papillae are too poorly developed to assess the critical character. A specimen from Tabasco, Gliessman et al. 1694 (CSAT, MO p.p., NY, TEX) has the papillae very reduced or obsolete; it has the blade size of S. minima, but the adaxial papillae are similar to S. auriculata. Most New World specimens identified as S. natans (L.) All., a Eurasian species, are actually S. minima. Salvinia natans resembles S. minima in its spreading hairs at the tips of short papillae and round sporocarps.

The Salvinia auriculata complex, with the hairs joined at their tips, consists of at least four species, three of which have not been reported north of South America (de la Sota, 1976; Forno, 1983; Mitchell, 1972; Mitchell & Thomas, 1972). The species in the complex other than S. auriculata itself have the main axis of the submerged leaf straight, rather than curved, and sporocarps are apiculate on long, many-branched fertile axes.

2. SALVINIA MINIMA Salvinia minima Baker, J. Bot. 24: 98. 1886. Type. Brazil. Santa Catarina: Mu¨ller 479 (K!). Figs. 262A–D. Salvinia rotundifolia auct., non Willd., a misapplied name, discussed by Morton (see citation under Salvinia rotundifolia Willd., a synonym of S. auriculata).

Rhizomes with circular stele in cross section; floating leaves orbicular to slightly oblong, wider than long, 0.6–1.5 ⫻ 0.6–2 cm, cordate at bases, obtuse (to rarely notched) at apices, adaxial surfaces with numerous conical papillae (2–3 mm long) over entire surface in rows between main veins, with 4 hairs spreading and free at their tips; sporocarps sessile on unbranched, pendent main axes of fertile leaves, lemon-shaped, somewhat apiculate.

1 1 0 . SC H A F F N E R I A Schaffneria Fe´e, Me´m. Foug. 7: 56, t. 17, f. 1. 1857. Type: Schaffneria nigripes Fe´e. Epipetric, or less commonly terrestrial; rhizomes short, compact, ascending, with dark, clathrate scales; fronds small, clumped, monomorphic; stipes black, lustrous, with minute white to tan, appressed hairs; blades simple, irregularly orbicular, flabellate, glabrous, fleshy, margins entire; veins obscure, netted, without included veinlets; sori linear, sometimes paired, facing each other, indusiate, lacking paraphyses; spores bilateral, with cristate perispores; x⫽?. Schaffneria is a monotypic genus of Middle America and Cuba, named for William Schaffner, not to be confused with J. G. Schaffner, who also collected in Mexico in the 1800s. Schaffneria is closest to Sinephropteris of China (Mickel, 1976), the type of which had been placed in Schaffneria by TardieuBlot (1957). Schaffneria is distinguished by its small, fleshy, fla-

548

SCHIZAEA

bellate blades, black stipes, and paired sori face to face on adjacent veins. Although molecular data are inconclusive, it appears almost a certainty that Schaffneria is nested within Asplenium s.l., where it was treated as a synonym by Kramer and Viane (in Kubitzki, 1990: 56). Despite its distinctive and striking appearance, there is no single morphological character in Schaffneria that cannot be found in other species of Asplenium. Thus, the generic separation of Schaffneria is strongly in doubt. References Mickel, J. T. 1976. Sinephropteris, a new genus of scolopendrioid ferns. Brittonia 28: 326–328; Riba, R., B. Pe´rez-Garcı´a & M. Pe´rez-Garcı´a. 1992. Schaffneria nigripes (Aspleniaceae): morfoge´nesis del gametofito y anatomı´a y morfologı´a del esporofito. Bol. Soc. Bot. Me´xico 52: 105–113; Tardieu-Blot, M.-L. 1957. Sur un Antigramma de Madagascar et sur la re´partition ge´ographique des genres Antigramma et Schaffneria. Naturaliste Malgache 9: 29–32.

1. SCHAFFNERIA NIGRIPES Schaffneria nigripes Fe´e, Me´m. Foug. 7: 56, t. 17, f. 1. 1857. Figs. 263A, B. Asplenium nigripes (Fe´e) Hook., Hooker’s J. Bot. Kew Gard. Misc. 9: 268, t. 9. 1857. Scolopendrium nigripes (Fe´e) Hook., Sp. Fil. 4: 3. 1862. Antigramma nigripes (Fe´e) J. Sm., Hist. Fil. 331. 1875. Type. Cited only as collected by Schaffner in Mexico (Me´m. Foug. 7: 56. 1857). In Me´m. Foug. 9: 18. 1857, Fe´e cited only one collection, W. Schaffner 47, from Orizaba. This collection at P would logically be the type.

Rhizomes erect; rhizome scales black, clathrate, 2–2.5 ⫻ 0.3– 0.5 mm, entire; stipes black, lustrous, color ending abruptly at blade base, 2–8 cm long, ca. 3⁄5 the frond length, with occasional linear, clathrate scales and minute, 1- to few-celled hairs; blades fleshy, very dark green adaxially, undivided, suborbicular to subrhombic, 2.5–5.5 cm wide, apices acute to rounded; sori mostly simple, a few in facing pairs, indusia 2–18 ⫻ 0.5–0.8 mm, entire.

UC, US). Qro (Ferna´ndez N. 3466, ENCB, IEB; Rzedowski 46617, IEB). SLP (Pringle 3366, F, GH, MEXU, NY, UC, US, 4090, MICH, UC). Tam (Harrell 186, US). Ver (Copeland herb. 68, BM, GH, MICH, UC, US; Mu¨ller 693, NY).

1 1 1 . SC H I Z A E A Schizaea Sm., Me´m. Acad. Roy. Sci. (Turin) 5: 419. 1793, nom. cons. Type (implicit in Smith’s publication; see Webbia 12: 6. 1956): Schizaea dichotoma (L.) Sm. [⬅ Acrostichum dichotomum L.]. Lophidium Rich., Actes Soc. Hist. Nat. Paris 1: 114. 1792, nom. rejic. Type: Lophidium latifolium Rich. ⫽ Schizaea elegans (Vahl) Sw.

Rhizomes short-creeping or ascending, subterranean, hairy; fronds erect, undivided and grass-like, or the veins dichotomously forked and forming a fan-like divided or undivided blade; sporangia on terminal pinnate fertile pinnules of fertile blades, either at the apices of undivided blades or on terminal flag-like projections on fan-shaped blades; sporangia in two rows on each fertile segment, often mixed with hairs, sporangia erect, oblong with terminal annuli, exindusiate; spores bilateral, foveolate; x⫽77, 94, 96, 103. Schizaea is a genus of about 20 species, largely of the Old World tropics. It is distinct by its erect apically annular sporangia on flag-like projections of the blade and grass-like to flabellate lamina. It is related most closely to Actinostachys, which is sometimes included in Schizaea, and more distantly to Anemia and Lygodium (e.g., Hasebe et al., 1995). They agree in having terminal annuli. Key to the Mexican Species of Schizaea 1. Sterile blade segments expanded, 3–28 mm wide; fertile pinnules 5–12 mm long; fronds monomorphic, the fertile ones about the same length as the sterile. ............. 1. S. elegans. 1. Sterile blade segments linear, 1 mm wide; fertile pinnules (1–)1.5–4 cm long; fronds dimorphic, the fertile ones far exceeding (2–3 times) the sterile in length. ......................................................................................... 2. S. poeppigiana.

1. SCHIZAEA ELEGANS Schizaea elegans (Vahl) Sw., J. Bot (Schrader) 1800(2): 103. 1801. Figs. 264A, B. Acrostichum elegans Vahl, Symb. Bot. 2: 104, t. 50. 1791. Lophidium elegans (Vahl) C. Presl, Gefa¨ssbu¨ndel Farrn 337. 1847. Type. Trinidad. von Rohr s.n. (C!).

Distribution. On moist, shaded limestone rocks, in wet hardwood forests on Atlantic slopes; 800–1400 m. Mexico; Guat; Cuba. Also reported from Costa Rica by Go´mez (Revista Biol. Trop. 21: 91–101. 1973, but not verified). Selected Specimens Examined. Chis (Breedlove 29133, DS, F, MEXU, NY; Ghiesbreght 302, K, NY). Oax (Mickel 7333, ENCB, NY,

Rhizomes ascending, underground, 1.5–2 mm diam., hairs tan, roots wiry (ca. 0.5 mm diam.); fronds monomorphic, erect, 25– 42 cm long; stipes 3⁄4 the frond length, stramineous, brown at bases, glabrous; blades flabellate, dichotomously divided 3–6 times, the broadest segments 1–2.8 cm wide, glabrous; veins free; fertile fronds with segments terminated with flag-like fertile pin-

SCOLIOSORUS

549

nules 5–12 mm long folded lengthwise in half; sporangia along the veins, intermixed with golden hairs.

Distribution. On sand or rocks in dense wet forests on Atlantic lowlands; 100–950 m. Mexico; Guat, Bel, Hond, Nic, CR, Pan; Jam, Hisp; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol, Parag. Selected specimens examined. Chis (Breedlove 57353, CAS; Martı´nez S. 7690, NY, 13951, MEXU; Matuda 3662, CAS, NY). Oax (Galeotti 6536 B, BR, LE; Hallberg 1330, NY; Herna´ndez G. 2565, NY; Martı´nez-C. 860, MICH, US). Tab (Zamudio & Magan˜a 106, MEXU). Ver (Orcutt 2913, UC).

This species is quite variable in its dissection, especially in South America, where it has been treated as several species by some authors. The forking fronds and flag-like fertile pinnules are distinctive for this species.

2. SCHIZAEA POEPPIGIANA Schizaea poeppigiana J. W. Sturm in Martius, Fl. Bras. 1(2): 181. 1859. Lectotype (chosen by Proctor, 1985: 80). Peru. “ad Ventanilla de Cassapillo,” Poeppig s.n. (BR?, not found). Figs. 264C–E. Similar to S. elegans except: sterile segments linear, only 1 mm wide (vs. 1–2.8 mm), the fertile fronds dimorphic, 2–3 times longer than the sterile (vs. about equal in length), and the fertile pinnae (1–)1.5–4 cm long (vs. 0.5–1.2 cm).

Distribution. Terrestrial and epipetric in lowland subperennial forests, sometimes flooded; 180 m. Mexico; Bel, Hond, Nic, CR, Pan; Bah, Cuba, Hisp, PR; Col, Ven, Guy, Ec, Peru, Braz, Bol. Specimens examined. Chis (Martı´nez S. 13953, 14895, 16325, 17877, MEXU).

1 1 2 . SC O L I O SO R US Scoliosorus T. Moore, Index Fil. xxix. 1857. Type: Scoliosorus ensiformis (Hook.) T. Moore [⬅ Antrophyum ensiforme Hook.] Epiphytic; rhizomes short-creeping, solenostelic, dorsiventral, with clathrate scales; roots densely tomentose with abundant root hairs; fronds mostly ca. 15–30(–70) cm long; stipes short to absent; blades simple, linear to elliptic or oblanceolate [to orbicularacuminate], entire, glabrous; veins anastomosing and forming 2– several series of areoles, sometimes with free marginal branches; epidermis with linear idioblasts; sori linear, in 2 or 4(–6) lines parallel to the midvein or in shorter, irregular, anastomosing lines, these or the areoles thus formed oblique to the midrib, either on the surface or more commonly immersed in the blades; indusia absent; paraphyses present and with a spherical apical cell; spores bilateral, reniform, yellow-hyaline or rarely green; gametophytes with paired gemmae; x⫽? As redefined by Crane (1997), Scoliosorus comprises three species, a single one in the neotropics and two or a few more in Africa and islands of the southwestern Indian Ocean. Scoliosorus is most closely related to Ananthacorus, and differs from that chiefly by the sporangia in reticulating soral lines (vs. two submarginal lines), the spherical apical cell of the paraphyses (vs. clavate, narrow apical cell), and, in our species at least, by the paired gemmae on the gametophytes (vs. gametophytes lacking gemmae). Scoliosorus has often been included in the genus Polytaenium or Antrophyum s.l. References Benedict, R. C. 1907. The genus Antrophyum I. Synopsis of the subgenera, and the American species. Bull. Torrey Bot. Club 34: 445–458; Benedict, R. C. 1911. The genera of the fern tribe Vittarieae. Their external morphology, venation, and relationships. Bull. Torrey Bot. Club 38: 153–190; Crane, E. H. 1997 [1998].

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SELAGINELLA

A revised circumscription of the genera of the fern family Vittariaceae. Syst. Bot. 22: 509–517; Tryon, R. M. 1964a. Taxonomic fern notes IV. Some American vittarioid ferns. Rhodora 66: 110–117.

1. SCOLIOSORUS ENSIFORMIS Scoliosorus ensiformis (Hook.) T. Moore, Index Fil. xxix. 1857. Figs. 265A–E. Antrophyum ensiforme Hook. in Bentham, Pl. Hartw. 73. 1841. Polytaenium ensiforme (Hook.) Benedict, Bull. Torrey Bot. Club 38: 169. 1911. Type. Mexico. Cumbre de Totontepeque, Hartweg [522] (K!). Antrophyum falcatum M. Martens & Galeotti, Me´m. Foug. Mexique 49. 1842, hom. illeg., non Blume, 1828. Antrophyum carnosum Liebm., Mexic. Bregn. 161 (reprint 9). 1849. Antrophyum galeottii Fe´e, Me´m. Foug. 4: 51, pl. 5, f. 4. 1852. Type. Mexico. Oaxaca: Llano Verde, Galeotti 6385 (BR!, P! photo ex BR, BM! photo ex BR).

Rhizomes short-creeping, 2–4 mm diam.; rhizome scales clathrate, linear-lanceolate, long-attenuate at apices, 10–12 ⫻ 1.5–2 mm; fronds 12–45 cm long, clumped; stipes absent; blades linearoblanceolate, entire, 10–20 mm wide at the widest point, (2–)5–8 mm wide at very base, apices acuminate, glabrous, fleshy; veins obscure, netted, 2–3 rows of oblique areoles between midveins and blade margins, veins near margins forming areoles; sori in grooves, rarely surficial, 1–10 mm long, somewhat following vein pattern, forming a loose and irregularly connecting pattern on each side of the midvein; paraphyses clavate, abundant to rarely scattered, reddish, 0.3–0.5 mm long; spores bilateral, whitish to cream-colored.

Distribution. Epiphytic or epipetric in cloud forests on both Atlantic and Pacific slopes, montane cloud forests, damp oak forests, evergreen cloud forests, semi-evergreen montane forests, fir-oak-Podocarpus forests; 550–2800 m. Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan. Selected Specimens Examined. Chis (Breedlove 57770, CAS, ENCB; Martı´nez S. 17619, BM, MEXU). Gro (Hinton 11174, K, US; Rzedowski & McVaugh 188, ENCB, MEXU). Oax (Mickel 3817, ENCB, MEXU, NY, 7252, NY, UC). Pue (Vera Santos 3357, US). SLP (Schaffner 71, NY). Ver (Calzada 425, CAS, ENCB, MEXU; Ventura A. 8212, ENCB, MEXU). Unverified, Doubtful, or Mistaken Reports. Me´x (reported by Mickel & Beitel, 1988, but not verified; see comments below).

This species is mostly likely to be confused with Loxogramme mexicana, which differs in having fewer sori closer to the midrib and more nearly parallel to it. From species of Polytaenium it differs by the numerous paraphyses with a spherical tip and the bilateral spores. It is relatively common in Chiapas, Oaxaca, and Veracruz, in suitable habitats, but rare or uncommon elsewhere in Mexico. Schaffner 71 (NY), “Valley of Mexico,” 1875, is no doubt the basis for citation of this species from Edo. Me´xico by Mickel and Beitel (1988), but the specimen is most likely from San Luis Potosı´ (Rzedowski, 1959), where Schaffner is known to have collected extensively during this time. Another specimen with the same number at NY, but without year, is labelled as from San Luis Potosı´, and a third collection, dating from 1876, “Santa Marı´a de Rey,” but without state, is probably also from San Luis Potosı´ (cuenca del rı´o de Santa Marı´a; Rzedowski, 1959: 102).

1 1 3. SE L A GI N E L L A by John T. Mickel, Alan R. Smith, and Iva´ n A. Valdespino Q.

Selaginella P. Beauv., Mag. Encycl. 5: 478. 1804; and Prod. Aethe´ogam. 101. 1805., nom. cons. Type: Selaginella spinosa P. Beauv., nom. nov. for Lycopodium selaginoides L. [⬅ Selaginella selaginoides (L.) Link]. Stachygynandrum P. Beauv. ex Mirb., in Lamarck & Mirbel, Hist. Nat. Veg. 3: 477. 1802. Selaginella P. Beauv. subg. Stachygynandrum (P. Beauv.) Baker, J. Bot. 21: 3. 1883. Type: Stachygynandrum flabellatum (L.) P. Beauv. [⬅ Lycopodium flabellatum L.] ⬅ Selaginella flabellata (L.) Spring. Selaginella P. Beauv. subg. Heterostachys Baker, J. Bot. 21: 4. 1883. Lectotype (chosen by Jermy, 1986): Selaginella heterostachys Baker. Selaginella P. Beauv. subg. Tetragonostachys Jermy, Fern Gaz. 13: 118. 1986. Type: Selaginella rupestris (L.) Spring [⬅ Lycopodium rupestre L.]. Bryodesma Soja´k, Preslia 64: 154. 1992. Type: Bryodesma rupestris (L.) Soja´k. [⬅ Lycopodium rupestre L.] ⬅ Selaginella rupestris (L.) Spring.

Terrestrial or epipetric, rarely epiphytic, prostrate and creeping to somewhat ascending or fully erect, slightly to much branched; stems with abundant very small leaves, isophyllous and tightly imbricately arranged spirally in several rows, or more commonly anisophyllous in four rows, two lateral rows of larger spreading leaves and two rows of appressed and ascending median leaves on upper side of stems, also with axillary leaves in axils of branches; rhizophores (roots) either basal or from nodes (branchings), dichotomously branched on touching the ground; leaves orbicular to linear, each with a single vein and a minute ligule, sometimes ending in hair-like tips, entire to ciliatedenticulate, glabrous or rarely hispid on upper leaf surfaces, generally green above, silver beneath; sporangia sessile, generally in 4-sided strobili terminal on branches, the sporophylls 4-ranked and slightly modified from the vegetative leaves, usually monomorphic, rarely dimorphic; spores of two kinds (plants heterosporous); microspores and megaspores in separate sporangia but within the same strobilus; microsporangia each with hundreds of microspores (18–31(–63) ␮m diam.); megasporangia each with four large megaspores (150–1500 μm diam.); spores tetrahedralglobose, with an equatorial flange, in some species the microspores shed as tetrads.

SELAGINELLA

Selaginella is a genus of 600–700 species, mostly of tropical regions. Two major groups are recognized: the isophyllous species, which are mostly of dry regions of Mexico and western United States; and the anisophyllous species, which represent by far the largest portion of the genus, these largely of wet tropical forests. Eighty species of Selaginella occur in Mexico. In the key and descriptions below, the following conventions and terminology are used. Exceptions to these conditions are mentioned when significant. In all species, the root-like structures (rhizophores) are straight aboveground, but fork dichotomously in the soil. The green expanded portion of the leaves is referred to as the lamina, excluding the arista or apiculum at the tip. The midveins of leaves and sporophylls are sometimes prominent or “keeled.” Megaspore and microspore colors were determined from air-dried spores of recent herbarium specimens mostly using Ridgway’s (1912) color standards. The isophyllous species have their leaves basically all alike (linear to lanceolate), thick, often with marginal cilia and pale aristae. Some isophyllous species have the leaves close to the substrate surface morphologically different from those on the side away from the substrate. In this case, the leaves on the side of the axis away from the surface are called upperside leaves, and those on the side toward the surface are called underside leaves. In this case they are called “dimorphic,” as opposed to anisophyllous. Similarly, some species have slightly or strongly different leaves on underground (rhizomatous) and aerial stems. In this case, the leaves are called rhizomatous stem leaves and aerial stem leaves respectively. In the anisophyllous species both the lateral and median leaves are oblique at base, somewhat distant in the lower parts of the stems, and imbricate in the distal regions. The lateral leaves are often held at right angles to the stem in the lower parts of the plant, but are more ascending distally. The acroscopic bases are usually more exaggerated and overlapping the underside than the basiscopic bases on the stems. The median leaves are strongly ascending, and the outer side of their bases is more exaggerated (sometimes auriculate). Axillary leaves are located at the points of stem branching and are best seen from the underside. They closely resemble the lateral leaves but are more symmetric. The leaves often possess distinct, elongate, often white, thickwalled cells, called idioblasts or “false veins,” on one or both surfaces. Leaf margins are often differentiated (green, hyaline, or white), and are usually short- or long-ciliate, or sometimes denticulate. In one group of anisophyllous species stem swellings (“articulations”) occur at points of branching. Strobili are most commonly quadrangular, though some are dorsiventral with dorsal rows of sporophylls green and spreading, the ventral ones pale green to hyaline and ascending. Megasporangia and microsporangia vary in their distribution within the strobili; the anisophyllous species often have few to only one megasporangia per strobilus, and this is basal and ventral. Selaginella, the single genus of the family of the spikemosses (Selaginellaceae), is clearly isolated and monophyletic, in all molecular analyses so far attempted (Korall & Kenrick, 1999; Korall

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et al., 1999; Therrien et al., 1999). It is the sister group of the quillworts, Isoe¨taceae, in all “global” analyses performed on pteridophytes (e.g., Nickrent et al., 2000; Pryer et al., 2001). Selaginella and Isoe¨tes, both heterosporous, are in turn the sister group of the homosporous lycopods (Lycopodiaceae). These three families are the basalmost extant group of tracheophytes (vascular plants), and are sister to all remaining vascular plants. Relationships within Selaginella have recently been addressed in preliminary work using the chloroplast rbcL gene (Korall & Kenrick, 1999; Korall et al., 1999). Evidence suggests that the type of Selaginella and a few close allies (subg. Selaginella; this group not represented in Mexico) form the basal element in the family. The isophyllous (homophyllous) species (subg. Tetragonostachys, in the nomenclature of Jermy, 1986; the genus Bryodesma, according to Soja´k, 1992) form a monophyletic group. The primary center for the diversity of subg. Tetragonostachys is in Mexico and the southwestern United States. The following 21 species are known from Mexico: S. arizonica, S. arsenei, S. asprella, S. basipilosa, S. bigelovii, S. carnerosana, S. cinerascens, S. eremophila, S. extensa, S. landii, S. macrathera, S. mutica, S. parishii, S. peruviana, S. rupincola, S. sartorii, S. sellowii, S. steyermarkii, S. underwoodii, S. viridissima, and S. wrightii. From the molecular data published so far, the sister group to subg. Tetragonostachys appears to be a group of so-called “articulate” species (series Articulatae, an unpublished name; for circumscription, see Somers, 1982, and Somers & Moran in Davidse et al., 1995). These species are anisophyllous (two different kinds of leaves) and also well represented in Mexico, with seven species: S. eurynota, S. marginata, S. schizobasis, S. sertata, S. silvestris, S. stellata, and S. tarda. The group is characterized by having articulations or swollen joints just below each bifurcation of the stems. In the living state, the articulations appear as prominent reddish areas; in dried specimens, they often appear blackish, with a distinct abscission line. Other characteristics of the articulate group are that (1) each strobilus has a single large megasporangium at the base; (2) rhizophores are borne on the dorsal side of the stems; (3) megaspores are exceptionally large and strongly cristate-reticulate; and (4) microspores are yellowish to brownish and echinate. The exact relationship of the articulate species to the homophyllous species (subg. Tetragonostachys) and to the species in subg. Stachygynandrum (see below) are still uncertain. All 52 remaining species of Selaginella in Mexico fall within subg. Stachygyandrum (sensu Jermy (1986), which is the most species-rich clade in the genus. Subgenus Stachygynandrum comprises anisophyllous (heterophyllous) species, characterized by having ascending, relatively short median leaves and spreading, usually longer lateral leaves. Although the natural groups within Stachygynandrum are not yet clear, the group called Heterostachys (recognized at subgeneric rank by Valdespino, 1995) is delimited by having dimorphic sporophylls. In Mexico, there are 14 species in this group: S. bernoullii, S. finitima, S. flagellata, S. lineolata, S. minima, S. mixteca, S. moritziana, S. popayanensis, S. porphyrospora, S. prolifera, S. simplex, S. stenophylla, S. tenella, and S. tuberosa. In addition to having the dorsal and ventral sporophylls oriented differently, species of Heterostachys tend to be prostrate,

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have flagelliform branch tips or sobols (filiform, leafy, horizontal stoloniferous stems from near the bases of main stems) and rather thin leaves. Another probably natural group includes those species that have a rosette habit with fascicled stems, and strongly imbricate and relatively thick-textured leaves. This group includes S. convoluta, S. gypsophila, S. lepidophylla (resurrection plant), S. nothohybrida, S. novoleonensis, S. pilifera, and S. ribae. A related group, also with imbricate leaves but not as strongly rosette-forming, includes S. acutifolia, S. harrisii, S. pallescens, S. polyptera, and S. pulcherrima. References Abundiz-Bonilla, L. A. M. & D. Tejero-Dı´ez. 1990. El ge´nero Selaginella P.Beauv. (Selaginellaceae, Lycophyta) en el Oeste del Estado de Me´xico. Acta Boyt. Mex. 11: 23–97. Alston, A. H. G. 1951. A revision of the West Indian species of Selaginella. Bull. Brit. Mus. (Nat. Hist.), Bot. 1: 25–50; Alston, A. H. G. 1955. The heterophyllous Selaginellae of continental North America. Bull. Brit. Mus. (Nat. Hist.), Bot. 1: 222–274; Alston, A. H. G., A. C. Jermy, & J. M. Rankin. 1981. The genus Selaginella in tropical South America. Bull. Brit. Mus. (Nat. Hist.), Bot. 9: 233–330; Buck, W. R. 1977. A new species of Selaginella in the S. apoda complex. Canad. J. Bot. 55: 366–371; Dı´az, H. & M. Palacios. 1992. Listado preliminary de especies pterido´fitas de los estados de Guanajuato, Michoaca´n y Quere´taro. Flora del Bajı´o y Regiones Adyacentes, Fasciculo Complementario 3: 1–58. Instituto de Ecologı´a A.C.; Gregory, D. & R. Riba. 1979. Selaginellaceae. Flora de Veracruz 6: 1–35; Hellwig, R. L. 1969. Spores of the heterophyllous Selaginellae of Mexico and Central America. Ann. Missouri Bot. Gard. 56: 444–464; Jermy, A. C. 1986. Subgeneric names in Selaginella. Fern Gaz. 13: 117–118; Korall, P. & P. Kenrick. 1999. Phylogeny of Selaginellaceae

based on rbcL gene sequences. XVI International Botanical Congress, Abstracts. p. 335 (Poster 257); Korall, P. & P. Kenrick. 2002. Phylogenetic relationships in Selaginellaceae based on rbcL sequences. Amer. J. Bot. 89: 506–517; Korall, P., P. Kenrick & J. P. Therrien. 1999. Phylogeny of Selaginellaceae: evaluation of generic/subgeneric relationships based on rbcL gene sequences. Int. J. Pl. Sci. 160: 585–594; Mickel, J. T. & I. A. Valdespino. 1992. Five new species of pteridophytes from Oaxaca, Mexico. Brittonia 44: 312–321; Nickrent, D. L., C. L. Parkinson, J. D. Palmer & R. J. Duff. 2000. Multigene phylogeny of land plants with special reference to bryophytes and the earliest land plants. Molec. Biol. Evol. 17: 1885–1895; Riba, R., L. Pacheco, A. Valde´s & Y. Sandoval. 1996. Pteridoflora del estado de Morelos, Me´xico. Lista de familias, ge´neros y especies. Acta Bot. Mex. 37: 45–65; Ridgeway, R. 1912. Color standards and color nomenclature. Published by the author, Washington, D.C.; Soja`k, J. 1992. Generische Problematik de Selaginellaceae. Preslia 64: 151–158; Somers, P. 1982. A unique type of microsporangium in Selaginella series Articulatae. Amer. Fern J. 72: 88–92; Therrien, J. P., C. H. Haufler & P. Korall. 1999. Phylogeny and biogeography of Selaginella subg. Tetragonostachys. XVI International Botanical Congress, Abstracts. p. 91; Tryon, A. 1949. Spores of the genus Selaginella in North America, north of Mexico. Ann. Missouri Bot. Gard. 36: 413–436; Tryon, R. M. 1955. Selaginella rupestris and its allies. Ann. Missouri Bot. Gard. 42: 1– 99; Valdespino, I. A. 1993. Notes on neotropical Selaginella (Selaginellaceae), including new species from Panama. Brittonia 45: 315–327; Valdespino, I. A. 1994. A new Selaginella (Selaginellaceae) from Quere´taro and Veracruz, Mexico. Amer. Fern J. 84: 99–101; Valdespino, I. 1995. A monographic revision of Selaginella P. Beauv. subgenus Heterostachys Baker in Central and South America. Ph.D. thesis, The City University of New York.; Wagner, W. H. Jr., J. M. Beitel & F. S. Wagner. 1982. Complex venation patterns in the leaves of Selaginella: megaphyll-like leaves in lycophytes. Science 218: 793–794; Weatherby, C. A. 1944. The group of Selaginella oregana in North America. J. Arnold Arbor. 25: 407–419.

Key to the Mexican Species of Selaginella 1. Plants isophyllous; leaves linear to lanceolate, spirally arranged on the stem in several indistinct rows. 2. Aerial stems prostrate, dorsiventral; leaves strongly dimorphic to subdimorphic; rhizophores throughout length of stems. 3. Underside and upperside leaves abruptly adnate; leaves slightly dimorphic. ............................................................................80. S. wrightii. 3. Underside leaves decurrent, upperside leaves abruptly adnate; leaves strongly to moderately dimorphic. 4. Base of underside leaves pubescent. ....................................................................................................................................... 8. S. basipilosa. 4. Base of underside leaves glabrous or glabrescent. 5. Apices of upperside leaves broadly acute to muticous or short-aristate; sporophylls linear to lanceolate with acuminate to short-aristates apices, or if ovate-deltate, at least some leaves with long, twisted apices. 6. Apices of underside leaves mostly short-aristate. ...................................................................................................... 33. S. landii. 6. Apices of underside leaves acute to obtuse. ........................................................................................................... 51. S. parishii. 5. Apices of upperside leaves aristate, mucronate, or acute. 7. Leaves with a twisted arista (at branch tips or buds) or acute to mucronate (by breaking off); upperside leaves lanceolate; plants forming compact mats. 8. Leaf aristae caducous, leaf apices becoming acute to mucronate; sporophylls attenuate apically and usually mucronate. ........................................................................................................................................ 20. S. eremophila. 8. Leaf aristae usually persistent; sporophylls each abruptly tipped by a twisted and persistent arista. ......................................................................................................................................................................... 13. S. carnerosana.

7.

2.

Leaves acute to aristate, the arista straight; upperside leaves linear-lanceolate; plants forming rather loose mats. 9. Underside leaves lanceolate, widest at middle; apices acute or each with a short and flattened arista, aristae 0.1–0.3 mm long (mostly at branch tips or buds); sporophylls acute to acuminate. .................. 3. S. arizonica. 9. Underside leaves linear-lanceolate, widest at bases; apices each with a long, terete arista, aristae 0.3–0.8 mm long; sporophylls aristate. ...................................................................................................................... 52. S. peruviana. Aerial stems erect, ascending, creeping, or decumbent, and radially symmetric; leaves monomorphic, rarely subdimorphic; rhizophores throughout or restricted to stem bases. 10. Aerial stems erect or ascending, seldom decumbent or creeping; main stems rhizomatous; bud-like arrested branches usually present on rhizomes or proximal part of aerial stem systems. 11. Leaves acute to obtuse, the margins denticulate; bases of aerial stem leaves and rhizome scale-like leaves decurrent or long-decurrent. ...................................................................................................................................................................... 79. S. viridissima. 11. Leaves aristate, the margins ciliate; bases of aerial stem leaves and rhizome scale-like leaves abruptly adnate.

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12. Leaf margins short-ciliate at leaf bases, cilia 0.02–0.08 mm long; rhizomes conspicuous; scale-like leaves persistent, bases cordate to almost peltate. .................................................................................................................... 10. S. bigelovii. 12. Leaf margins long-ciliate at leaf bases, cilia 0.06–0.2 mm long; rhizomes sometimes inconspicuous; scale-like leaves persistent or caducous, bases rounded. .............................................................................................................. 61. S. rupincola. 10. Aerial stems creeping, decumbent (or slightly dorsiventral), never erect; main stems not rhizomatous; bud-like arrested branches absent. 13. Leaf apices blunt, acute, or slightly acuminate to short-aristate, if short-aristate then the aristae reflexed, to 0.4 mm long. 14. Leaf apices blunt, acute to slightly acuminate or mucronate, not distinctly aristate, the acumen or mucro, if present, ca. 0.05 mm long; plants terrestrial or epipetric; leaves dull or grayish green. 15. Leaves loosely appressed, margins denticulate, apices plane. ............................................................................ 15. S. cinerascens. 15. Leaves tightly appressed, the margins proximally long-ciliate, apices keeled. .................................................... 45. S. mutica. 14. Leaf apices acuminate to short-aristate, tips often reflexed, 0.1–0.4 mm long or less. 16. Leaves dull, grayish green or beige when old; apices short-aristate, the aristae hyaline to whitish, (0.1–)0.2– 0.4 mm long; plants epipetric, forming short, loose mats to 10 cm long. ........................................................... 4. S. arsenei. 16. Leaves lustrous, dark green or brownish, to brownish with red spots when old; apices acuminate to shortaristate, aristae 0.1–0.2 mm long; plants epipetric or epiphytic, forming long, loose mats to 35 cm long. .. 22. S. extensa. 13. Leaf apices short- to long-aristate, aristae straight, 0.2–2 mm long. 17. Leaf bases decurrent. 18. Leaves dull green or pale green, usually distinctly keeled; aristae yellowish, whitish, or hyaline, not distinctly denticulate; lateral branches mostly ascending; plants forming compact, cushion-like (rarely loose) mats. ............................................................................................................................................................................... 6. S. asprella. 18. Leaves lustrous, bright green, not distinctly keeled; aristae green-yellow to rarely white, distinctly denticulate; lateral branches usually horizontal; plants forming mostly long and loose (rarely compact) mats. ...................................................................................................................................................................... 78. S. underwoodii. 17. Leaf bases abruptly adnate to stem. 19. Leaf bases glabrous or infrequently puberulent, if puberulent then aristae hyaline and more than 1⁄2 the lamina length. 20. Leaf apices long-aristate, aristae 1.4–2.6 mm long. .................................................................................. 37. S. macrathera. 20. Leaf apices short-aristate, aristae 0.2–0.8 mm long. 21. Stems 0.3–0.4 mm diam.; leaves denticulate to entire; aristae hyaline, 0.2–0.4 mm long. ............... 4. S. arsenei. 21. Stems 0.5–0.6 mm diam.; leaves ciliate-denticulate; aristae white, 0.5–0.8 mm long. .................... 67. S. sellowii. 19. Leaf bases pubescent to puberulent; aristae white, yellowish, or greenish hyaline, if greenish hyaline then less than 1⁄2 the lamina length. 22. Main stems suberect to erect, rarely decumbent; leaves of axis bluish to grayish green, the marginal cilia white, spreading, usually more than 15 per side; aristae white, distinctly denticulate. ................. 61. S. rupincola. 22. Main stems never ascending or erect; leaves light green with red spots or reddish when old, marginal cilia hyaline or white, ascending, usually fewer than 15 per side; aristae hyaline or whitish, not distinctly denticulate. 23. Leaf cilia and aristae white to whitish, aristae 0.8–1 mm, usually 1⁄2 the lamina length. ................. 63. S. sartorii. 23. Leaf cilia and aristae hyaline; aristae 0.5–0.8 mm, usually 1⁄3 the lamina length. ...................... 73. S. steyermarkii. 1. Plants anisophyllous; leaves 4-ranked, with 2 rows of spreading lateral leaves and 2 rows of appressed, strongly ascending median leaves. 24. Stems articulate; stele of 2 strands (1 in S. silvestris); megasporangia one basal per strobilus; megasporophylls usually twice or more as long as microsporophylls; megaspores with high ridges; microspores with sharp-pointed spines, tan or cream. 25. Median leaves with basal attachment, biauriculate, auricles either subequal or one much larger; axillary leaves biauriculate or exauriculate. 26. Leaf margins denticulate to entire; median leaves each with 2 small, subequal auricles; lateral leaves with rounded bases (not auriculate); axillary leaves with rounded bases, not auriculate. ....................................................................... 69. S. silvestris. 26. Leaf margins denticulate or ciliate; median leaves each with 1 large auricle, to 1⁄4 the lamina length; lateral leaves auriculate or truncate; axillary leaves auriculate or rounded at bases. 27. Auricles of median leaves ciliate; axillary leaves slightly biauriculate. .......................................................................... 71. S. stellata. 27. Auricles of median leaves denticulate; axillary leaves exauriculate. ................................................................................ 75. S. tarda. 25. Median leaves peltate with single auricle; axillary leaves peltate or biauriculate. 28. Lateral leaf bases peltate, each with a long, ciliate auricle extending lengthwise along stems. ....................................38. S. marginata. 28. Lateral leaves basifixed, bases with 1 or 2 short-ciliate or denticulate auricles, these clasping but not extending lengthwise along stems. 29. Bases of axillary leaves usually truncate, exauriculate on main, non-stoloniferous stems; lateral leaves on main stems oblong, bases truncate, exauriculate. ...................................................................................................................... 68. S. sertata. 29. Bases of axillary leaves auriculate, with two auricles on main, non-stoloniferous stems; lateral leaves on main stems oblong-ovate or oblong-elliptic, bases cordate to subcordate with two auricles.

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SELAGINELLA 30. Axillary leaf bases with 2 incurved ciliate auricles on main stems; lateral leaves on main stems oblong-ovate, widest at bases, each leaf base with an acroscopic auricle curving downward on stems. ............................... 21. S. eurynota. 30. Axillary leaf bases with 2 divergent, ciliate auricles on main stems; lateral leaves on main stems oblongelliptic, widest at middle, each leaf base with an acroscopic auricle not curving downward. .................... 66. S. schizobasis. 24. Stems not articulate; monostelic with single ribbon-shaped stele; megasporangia more than 1 per strobilus, or if one, not located at base; megasporophylls and microsporophylls usually of the same size but if dimorphic usually the longer sporophylls present on the upper side of the stems; megaspores with low ridges; microspores without spiny projections, variously sculptured, often capitate to clavate, orange or tan. 31. Aerial stems fasciculate, rosette-forming, or if upright and elongate, neither stoloniferous nor flagelliform; stem leaves with long-ciliate margins. 32. Plants upright, rarely rosette-forming (S. acutifolia, S. pallescens); median leaf bases truncate, rounded, cordate, or short-auriculate. 33. Axillary leaves on main stems and primary branches with rounded or truncate bases. 34. Median and lateral leaves and sporophylls acute; stems and some lateral leaves red to reddish at least on the proximal 1⁄2 of stems. ................................................................................................................................................. 1. S. acutifolia. 34. Median and lateral leaves and sporophylls long-acuminate to long-aristate, aristae 1⁄3 or more the lamina length; stems stramineous to greenish throughout, lateral leaves never reddish. ........................................... 50. S. pallescens. 33. Axillary leaves on main stems and primary branches with cordate bases, each leaf base with 2 distinct auricles. 35. Leaves on main stems below the first branch conspicuously dimorphic with the lateral leaves spreading and the median leaves ascending. ...................................................................................................................................54. S. polyptera. 35. Leaves on main stems below the first branch seemingly monomorphic with both lateral and median leaves appressed to the stems and ascending. 36. Median and lateral leaves long-ciliate at bases and noticeably ciliate most of the lamina length, especially on acroscopic (lateral leaves) or inner (median leaves) margins; 600–1550 m. ........................ 28. S. harrisii. 36. Median and lateral leaves long-ciliate at bases, otherwise denticulate or entire; 1200–2400 m. ...... 58. S. pulcherrima. 32. Plants rosette-forming; median leaf bases subpeltate to peltate. 37. Lateral leaf bases peltate with a single auricle; stems of branches glabrous; lower surfaces of lateral leaves with a brown band of cells near acroscopic margin and/or brownish throughout with age; leaf margins green to slightly green-hyaline, coarsely dentate or ciliate. ..................................................................................................................... 16. S. convoluta. 37. Lateral leaf bases basifixed, rounded or biauriculate; stems of the branches pubescent or glabrous; lower surfaces of the lateral leaves with age wine-red, reddish brown, green to yellowish, or with a yellow-brown band of cells near acroscopic margin, or frequently throughout; leaf margins green to broadly hyaline, long- to short-ciliate, fimbriate, or dentate. 38. Axillary leaves broadly ovate to ovate-deltate, bases rounded and without auricles; lower surface of the leaves with at least a wine-red or reddish brown band of cells along acroscopic margin; lateral and median leaves similar in size and shape. 39. Apices of lateral, median, and axillary leaves rounded to broadly obtuse. .................................................... 60. S. ribae. 39. Apices of lateral, median, and axillary leaves acute, mucronate, or aristate. 40. Median leaves acute to mucronate; leaf margins broadly hyaline, ca. 0.1 mm wide; median leaf bases pubescent to villous, the hairs long, distributed throughout; stems of branches pubescent to velutinous on the upper surface, the hairs conspicuous. ............................................................... 34. S. lepidophylla. 40. Median leaves long-acuminate to aristate; leaf margins narrowly hyaline, ca. 0.05 mm wide; median leaf bases puberulent, the hairs short, mostly restricted to the center and outer base; stems of branches puberulent on the upper surface, the hairs frequently inconspicuous. ................. 47. S. novoleonensis. 38. Axillary leaves lanceolate or ovate-lanceolate, bases usually cordate with two incurved ciliate auricles; lower surface of the leaves dull green to yellowish, yellow-brown or brown, never red; lateral and median leaves differing in size and shape. 41. Leaf margins broadly hyaline, short-ciliate; median leaves ovate, broadly ovate-lanceolate to ovatedeltate, the bases glabrous or weakly puberulent. ................................................................................. 46. S. nothohybrida. 41. Leaf margins green or inconspicuously narrowly hyaline, entire, with scattered teeth, or with long hairs near the bases and weakly dentate distally; median leaves lanceolate, the bases pubescent to velutinous. 42. Leaf apices rounded. ............................................................................................................................... 27. S. gypsophila. 42. Leaf apices long-aristate, aristae 1⁄3 to 1⁄2 or more the lamina length. .................................................. 53. S. pilifera. 31. Aerial stems simple, not fasciculate; main stems creeping, or if erect, stoloniferous; stem leaves denticulate or ciliate. 43. Bases of lateral, median, and axillary leaves peltate; leaves coriaceous; veins not visible. ............................................ 64. S. schaffneri. 43. Bases of lateral, median, and axillary leaves basally attached (asymmetric, oblique, rounded, cordate, sometimes auriculate); leaves chartaceous; veins apparent. 44. Stems reddish at base, pale reddish throughout main stems; stolons red and numerous; main stems erect, not creeping; axillary leaves denticulate, exauriculate. .................................................................................................... 30. S. hoffmannii. 44. Stems stramineous or green (if reddish patches present, then stems prostrate, rooting along main axis, nonstoloniferous; axillary leaves long-ciliate and biauriculate).

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45. Leaves on main stems uniform; stems stoloniferous (often flagelliform in S. mickelii), main stems erect, simple, with distal portion frond-like; rhizophores only near bases of stems. 46. Stem, median, lateral, and axillary leaves denticulate; microspores yellow-orange to orange; stems nonflagelliform; leaves on main stem appressed. 47. Lateral leaf bases rounded to subcuneate; lateral leaves with no visible idioblasts; median leaves with 1–2 short idioblasts. ........................................................................................................................... 18. S. cuneata. 47. Lateral leaf bases rounded acroscopically, truncate or subauriculate basiscopically; lateral and median leaves with numerous idioblasts. ............................................................................................. 48. S. oaxacana. 46. Stem, median, lateral, and axillary leaves conspicuously long-ciliate, or with short to long cilia restricted to leaf bases; microspores yellow-orange or tan; stems flagelliform or not; leaves on main stems appressed or spreading. 48. Lateral leaves on main stems spreading, median leaves on main stems appressed; median leaves with prominent basiscopic auricles; microspores tan. ................................................................. 44. S. mosorongensis. 48. Lateral and median leaves on main stem appressed; median leaves exauriculate; microspores yelloworange. 49. Leaves on main stems ciliate only at bases; median leaves entire except for few long cilia at basiscopic bases; axillary leaves long-ciliate at bases; stems non-flagelliform. ........................ 32. S. illecebrosa. 49. Leaves on main stems short-ciliate along margins; median leaves long-ciliate along margins; axillary leaves long-ciliate for most of length; stems often flagelliform. ................................... 40. S. mickelii. 45. Leaves everywhere dimorphic; stems stoloniferous or not, main stems prostrate, ascending, suberect or erect with rhizophores throughout, or suberect with rhizophores confined to proximal 1⁄2 of stems. 50. Upper surfaces of leaves verruculose or hispid, at least near basiscopic margins of lateral leaves. 51. Upper surfaces of lateral leaves distinctly hispid, at least near basiscopic margins. 52. Upper surfaces of lateral leaves hispid throughout. ....................................................................... 35. S. lindenii. 52. Upper surfaces of lateral leaves hispid near basiscopic margins and occasionally near apices. 53. Acroscopic margins of lateral leaves long-ciliate, at least on proximal 1⁄4; median leaves usually with an outer auricle. 54. Margins of median leaves long-ciliate; upper surfaces corrugate and frequently verruculose; lateral leaves with short hairs along basiscopic half (particularly near margins). ........................................................................................................................... 29. S. hirtifolia. 54. Margins of median leaves ciliate-denticulate to entire; upper surfaces of lateral leaves with hairs mostly submarginal, otherwise smooth. 55. Outer bases of median leaves glabrous; megaspores snow-white. ....................... 41. S. minima. 55. Outer bases of median leaves short-ciliate; megaspores orange. .................... 43. S. moritziana. 53. Acroscopic margins of lateral leaves entire to denticulate; median leaves without a distinct auricle. 56. Plants not stoloniferous, creeping; rhizophores borne mostly throughout the stems; lateral leaves ovate-elliptic to ovate-oblong, abruptly tapering at apices, without false veins. ............................................................................................. 55. S. popayanensis. 56. Plants stoloniferous, creeping to suberect; rhizophores borne mostly restricted to proximal 1⁄4 to 1⁄2 of stems; lateral leaves ovate to ovate-lanceolate, gradually tapering toward apices, with or without false veins parallel to midvein. 57. Median leaves gradually short- to long-acuminate or short-aristate, aristae 1 ⁄4 or less the lamina length, 0.1–0.3 mm long; upper surfaces of lateral and median leaves with conspicuous or inconspicuous idioblasts. ............... 43. S. moritziana. 57. Median leaves abruptly long-aristate, aristae 1⁄4 to 1⁄2 or more the lamina length, 0.3–1 mm long; upper surfaces of lateral and median leaves without idioblasts. 58. Median leaves on main stems broadly ovate-elliptic; lateral leaves without false veins, the upper surfaces usually hispid only near basiscopic margin, apices acute; megaspores cream-yellow; apices of stem and branches only occasionally flagelliform or gemmiferous. ............................................................................................. 9. S. bernoullii. 58. Median leaves on main stem ovate to ovate-lanceolate; lateral leaves usually with 1 or 2 false veins parallel to midvein, upper surfaces hispid near basiscopic margin and apices, apices acuminate; megaspores orange; apices of stem and branches usually flagelliform and gemmiferous. ..................................................................................... 56. S. porphyrospora. 51. Upper surfaces of lateral leaves verruculose (with papilla-like projections) or with prominent round cells throughout. 59. Apices of median leaves long-aristate, aristae 1⁄2 or more the lamina length.

556

SELAGINELLA 60. Margins of median leaves denticulate to entire, upper surfaces striate to weakly verruculose; axillary leaves and acroscopic margins of lateral leaves denticulate to entire. .......................................................................................................................................... 25. S. flexuosa. 60. Margins of median leaves long-ciliate, upper surfaces strongly verruculose to corrugate; axillary leaves and acroscopic margins of lateral leaves long-ciliate, at least on proximal 1 ⁄2. 61. Median leaves suborbicular to broadly ovate, 0.7–1 mm wide, margins greenish proximally to hyaline (1–2 cells wide) distally, aristae 1⁄3–1⁄2 the lamina length; 1000– 1400 m. ............................................................................................................................ 7. S. barnebyana. 61. Median leaves ovate-lanceolate or broadly ovate, 0.5–0.7 mm wide, margins broadly hyaline throughout (3 or more cells wide), arista 1⁄2 or more the lamina length; 160– 400 m. .................................................................................................................. 31. S. huehuetenangensis. 59. Apices of median leaves acuminate to short-aristate, arista 1⁄4 or less the lamina length. 62. Lateral leaves ca. 1 mm long, broadly ovate, acroscopic margins denticulate, apices obtuse to rounded; rhizophores filiform, ca. 0.1 mm diam. ................................................ 17. S. corrugis. 62. Lateral leaves 1.7–3 mm long, ovate-oblong to suborbicular, acroscopic margins denticulate or ciliate near base or throughout, apices acute to broadly acute; rhizophores stout, 0.2–0.5 mm diam. 63. Median leaves denticulate, surfaces conspicuously verruculose. .............................. 74. S. subrugosa. 63. Median leaves distinctly ciliate, surfaces smooth or inconspicuously verruculose. 64. Median leaves broadly ovate to orbicular. ........................................................... 14. S. chiapensis. 64. Median leaves ovate. ......................................................................................... 26. S. guatemalensis. 50. Upper surfaces of leaves smooth, wrinkled, or striate, not verruculose or hispid. 65. Lateral leaves denticulate, ciliate-denticulate, or short-ciliate only at acroscopic base. 66. Plants prostrate, rhizophores borne throughout stem length (or if restricted to proximal 1⁄2 of stems, then upper surface of leaves with idioblasts), creeping, never flagelliform or stoloniferous. 67. Strobili quadrangular, 1–2 mm wide; median leaves each with a prominent, raised midrib. 68. Median leaves long-aristate, bases glabrous; rhizophores usually borne on the lower side of the stem, stramineous to green; megaspores white. ......................................... 25. S. flexuosa. 68. Median leaves acute, bases ciliate; rhizophores borne on the upper side of the stem, dark brown; megaspores pale orange to deep yellow. ............................................... 62. S. rzedowskii. 67. Strobili flattened, 2–4 mm wide; median leaves each without a prominent, raised midrib. 69. Upper and lower sporophylls spreading, green, symmetric, ventral ones longer than dorsal ones; lateral leaves 1.4–2.3 mm long, light green; megaspores bone-white to pale cream. ...............................................................................................................................2. S. apoda. 69. Upper sporophylls spreading, green, asymmetric; lower sporophylls ascending, hyaline, smaller than dorsal ones; lateral leaves 2–3.7 mm long, silvery beneath; megaspores white or orange. 70. Median leaves frequently with two auricles at the bases or with only well developed outer auricles; margins long-ciliate along proximal 1⁄2; rhizophores 0.5– 0.8 mm diam.; lateral leaf apices broadly acute to rounded; megaspores white to cream. ..................................................................................................................... 72. S. stenophylla. 70. Median leaves with an asymmetric base, without auricles; margins denticulate; rhizophores 0.1–0.3 mm diam.; lateral leaves with acute to long-acuminate apices; megaspores orange. 71. Median leaves gradually acuminate to short-aristate, aristae 1⁄4 or less the lamina length; lateral leaves with or without idioblasts, if present then with many short idioblasts, these not distinctly parallel to midrib. ............... 43. S. moritziana. 71. Median leaves abruptly aristate, aristae 1⁄2 or more the lamina length; lateral leaves with 1 or 2 elongate idioblasts parallel to midrib along most of its length. ......................................................................................................... 56. S. porphyrospora. 66. Plants erect or suberect, rhizophores restricted to proximal 1⁄2 of stems, or plants with combination of suberect and prostrate habit with flagelliform and stoloniferous stems. 72. Lateral leaves 2.8–5 mm long; megaspores white. 73. Rhizophores 0.8–1.2 mm diam.; stems not stoloniferous, 2–2.5 mm diam. ........... 39. S. martensii. 73. Rhizophores 0.1–0.3 mm diam.; stems stoloniferous, 0.5–1.4 mm diam. ................ 77. S. tuberosa. 72. Lateral leaves 1.2–3 mm long; megaspores white or yellow. 74. Leaves with hyaline to whitish idioblasts on upper surfaces. 75. Median leaf bases with an outer auricle. ............................................................... 36. S. lineolata.

SELAGINELLA

557

75. Median leaf bases without an outer auricle. ............................................................ 76. S. tenella. 74. Leaves without hyaline or whitish idioblasts on upper surfaces. 76. Mature, strobilus-bearing plants 1–2 cm tall, ascending to suberect, not flagelliform or stoloniferous; megaspores yellow. .................................................. 70. S. simplex. 76. Mature, strobilus-bearing plants 5–32 cm tall, suberect to erect, usually flagelliform or stoloniferous; megaspores white or cream. 77. Median leaves marginally short-ciliate along distal 1⁄2; stems only occasionally flagelliform. ................................................................................... 23. S. finitima. 77. Median leaves marginally denticulate; stems regularly flagelliform. ......... 24. S. flagellata. 65. Lateral leaves short-ciliate for entire length, or long-ciliate for portion of margin. 78. Axillary leaves auriculate and long-ciliate; older leaves turning reddish. ............................ 19. S. delicatissima. 78. Axillary leaves exauriculate and long-ciliate to denticulate; older leaves green, white or tan, not reddish. 79. Plants prostrate; rhizophores borne throughout stem length. 80. Median leaf apices broadly acute, margins short-ciliate at bases, denticulate distally; lateral leaves strongly imbricate; branch tips ascending. ............................................... 5. S. arsiclada. 80. Median leaves cuspidate to aristate, margins ciliate throughout leaf length; lateral leaves not overlapping; branch tips horizontal. 81. Axillary leaves ovate to ovate-deltate; median leaves ovate to ovate-deltate, distant or not strongly imbricate. ........................................................................ 65. S. schiedeana. 81. Axillary leaves obovate; median leaves ovate to suborbicular, usually strongly imbricate. 82. Median leaf bases subcordate, each with a distinct inner auricle, auricles tufted with 3–6 whitish hairs; acroscopic base of lateral leaves with 5–8 hairs; median leaf margins and basiscopic margin of lateral leaves 4–6 (or more) cells wide. .................................................................................................... 12. S. carioi. 82. Median leaf bases oblique to subcordate, inner bases not auriculate, glabrous; acroscopic bases of lateral leaves short-ciliate; median leaf margins and the basiscopic margin of lateral leaves 3 or fewer cells wide. ................. 59. S. reflexa. 79. Plants suberect to erect, rarely creeping; rhizophores borne only in the proximal third of stems. 83. Mature, strobilus-bearing plants 1–2 cm tall, never flagelliform or stoloniferous; sporophylls asymmetric, spreading, each with a prominent keel. 84. Megaspores white; lateral leaves long-ciliate along proximal 1⁄4–1⁄2 of acroscopic margin; axillary leaves long-ciliate in proximal 1⁄4–1⁄2. .......................................... 41. S. minima. 84. Megaspores yellow; lateral leaves short-ciliate along proximal 1⁄4–1⁄2 of acroscopic margin; axillary leaves short-ciliate in proximal 1⁄4–1⁄2. ........................................ 70. S. simplex. 83. Mature, strobilus-bearing plants over 5 cm tall, occasionally flagelliform or stoloniferous; sporophylls keeled or not. 85. Rhizophores stout and straight, 0.4–1.2 mm diam.; sporophylls lacking keels. 86. Rhizophores 0.4–0.5 mm wide; surfaces of lateral leaves hispid; axillary leaves ciliate in proximal 1⁄4; median leaves acuminate. ...................... 26. S. guatemalensis. 86. Rhizophores 0.8–1.2 mm diam.; surfaces of lateral leaves smooth; axillary leaves denticulate; median leaves long-aristate. ........................................... 39. S. martensii. 85. Rhizophores filiform, 0.3 mm or less diam.; sporophylls each with slightly to well developed keel. 87. Megaspores white; sporophylls short- to long-ciliate. 88. Lateral leaves lacking white idioblasts; stems upright and frond-like to creeping, commonly flagelliform; Atlantic slope and Isthmus of Tehuantepec. .............................................................................................. 40. S. mickelii. 88. Lateral leaves with scattered white idioblasts; stems suberect to erect, rarely flagelliform (or upright and frond-like); Pacific slope and Isthmus of Tehuantepec. .......................................................................... 57. S. prolifera. 87. Megaspores yellow; sporophylls denticulate to very short-ciliate. 89. Median leaves on main stems narrowly lanceolate, entire to denticulate; lateral leaves acuminate, short-ciliate at bases. ................................... 11. S. breedlovei. 89. Median leaves on main stems ovate, long- to short-ciliate or denticulate; lateral leaves acute to obtuse, long-ciliate at bases. 90. Median leaves short-ciliate to denticulate; Oaxaca. ..................... 42. S. mixteca. 90. Median leaves long- to short-ciliate; Veracruz. ........................ 49. S. orizabensis

558

SELAGINELLA

1. SELAGINELLA ACUTIFOLIA Selaginella acutifolia (Stolze) Valdespino, comb. et stat. nov. Figs. 284A–G. Selaginella pallescens (C. Presl) Spring var. acutifolia Stolze, Amer. Fern J. 71: 51. 1981. Type. Guatemala. Zacapa: near Santa Rosalı´a, 2 mi S of Zacapa, Steyermark 29293 (F!).

Stems fasciculate from a rhizomatous base, rosette-forming or occasionally erect, 3–30 cm long, 0.5–1 mm diam., not articulate, green to reddish, 2–3 times branched; rhizophores restricted to stem bases, filiform, 0.2–0.5 mm diam.; leaves of two kinds (anisophyllous) throughout, green or reddish above, silvery to lustrous green, golden, or reddish below; lateral leaves imbricate, ascending, broadly ovate to ovate-deltate, 1–2 ⫻ 0.6–1.2 mm, bases rounded to subcordate, ciliate, margins hyaline, short- to long-ciliate proximally, denticulate distally, apices acute; median leaves imbricate to slightly distant, broadly ovate to ovate-deltate, 1–1.5 ⫻ 0.8–1.2 mm, glabrous, bases subcordate, margins hyaline, short-ciliate proximally, denticulate distally, apices acute; axillary leaves ovate to lanceolate, bases cordate, long-ciliate, margins hyaline, ciliate throughout or denticulate distally, apices acute; strobili terminal on branches, quadrangular, 3–10 mm long; sporophylls monomorphic; megasporangia mostly basal in two ventral rows; microsporangia in two dorsal rows and distally also in ventral rows; megaspores lemon-yellow, reticulate, with or without a low equatorial flange, 240–260 ␮m diam.; microspores orange.

Lycopodium apodum L., Sp. Pl. 2: 1105. 1753. Type. U.S.A. “Carolina, Virginia, Pensylvania,” Kalm s.n. (LINN 1257.27).

Stems prostrate to short-creeping, forming rather loose or clustered mats 3–7 cm long, 0.3–0.5 mm diam., greenish to stramineous, not articulate, not flagelliform, not stoloniferous, sparsely branched, the branches mostly simple or 1-forked; rhizophores borne throughout stem length or restricted to proximal 1 ⁄3, less than 0.1 mm diam.; leaves of two kinds (anisophyllous) throughout; lateral leaves ovate to ovate-lanceolate, 1.4–2.3 ⫻ 0.8–1.4 mm, bases rounded, margins green or with 1 row of hyaline cells, denticulate, apices acute; median leaves lanceolate, surfaces glabrous, 1–1.6 ⫻ 0.4–0.7 mm, bases rounded, margins sparingly denticulate or subentire, apices acuminate to longacuminate; axillary leaves similar to lateral ones, exauriculate; strobili lax and dorsiventral, 1–2 cm long; sporophylls subdimorphic, dorsal ones spreading, ventral ones ascending; megaspores pale yellow to bone-white, reticulate, 330–390 ␮m diam.; microspores orange, rugulose-reticulate to verruculose, 25–30 ␮m diam.

Distribution. Terrestrial in swamps, meadows, marshes, pastures, damp lawns, shaded slopes, light woods, and stream banks, in basic to acidic soil conditions; 1700–2700 m. E USA; Mexico; Guat.

Distribution. Terrestrial or epipetric on shaded trail banks and road banks; 0–150 m. Mexico; Guat, Bel, Hond, Salv, Nic, CR. Specimens Examined. Jal (Lott & Wendt 1593, UC). Mich (Guerrero 991, 1216, XAL). Oax (Mickel 6905, NY).

Selaginella acutifolia is characterized throughout its range by acute leaf apices and red-pigmented leaves and stems; it is here considered a species distinct from S. pallescens.

2. SELAGINELLA APODA Selaginella apoda (L.) Spring in Martius, Fl. Bras. 1: 119. 1840 [“apus”]. Figs. 285A–F.

Selected Specimens Examined. Chis (Breedlove 21287, DS, NY, 39780, DS, ENCB, NY; Little & Sharp 9876, US; Mu¨nch s.n., DS). Hgo (Pringle 8797, ARIZ, CAS, MSC, NY, UC, US). Mich (Garcı´a & Dı´az Barriga 3854, NY; Rzedowski 48537, NY; Zamudio & Becerra 10440, NY). Pue (Pringle 10815, ARIZ, CAS, LL, MICH, MSC, UC, US). Ver (Nee & Taylor 26781, F, NY).

Selaginella apoda is the central component of a taxonomically difficult species complex of the eastern and southeastern United States. Its closest relative is S. eclipes W. R. Buck. It is also related to S. ludoviciana (A. Braun) A. Braun, from which it can be distinguished by its green or less often narrowly hyaline leaf margins, stomata scattered throughout the leaves, tendency to form rather short mats of prostrate or creeping stems, and rugulose-reticulate to verruculose microspores. Selaginella ludoviciana has lateral leaves with hyaline margins 3–5 cells wide, stomata restricted to the midrib region, long-creeping to ascending stems, and rather smooth microspores.

SELAGINELLA ARSICLADA

3. SELAGINELLA ARIZONICA Selaginella arizonica Maxon, Smithsonian Misc. Collect. 72: 5. 1920. Figs. 274U–Z. Bryodesma arizonica (Maxon) Soja´k, Preslia 64: 154. 1992. Type. U.S.A. Arizona: Foot of Soldier Trail, Santa Catalina Mountains, Shreve s.n. (US!; isotypes GH, MO!, NY!).

Stems prostrate, forming rather loose mats, not readily fragmenting, dorsiventral, tips upturned in extremely dry conditions, 6–15 cm long, 0.5–0.7 mm diam., not articulate, not flagelliform, not stoloniferous, irregularly branched, branches determinate, 2–3 times branched; rhizophores borne throughout stem length, 0.2–0.3 mm diam.; leaves of one kind (isophyllous) throughout, tightly appressed to ascending, green, apices aristate, sometimes becoming acute by breaking off of aristae, aristae flattened, transparent to opaque, 0.1–0.3 mm, dentate at margins; upperside leaves linear-lanceolate, slightly falcate, 1.3–2.3 ⫻ 0.4– 0.6 mm, bases abruptly adnate, pubescent or glabrous, margins green, ciliate, the cilia hyaline to opaque, spreading; underside leaves lanceolate, falcate, 2–2.5 ⫻ 0.5–0.6 mm, bases decurrent, glabrous, margins ciliate, the cilia hyaline to opaque, spreading or ascending; strobili quadrangular, 5–10 mm long; sporophylls monomorphic; megasporangia in two ventral rows; microsporangia in two dorsal rows; megaspores yellow, rugose to reticulate, 320–340 ␮m diam.; microspores orange.

559

Bryodesma arsenei (Weath.) Soja´k, Preslia 64: 154. 1992. Type. Mexico. .Quere´taro: Quere´taro, Arse`ne 10641 (US!, frag. GH)

Stems long-creeping, forming loose mats, to 10 cm long, 0.3– 0.4 mm diam., radially symmetric, not readily fragmenting, tips straight, 0.3–0.4 mm diam., not articulate, not flagelliform, not stoloniferous, 1–2 times branched; rhizophores borne throughout stem length, stout, (0.1–)0.2–0.4 mm diam.; leaves of one kind (isophyllous) throughout, dull, grayish green (beige when old), ascending, linear, 1.8–3 ⫻ 0.3–0.4 mm, bases truncate, abruptly adnate to the stems, glabrous or infrequently pubescent, margins green, denticulate to entire, the teeth hyaline, ascending, apices short-aristate, aristae hyaline to whitish, (0.1–)0.2–0.4 mm long; strobili quadrangular, 0.5–1.3 cm long; sporophylls monomorphic; megasporangia few, scattered on ventral side; microsporangia in two dorsal rows and scattered on ventral rows; megaspores lemonyellow, rugose to reticulate, 360–400 ␮m diam.; microspores orange, 40–50 ␮m diam.

Distribution. Epipetric or epiphytic on rock crevices; 2700 m fide R. Tryon (1955). Mexico. Specimens Examined. Gro (Frye & Frye 3146, UC, US). Hgo (Frye & Frye 3149, DS, UC; Hitchcock & Stanford 7248, US; Stuessy 296, TEX; Tryon & Tryon 5133, MEXU). Qro (Arse`ne 9983, US, 10643, US; Basile 2, NY). SLP (Frye & Frye 2983, F, LL).

Distribution. Terrestrial or epipetric, in rock crevices or on gravel, on sandstone, igneous, or rarely limestone substrates; 500–2200 m. USA; Mexico. Selected Specimens Examined. BCS (Jones s.n., NY; Moran 18820, DS). NL (Hinton et al. 20084, MEXU, TEX, 21112, TEX). Son (Felger & Harris 16719, MEXU; Fischer 6858, ENCB; Mearns 2701, NY; Strickland & Felger 20481, ENCB; Wilkinson s.n., ENCB).

Selaginella arizonica can be distinguished from the similar S. peruviana by its underside leaves broad and thin, versus narrow and fleshy in S. peruviana.

4. SELAGINELLA ARSENEI Selaginella arsenei Weath., J. Arnold Arbor. 25: 417, pl. II, f. 8. 1944. Figs. 276M–O.

Selaginella arsenei is characterized by its abruptly adnate leaf bases and by its relatively short aristae. It can be separated from S. sellowii by the characters given in the key, couplet 21. Selaginella arsenei may occasionally grow as an epiphyte on trees (Frye & Frye 2983).

5. SELAGINELLA ARSICLADA Selaginella arsiclada Valdespino, Amer. Fern J. 84: 99. 1994. Type. Mexico. Quere´taro: Landa, Tangojo cerca del Rı´o Moctezuma, Dı´az Barriga 4627 (NY!; isotypes IEB!, NY!, PMA!). Figs. 284U–Z. Stems creeping, green to stramineous, 2–12 cm long, 0.3–0.5 mm diam., not articulate, not flagelliform, not stoloniferous, 2–3 times branched; rhizophores borne throughout stem length, 0.2– 0.3 mm diam.; leaves of two kinds (anisophyllous) throughout,

560

SELAGINELLA

surfaces green or yellowish to brown when old; lateral leaves broadly ovate to ovate-deltate on main stems, ovate-oblong to oblong on branches, 1–1.4 ⫻ 0.5–0.9 mm, margins whitish in a band 1(–2) cells wide, short-ciliate (to long-ciliate basiscopically), denticulate distally, apices obtuse; median leaves ovate, 0.6–0.8 ⫻ 0.4–0.6 mm, bases oblique to slightly cordate, margins whitish in a band 3–4 cells wide, short-ciliate most of the length, denticulate distally, apices broadly acute; axillary leaves similar to lateral ones except bases more obviously cordate; strobili quadrangular, 2–5 mm long; sporophylls monomorphic; megasporangia in two ventral rows; microsporangia in two dorsal rows; megaspores white, smooth, 200–240 ␮m diam.; microspores orange with capitate projections, 21–28 ␮m diam.

Stems decumbent to short-creeping, to 10 cm long, forming cushion-like or loose mats, main stems dorsiventral, lateral branches radially symmetric, often strongly ascending, readily fragmenting, tips straight, 0.3–0.4 mm diam., not articulate, not flagelliform, not stoloniferous, irregularly branched 1–2 times; rhizophores borne throughout stem length, 0.2–0.3 mm diam.; leaves of one kind (isophyllous) throughout, tightly appressed, lanceolate to linear-lanceolate, (2–)2.5–4 ⫻ 0.5–0.7(–0.8) mm (smaller on young shoots), bases cuneate and decurrent or sometimes rounded and adnate on young shoots, pubescent (hairs often covering 1⁄4 of lamina length abaxially), margins ciliate, cilia transparent to whitish, spreading, apices attenuate or obtuse, aristate, aristae yellowish, whitish or hyaline, denticulate, 0.5–1.4 mm; strobili quadrangular, 0.4–1.5(–2) cm long; sporophylls monomorphic; megasporangia few, basal; microsporangia occupying most of strobilus; megaspores lemon-yellow, rugose to reticulate, 290–380 ␮m diam.; microspores light orange, 33–35 ␮m diam.

Distribution. Epipetric or terrestrial on dry clay at river banks in seasonally dry areas, or on soil and rocks in moist forests; 200–600 m. Mexico. Specimens Examined. Qro (Dı´az Barriga 4267, NY). Ver (Purpus 8463, MO p.p., 8464 p.p., UC, US, 117 [10117], US; Sperling 4049, XAL).

Selaginella arsiclada does not seem closely related to other Mexican Selaginellas. It may be confused with S. reflexa and S. schiedeana because of its creeping habit, axillary rhizophores that are borne throughout the stems, ovate median leaves, and whitish leaf margins. Selaginella arsiclada can be distinguished from the other two species by its median leaves with broadly acute (vs. long-acuminate to aristate) apices, short-ciliate or denticulate margins proximally, and denticulate margins distally (not long- to short-ciliate throughout). It differs further from S. reflexa in the axillary leaves on the main stems being similar to the lateral leaves (vs. obovate in S. reflexa), and from S. schiedeana by its lateral leaves with apices gradually tapering (vs. almost equal in width to the base in S. schiedeana).

6. SELAGINELLA ASPRELLA Selaginella asprella Maxon, Smithsonian Misc. Collect. 72: 6.1920. Figs. 273P–S. Bryodesma asprella (Maxon) Soja´k, Preslia 64: 154. 1992. Type. U.S.A. California: west end of Ontario Peak, San Antonio Mountains, Johnston 1815 (US!; isotype MO!).

Distribution. Epipetric or terrestrial on limestone ridges, dry rocky slopes, igneous rocks, rock crevices, exposed cliffs, or gravelly soil; 2600–3100 m. USA; Mexico. Specimens Examined. BCN (Moran 15276, ARIZ, MEXU, 15451, UC, 21132, MSC, 25602, UC; Wiggins 16658, ENCB).

Selaginella asprella has readily fragmenting stems and has been reported in Mexico only in Baja California Norte, which sets it apart from the other selaginellas in the flora.

7. SELAGINELLA BARNEBYANA Selaginella barnebyana Valdespino, sp. nov. Type. Mexico. Chiapas: Mpio. El Bosque, 7 km NE of the Jitotol-Pichucalco junction on the road from Bochil to Simojovel, Breedlove 32475 (holotype MICH!; isotypes DS, NY!). Figs. 283M, N. A S. huehuetenangensi foliis medianis suborbicularibus vel orbicularibus, apicibus aristatis viridulis, aristis longitudine 1⁄3–1⁄2 folium aequantibus, basibus subcordatis biauriculatis, megasporis rugoso-reticulatis differt. (Dedicated to the late Rupert Barneby, distinguished student of legumes at The New York Botanical Garden.)

SELAGINELLA BASIPILOSA

Stems creeping, stramineous, ca. 30 cm long, 0.7–1.6 mm diam., not articulate, not flagelliform, not stoloniferous, 2–3 times branched; rhizophores borne throughout stem length, stout, 0.3–0.7 mm diam.; leaves of two kinds (anisophyllous) throughout, lacking idioblasts; lateral leaves ovate-oblong, 1.7– 2.5 ⫻ 0.7–1.1 mm, bases subcordate, margins greenish to slightly hyaline, long-ciliate on acroscopic bases, denticulate to entire distally, apices obtuse to broadly acute, upper surfaces corrugate, lower surfaces glabrous, without idioblasts; median leaves suborbicular to orbicular, 0.9–1.4 ⫻ 0.7–1 mm, bases subcordate to auriculate, auricles ciliate, margins greenish proximally, hyaline distally, long-ciliate (particularly the inner margin and the distal 1 ⁄2 of outer margin), apices long-acuminate, tips 0.4–0.7 mm (1⁄3–1⁄2 the lamina length), upper surfaces corrugate; axillary leaves ovate to ovate-lanceolate, bases subcordate, margins longciliate proximally, denticulate distally, apices acute to broadly acute; strobili quadrangular, 3–17 mm long; sporophylls monomorphic; megasporangia along ventral two rows; microsporangia in dorsal two rows or occupying most of strobili; megaspores white or cream, without equatorial flange, rugose-reticulate, 250–300 ␮m diam.; microspores orange.

Distribution. Terrestrial on slopes or epipetric on shaded, wet rocks in tropical montane forests; 1000–1400 m. Mexico. Specimens Examined. Chis (Gittins 4244, 4251, NY; Matuda 2782, NY; Me´ndez 5731, CAS; Mu¨nch 2, DS, 39, DS, NY, US).

Specimens of S. barnebyana were treated by Smith (1981) as S. huehuetenangensis. Selaginella barnebyana has long-aristate leaf apices resembling those of S. huehuetenangensis, but S. barnebyana seems to be restricted to high elevations (1000–1400 m), whereas S. huehuetenangensis is found at low elevations (600 m or less). Selaginella barnebyana differs further from S. huehuetenangensis by having median leaves suborbicular to orbicular (vs. ovate-lanceolate or broadly ovate), with green and stouter (vs. hyaline and delicate) apices that are 1⁄3–1⁄2 (vs. 1⁄2 or more) the lamina length, the bases subcordate (vs. oblique) with inner and outer (vs. outer only) auricular lobes, both auricles with long cilia, the distal 1⁄2 of the margins obscurely (vs. conspicuosly) hyaline. Furthermore, S. barnebyana is distinct in axillary leaves with greenish to obscurely (vs. broadly) hyaline margins; lateral leaves with acroscopic mar-

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gins greenish to obscurely (vs. broadly) hyaline; and megaspores rugose-reticulate (vs. granulate to tuberculate). Selaginella barnebyana can be confused with S. guatemalensis, from which it differs by its median leaves suborbicular to orbicular or broadly ovate (vs. ovate to ovate-lanceolate), with longaristate (vs. acute to acuminate) apices 1⁄3–1⁄2 (vs. 1⁄4) the lamina length. This species is part of the S. flexuosa group (see S. chiapensis for discussion).

8. SELAGINELLA BASIPILOSA Selaginella basipilosa Valdespino, Brittonia 44: 314. 1992. Type. Mexico. Oaxaca: Distrito Tlacolula, E of Oaxaca on Rt. 190 near Km 36, Mickel 4965 (NY!; isotypes NY!, PMA!, UC!). Figs. 266D–G. Stems prostrate, forming rather loose mats, dorsiventral, 5–17 cm long, 0.4–0.8 mm diam., not articulate, not flagelliform, not stoloniferous, 2–3 times branched, branches horizontal or curling upward (in dry conditions); rhizophores borne throughout stem length, 0.2–0.3 mm diam.; leaves of one kind (isophyllous) throughout; upperside leaves spreading, green to green-grayish, or light brown (in old leaves), narrowly deltate-lanceolate to linear-lanceolate, slightly falcate on lateral rows, 1.7–2.5 ⫻ 0.4– 0.6 mm (wider at base), bases abruptly adnate (distinct from stems), truncate to slightly rounded, strongly pubescent, hairs often running along ridges bordering abaxial grooves, margins ciliate, cilia hyaline, 0.1–0.3 mm, apices acuminate to shortcuspidate, hyaline, 0.1–0.2 mm; underside leaves buff to light brown or golden brown, lanceolate, 2.5–3.5 ⫻ 0.5–0.9 mm (wider at middle), bases rounded to subcordate, weakly decurrent, pubescent, rarely glabrescent, apices acuminate, shortaristate, aristae hyaline, whitish or yellowish, 0.1–0.4 mm, margins ciliate, cilia hyaline, denticulate distally; strobili quadrangular, 4–6 mm long; sporophylls monomorphic, green to grayish green; megasporangia usually in one ventral row; microsporangia in two dorsal rows and one ventral row; megaspores lemon-yellow, shallowly reticulate-rugose, 330–400 ␮m diam.; microspores light-orange, granulose, 40–53 ␮m diam., shed mostly in tetrads.

562

SELAGINELLA

and the outer bases auriculate or oblique, margins hyaline, sparsely denticulate, apices abruptly long-aristate, aristae 1⁄4–1⁄2 the lamina length, 0.3–1 mm long, both surfaces glabrous, without idioblasts; axillary leaves similar to lateral leaves, bases rounded; strobili dorsiventral, lax and flattened, 2–6 mm long; sporophylls dimorphic, the dorsal ones green, spreading, the ventral ones hyaline to green-hyaline, ascending; megasporangia few, 1–3 at the base of ventral rows; microsporangia in two dorsal rows or strobili almost wholly microsporangiate; megaspores creamyellow, usually without an equatorial flange or with a slightly developed one, rugulate to rugulate-reticulate, 250–300 ␮m diam.; microspores red-orange with capitate projections.

Distribution. Epipetric on dry, rocky slopes; ca. 1500 m. Mexico. Specimens Examined. Oax (Rowell et al. 17M489, MEXU, TEX; Valdespino et al. 1126, BM, MEXU, MICH, MO, NY, PMA, UC, US).

Selaginella basipilosa belongs to subg. Tetragonostachys series Eremophilae. Among the species of series Eremophilae it is most closely related and easily confused with S. landii. Selaginella basipilosa differs from S. landii by its narrowly lanceolate to linearlanceolate dorsal leaves (vs. ovate-deltate to ovate-lanceolate), with apices acuminate to short-cuspidate (vs. muticous, toothed tip); margins ciliate throughout (vs. short-ciliate at bases and denticulate toward apices), cilia and teeth hyaline (vs. white to white-hyaline); sporophylls with upper surface minutely papillose and seemingly smooth (vs. warty, especially along ridges bordering abaxial grooves); sporophyll apices acute to shortacuminate (vs. long-attenuate and aristate); and ventral leaf bases pubescent (vs. glabrous). Additionally, the distal face of megaspores in S. basipilosa has an open reticulum (vs. close reticulum) with low ridges (vs. high ridges) and the microstructure is foveolate with coarse elements (vs. smoother elements).

9. SELAGINELLA BERNOULLII Selaginella bernoullii Hieron., Hedwigia 41: 192. 1902. Lectotype (chosen here). Guatemala. Quiche´: between Cubulco and Joyabaj, Bernoulli 1121 (B!; isolectotypes B!, K!, P! [on same sheet with Bernoulli & Cario 160]). Figs. 272O–T. Stems suberect, stramineous or greenish, 9–25 cm long, 0.3– 1 mm diam., not articulate, occasionally flagelliform, stoloniferous with buds at tips, 2–3 times branched; rhizophores restricted to proximal 1⁄4–1⁄3 of stems, filiform, 0.1–0.3 mm diam.; leaves of two kinds (anisophyllous) throughout, or above proximal 1⁄4 of stems; lateral leaves distant, ovate to ovate-oblong, 2.4–5 ⫻ 1–2.4 mm, bases rounded to subcordate, margins hyaline, entire to sparingly denticulate, especially distally, apices acute, upper surfaces usually hispid, especially near basiscopic margins, without idioblasts, lower surfaces glabrous, usually with conspicuous idioblasts; median leaves broadly ovate-elliptic, 1.3– 2.2 ⫻ 0.7–1.2 mm, bases usually with the inner bases truncate

Distribution. Terrestrial or epipetric on wet and shady slopes and road banks, stream banks, or wet mossy rocks; 1300–3000 m. Mexico; Guat, Hond, Salv. Selected Specimens Examined. Chis (Breedlove 19967, DS, MO, NY, 22688, 22825, DS, NY, 29437, DS, F, NY, 31894, DS, NY, 46258, CAS, ENCB, MO, NY; Purpus 6720, US, 6721, BM, GH, NY, UC, US). Oax (Hellwig & Whitaker 1293, MEXU, NY; Mickel 5092, NY, UC, 6192, NY).

As pointed out by Valdespino (1995), S. bernoullii has been misunderstood and synonymized under S. porphyrospora because of its hispid upper surfaces of lateral leaves and sporophylls, abruptly aristate median leaves, and frequent tubers at the bases of upright stems. Selaginella bernoullii can be distinguished from S. porphyrospora by lack of idioblasts on the upper surface of lateral leaves (vs. usually with 2 false veins, one at each side of the midrib or at least at one side), lateral leaves usually acute (vs. acuminate), and occasional flagelliform branches ending in an illdifferentiated, elongate, and rather flattened propagule with denticulate leaves (vs. flagelliform branches frequently ending in a well differentiated, round, and globose propagule or bud with ciliate leaves in S. porphyrospora). In addition, S. bernoullii differs from S. porphyrospora by its cream (vs. orange) megaspores, with or without an ill-differentiated (vs. with a well differentiated) equatorial flange, and orange (vs. reddish) microspores. In addition to reproducing sexually, Selaginella bernoullii propagates vegetatively by sending out stolons that form a tuberlike bud at the tip. These buds are reddish (Moran 5548, NY), rather elongate, stout, and covered with hyaline, scale-like leaves.

SELAGINELLA BREEDLOVEI

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Upon germination they develop suberect, branched stems. This new plant in turn can produce new stolons, repeating the cycle. Selaginella bernoullii is also characterized by asymmetric lateral and axillary leaves. In both cases the broader side (acroscopic one in lateral leaves) is at least 1.2–1.5 times wider than the other side. Purpus 6720 (US) is S. bernoullii, whereas duplicates of the same number at BM, M, NY, and UC are S. porphyrospora. Presumably, both species were growing together and were collected as part of a single gathering.

Selaginella bigelovii is closely related to S. rupincola, but their ranges do not overlap.

10. SELAGINELLA BIGELOVII

Selaginella breedlovei Valdespino, sp. nov. Type. Mexico. Chiapas: Huixtla, 6–8 km NE of Huixtla along road to Motozintla, Breedlove 28528 (holotype DS!). Figs. 285G, H, J–M.

Selaginella bigelovii Underw., Bull. Torrey Bot. Club 25: 130. 1898. Figs. 275A–D. Bryodesma bigelovii (Underw.) Soja´k, Preslia 64: 154. 1992. Type. U.S.A. California: Fort Smith to Rı´o Grande, Bigelow s.n. (NY!; isotypes GH, NY!).

Selected Specimens Examined. BCN (Moran 20495, UC, 24534, ENCB; Raven et al. 12284, UC; Wiggins & Wiggins 14843, MEXU). BCS (Brandegee 672, NY, UC; Carter & Ernst 494, MEXU; Carter et al. 2050, ARIZ, MEXU, UC; Wiggins et al. 494, UC).

11. SELAGINELLA BREEDLOVEI

Selaginella orizabensi affinis, a qua imprimis differt foliis medianis lanceolatis, basibus obliquis, foliis lateralibus apicibus acuminatus, breviciliatis secus 1⁄3 basium. (Dedicated to Dennis Breedlove, pre-eminent collector of Chiapan plants.)

Stems forming clumps, radially symmetric, underground (rhizomatous) and aerial, not readily fragmenting, irregularly branched, not articulate, not flagelliform, not stoloniferous; rhizomatous and aerial stems often with one branch arrested, budlike, tips straight; aerial stems erect or occasionally ascending; rhizophores restricted to rhizomes and proximal 1⁄3 of aerial stems, 0.3–0.4 mm diam.; leaves of one kind (isophyllous) throughout; rhizomatous stem leaves persistent, tightly appressed, scale-like; aerial stem leaves appressed, ascending, green, linear-lanceolate to narrowly lanceolate, 2.2–3.8 ⫻ 0.3–0.4 (–0.8) mm, bases abruptly adnate, cordate to almost peltate, pubescent or sometimes glabrous, margins short-ciliate at leaf bases, denticulate distally, cilia 0.02–0.08 mm long, white to transparent or greenish, leaf apices aristate, aristae hyaline to whitish, 0.2–0.8 mm; strobili quadrangular, (0.4–)1–1.5 cm; sporophylls monomorphic; megasporangia in two ventral rows; microsporangia in two dorsal rows; megaspores lemon-yellow, reticulate or rugose to smooth (when old), 300–340 ␮m diam.; microspores orange.

Stems creeping, stramineous distally, reddish proximally, ca. 17 cm long, 0.3–0.4 mm diam., not articulate, not flagelliform, not stoloniferous, 2–3 times branched; rhizophores limited to basal third of plant, filiform, 0.1–0.2 mm diam.; leaves of two kinds (anisophyllous) throughout stem length, imbricate, lustrous, red proximally; lateral leaves ovate to ovate-oblong, 1.5– 2.3 ⫻ 0.7–1.2 mm, acroscopic bases rounded, strongly overlapping the stems, margins hyaline, denticulate, short-ciliate on acroscopic base, apices acuminate, upper surfaces hispid, without idioblasts, lower surfaces glabrous, with or without obscure idioblasts; median leaves lanceolate, 0.7–2.1 ⫻ 0.3–0.7 mm, margins hyaline, the inner margins short-ciliate proximally, denticulate distally, outer margins entire or denticulate along distal 1⁄4, apices long-aristate, both surfaces glabrous, without idioblasts; axillary leaves similar to lateral leaves or more ovate-lanceolate, bases rounded, 1.4 ⫻ 0.7 mm; strobili quadrangular to slightly dorsiventral, 1–5 mm long; sporophylls monomorphic; megasporangia few, on ventral side of strobili; microsporangia occupying most of strobili; megaspores light yellow, 200–250 ␮m diam.; microspores orange.

Distribution. Terrestrial or epipetric on exposed rock crevices, steep north-facing slopes, cliffs, boulders, sandstone or igneous rocks, serpentine, or gravelly soils; 200–2400 m. USA (Calif); Mexico.

Distribution. Terrestrial on riverbanks; 200 m; Mexico. Specimen Examined. Chis (Breedlove 22558, DS).

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SELAGINELLA

This species is characterized by having the stems reddish proximally and covered with imbricate, lustrous red leaves, acroscopic half of the lateral leaves almost twice as wide as the basiscopic half; and aristae of median leaves 1⁄3 to 1⁄2 as long as the lamina length. In addition, some stems of S. breedlovei seem to arise from red buds that develop at the tips of flagelliform branch apices. Selaginella breedlovei may be closely related to S. orizabensis, from which it can be distinguished by the characters given in the key. Specimens of S. breedlovei were treated by Smith (1981) as S. flagellata.

12. SELAGINELLA CARIOI Selaginella carioi Hieron. in Engler & Prantl, Nat. Pflanzenfam. 1(4): 688. 1901. Syntypes (cited in Hedwigia 41: 198. 1902). Guatemala. Zacapa: Rı´o Motagua inter Barbesco et Gualau, Bernoulli 955 (frag. B, BM, K, NY!, P); same locality, Bernoulli & Cario 154 (presumably B); Alta Verapaz: Rı´o Chisoy, Bernoulli & Cario 172 (P); same locality Bernoulli & Cario 1087 (presumably B). Figs. 284O–T. Stems creeping, stramineous, 10 cm long, 0.2–0.4 mm diam., not articulate, not flagelliform, not stoloniferous, 2–3 times branched; rhizophores borne throughout stem length, filiform, 0.1–0.2 mm diam.; leaves of two kinds (anisophyllous) throughout stem length, glabrous, without idioblasts, dark to bluish green above, yellow-green below; lateral leaves ovate-oblong, 1.2– 1.5 ⫻ 0.5–0.7 mm, acroscopic bases auriculate and strongly overlapping the stems, margins whitish to hyaline, long-ciliate proximally, denticulate distally, apices obtuse to broadly acute; median leaves ovate to suborbicular, 0.7–1.3 ⫻ 0.4–0.7 mm, glabrous, without idioblasts, bases subcordate to short-auriculate, margins whitish to hyaline, long-ciliate proximally to shortciliate distally, apices apiculate to short-aristate; axillary leaves ovate to obovate-oblong, bases subauriculate with long hairs, distally denticulate; strobili quadrangular, 2–5 cm long; sporophylls monomorphic; megasporangia on ventral side of strobilus; microsporangia occupying most of strobilus; megaspores yellow, rugulate to reticulate, 200–250 ␮m diam.; microspores orange.

Distribution. Terrestrial on dry slopes; 800–1000 m. Mexico; Guat.

Specimens Examined. Chis (Breedlove 36540, DS, NY, 37259, NY, TEX).

Selaginella carioi is characterized by its whitish to hyaline leaf margins, four or more elongate cells wide. This character is more obvious on median leaves, where there is a strong contrast between the whitish margins and the deep green laminae. This species is also characterized by having the bases of the axillary leaves, the inner margins of the median leaves, and the acroscopic bases of the lateral leaves with dense stiff hairs. Selaginella carioi is closely related to S. reflexa and S. schiedeana but can be distinguished by the characters given in the key. Some authors (Alston, 1955; Fraile in Davidse et al., 1995; Smith, 1981; Stolze, 1983) have synonymized S. carioi under S. reflexa. However, the characters stated above set them apart. Additionally, S. carioi is limited to Chiapas and Guatemala, whereas most specimens of S. reflexa are from the north-central part of Mexico. The three species mentioned above, S. microphylla (Kunth) Spring from South America, and a probable undescribed species from Guatemala seem to form a natural alliance. This species group is characterized by yellow megaspores; keeled, ciliate sporophylls, with keels denticulate; and axillary rhizophores. Of this group S. carioi, S. reflexa, and S. microphylla seem to be well adapted to dry areas, whereas S. schiedeana usually grows in more humid and wet conditions.

13. SELAGINELLA CARNEROSANA Selaginella carnerosana T. Reeves, Brittonia 32: 365. 1980. Type. Mexico. Coahuila: Carneros, Pringle s.n. (NY!, frag. GH; isotypes MSC!, UC!). Figs. 273A–E. Stems prostrate, forming dense mats, dorsiventral, 0.3–0.5 mm diam., not articulate, not flagelliform, not stoloniferous, 1–2 times branched; rhizophores borne throughout stem length, filiform, ca. 0.2 mm diam.; leaves of one kind (isophyllous) throughout stem length, grayish green, margins green, ciliate, cilia hyaline, apices long-aristate or broadly acute if aristae break off, aristae usually persistent, twisted; upperside leaves lanceolate, 0.7–2.3 ⫻ 0.3–0.5 mm, bases truncate, abruptly adnate to the stems, glabrous, aristae 0.2–0.5 mm long; underside leaves linearoblong to oblong-elliptic, 1.6–2.6 ⫻ 0.4–0.7 mm, bases abruptly adnate to decurrent, glabrous, aristae 0.5–0.7 mm long; strobili quadrangular, 3–6 mm; sporophylls monomorphic; megasporangia few, basal on ventral side; microsporangia in two dorsal rows and scattered on ventral rows; megaspores yellow, reticulate, 320–360 ␮m diam.; microspores tan-orange.

SELAGINELLA CINERASCENS

Distribution. Epipetric on faces of limestones; 300 m. Mexico. Known only from the type collection. Selaginella carnerosana is distinct by the long, persistent (at least near stem apices), twisted aristae of leaves and sporophylls.

14. SELAGINELLA CHIAPENSIS Selaginella chiapensis A. R. Sm., Amer. Fern J. 70: 25. 1980. Type. Mexico. Chiapas: Mpio. Ocozocoautla de Espinosa, 18– 20 km N of Ocozocoautla along road to Mal Paso, Breedlove 28159 (DS!; isotypes frag. F!, NY!, MO!, US!). Figs. 269H, J–M. Stems creeping, stramineous, ca. 20 cm long, 0.4–1.2 mm diam., not articulate, not flagelliform, not stoloniferous, 2–3 times branched; rhizophores borne throughout stem length, filiform to stout, 0.2–0.5 mm diam.; leaves of two kinds (anisophyllous) throughout stem length; lateral leaves ovate, 1.7–2.7 ⫻ 0.8–1.3 mm, bases rounded, margins slightly hyaline, long-ciliate proximally, denticulate to entire distally, apices obtuse to broadly acute, upper surfaces slightly corrugate, papillate, lower surfaces glabrous, with idioblasts; median leaves broadly ovate to orbicular, 0.9–1.5 ⫻ 0.6–1 mm, bases subcordate, margins hyaline to greenish, long-ciliate, apices acuminate, upper surfaces corrugate, papillate and with obscure idioblasts; axillary leaves ovatelanceolate, bases rounded, margins long-ciliate, entire to denticulate distally, apices obtuse to broadly acute; strobili quadrangular, 3–8 mm long; sporophylls monomorphic or slightly dimorphic; megasporangia few, basal in ventral rows; microsporangia in dorsal rows and distally in ventral rows; megaspores white, reticulate, 300–400 ␮m diam.; microspores tan-orange.

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Distribution. Terrestrial on steep slopes and roadsides and woods along streams in tropical montane forests; 100–820 m. Mexico. Specimens Examined. Chis (Carlson 2127, ENCB, US). Oax (Hallberg 1625b, NY, US, 1648, NY; Lawton TS-14b, NY; Mickel 5861, 6497, NY).

In this species, the megasporophylls tend to be slightly broader than the microsporophylls. Selaginella chiapensis belongs to a species group that includes, in Mexico, S. barnebyana, S. corrugis, S. flexuosa, S. guatemalensis, S. huehuetenangensis, and S. subrugosa, which are here informally termed the “S. flexuosa group.” They can be set apart by the characters given in the key. Additionally, S. chiapensis is separated from its close relative, S. guatemalensis, by its broadly ovate to orbicular (vs. ovate to ovate-lanceolate) median leaves and by its lateral leaves perpendicular to the stems (vs. ascending at 45–60⬚). Selaginella chiapensis differs further from S. barnebyana and S. huehuetenangensis by its acuminate (vs. long-aristate) median leaves. To the “S. flexuosa group” also belong S. idiospora Alston (Guatemala to Costa Rica), S. denudata (Willd.) Spring (Jamaica), S. krugii Hieron. (Puerto Rico), S. macrostachya (Spring) Spring (Brazil), and possibly S. mendocae Hieron. and S. contigua Baker (both from Brazil). This group is in need of revision to ascertain relationships and distributions of the species.

15. SELAGINELLA CINERASCENS Selaginella cinerascens A. A. Eaton, Fern Bull. 7: 33. 1899. Figs. 273L–O. Bryodesma cinerascens (A. A. Eaton) Soja´k, Preslia 64: 154. 1992. Type. U.S.A. California: National City (San Diego), Kimball s.n. (GH; isotype NY!).

Stems forming compact, cushion-like mats, creeping, not readily fragmenting, slightly dorsiventral, irregularly branched, without bud-like arrested branches, tips straight, main stems ascending, 0.3–0.4 mm diam., not articulate, not flagelliform, not stoloniferous, 1–2 times branched; rhizophores borne throughout stem length, 0.2–0.3 mm diam.; leaves of one kind (isophyllous) throughout stem length, loosely appressed, dull to grayish green, linear to linear-lanceolate (1–)2.5–3 ⫻ (0.3–)0.4–0.6 mm (leaves

566

SELAGINELLA

on secondary and tertiary branches smaller), bases rounded and adnate or cuneate and slightly decurrent, glabrous, seldom pubescent, margins denticulate to very shortly ciliate, teeth or cilia hyaline, scattered, acute to slightly acuminate; strobili quadrangular, 2–6 mm long; sporophylls monomorphic; megasporangia few in two ventral rows; microsporangia on two dorsal rows and scattered on ventral rows; megaspores yellow, reticulate, 320–340 ␮m diam.; microspores orange.

Distribution. Terrestrial in dry open places, rocky hilltops, clayey-sandy soil, or in shade under shrubs and trees; 0–350 m. USA; Mexico. Selected Specimens Examined. BCN (Broome et al. 1951, NY; Ferris 8474, DS, NY, US; Johnston 1304, CAS; Montgomery 83-38, NY; Moran 19269, ENCB, LL, UC, 27088, ARIZ, UC; Moran & Reveal 20283, MEXU, MSC; Wiggins 4213, ARIZ, DS, GH, NY, UC, US, 21553, ARIZ, DS, ENCB; Wiggins & Thomas 416, DS).

The light brown and grayish mats, short lateral branches, narrow stems, and short strobili distinguish S. cinerascens from all other species in the flora. Selaginella cinerascens is without close relatives in Mexico; R. Tyron (1955) related S. cinerascens to S. arsenei, and it also closely resembles S. nivea Alston from Madagascar.

16. SELAGINELLA CONVOLUTA Selaginella convoluta (Arn.) Spring in Martius, Fl. Bras. 1(2): 131. 1840. Figs. 279A–F. Lycopodium convolutum Arn., Mem. Wern. Nat. Hist. Soc. 5: 199. 1824, non P. Beauv.) Desv. ex Poir. (1814), nom. illeg., published only in synonymy. Type. Brazil. Rio de Janeiro: near Rio de Janeiro, Jameson s.n. (E). Selaginella longispicata Underw. ex Millsp., Publ. Field Columbian Mus. Bot. Ser. 1: 287. 1896. Type. Mexico. Yucata´n: Near Itzamal, Gaumer 825 (NY!; isotypes F!, NY).

Stems fasciculate from a rhizomatous base, rosette-forming or occassionally erect, green or sometimes reddish, 4–15 cm long, 0.7–1.2 mm diam., not articulate, not flagelliform, not stoloniferous, 2–3 times branched; rhizophores restricted to bases of stems, filiform, 0.2–0.5 mm diam.; leaves of two kinds (anisophyllous) throughout stem length, coriaceous; lateral leaves ascending, imbricate, broadly ovate, 2.5–3 ⫻ 1.5–2 mm, bases pel-

tate, truncate, short-ciliate, margins hyaline to greenish, with a submarginal brown band, short-ciliate to denticulate, apices acute to acuminate; median leaves imbricate, ovate to ovateoblong, 2.2–3 ⫻ 0.8–1.5 mm, bases peltate, truncate, denticulate to short-ciliate, margins greenish to hyaline, short-ciliate, denticulate distally, apices acute to acuminate; axillary leaves similar to lateral leaves or more lanceolate; strobili quadrangular, 5–25 mm long; sporophylls monomorphic; megasporangia mostly basal in two ventral rows; microsporangia in two dorsal rows and distal in ventral rows; megaspores and microspores not seen.

Distribution. Terrestrial or epipetric in dry, exposed conditions on roadsides and slopes; 0–100 m. Mexico; Guat, Hond; Cuba, Hisp; Braz, Bol, Parag, Arg. Specimens Examined. Yuc (Gaumer 1982, NY; Gaumer et al. s.n., ARIZ; Steere 2171, US; Valde´z 50, F, NY, US). Unverified, Doubtful, or Mistaken Reports. Camp (CebellosCambranis 136, UADY, cited by Palacios-Rios, 2002a, but not verified).

Among the rosette-forming selaginellas of Mexico, S. convoluta is distinct by its peltate lateral leaf bases with a single auricle and its occurrence only on the Yucatan Peninsula.

17. SELAGINELLA CORRUGIS Selaginella corrugis Mickel & Beitel, Mem. New York Bot. Gard. 46: 336. 1988. Type. Mexico. Oaxaca: Distrito Villa Alta, valley of the Yelagago River, about 20 mi NE of Villa Alta, Mickel 1043 (NY!; isotype MEXU!). Figs. 268N–S. Stems prostrate, stramineous to pale green, 4–10 cm long, ca. 0.3 mm diam., not articulate, not flagelliform, not stoloniferous, 2–3 times branched; rhizophores borne throughout stem length, filiform, ca. 0.1 mm diam.; leaves of two kinds (anisophyllous) throughout stem length; leaves verruculose above, with prominent midribs; lateral leaves broadly ovate, ca. 1 mm long, bases rounded, margins denticulate, apices obtuse; median leaves ovate, with a narrow white border, midribs prominent and raised, bases exauriculate, margins ciliate-denticulate, apices short-cuspidate; axillary leaves elliptical, exauriculate, margins ciliate-denticulate, apices obtuse; strobili quadrangular, 3–7 mm long; sporophylls

SELAGINELLA DELICATISSIMA

monomorphic and of two colors, dorsal rows green, ventral rows colorless, ascending; megasporangia several at base of ventral rows; microsporangia in dorsal two rows and distally in ventral rows; megaspores grayish white, with low reticulum, 210–260 ␮m diam.; microspores dull orange, with thin capitate papillae.

567

in ventral rows; megaspores white, with low narrow ridges, 190– 240 ␮m diam.; microspores yellow-orange.

Distribution. Terrestrial in wet montane forests; 750–1150 m. Mexico. Distribution. Terrestrial on rock cliffs in wet montane forests; 1150 m. Mexico; CR, Pan. Known in Mexico only from the type collection. Selaginella corrugis is distinct in its very small leaves with verruculose leaf surfaces, ciliate-denticulate margins, and cuspidate median leaves. Selaginella flexuosa has larger leaves with long aristae and white denticulate margins proximally. Selaginella subrugosa has intermediate-sized leaves with short aristae and white minutely denticulate margins. The complex needs further study.

18. SELAGINELLA CUNEATA Selaginella cuneata Mickel & Beitel, Mem. New York Bot. Gard. 48: 336. 1988. Type. Mexico. Oaxaca: Distrito Villa Alta, valley of the Yelagago River, ca. 20 mi NE of Villa Alta, Mickel 1086 (NY!; isotype MEXU!). Figs. 267P–T. Stems erect, stramineous to pale green, to 60 cm tall, to 2 mm diam., not articulate, not flagelliform, stoloniferous; rhizophores restricted to bases of stems, to 0.7 mm diam.; leaves uniform on main stems, anisophyllous otherwise, appressed below first branch, ovate, rounded at bases with finely denticulate margins, not curling at margins when dry, with conspicuous idioblasts only on median leaves; lateral leaves ovate on main stems, 3.5–4 mm long, narrowly ovate, on branchlets 1.5–2 mm long, bases rounded-subcuneate, entire to minutely denticulate, apices obtuse to acute; median leaves ovate, bases rounded to slightly oblique, margins entire or with a few minute ciliolate teeth, with two conspicuous idioblasts, acuminate to shortaristate; axillary leaves narrowly ovate to lanceolate, acute, exauriculate, bases cuneate to somewhat rounded, entire to minutely denticulate at bases; strobili quadrangular, 6–15 mm long; sporophylls monomorphic and of two colors, dorsal rows green, ventral rows colorless, ascending; megasporangia in basal 1⁄2 of ventral two rows; microsporangia in dorsal two rows and distally

Specimens Examined. Oax (Liebmann 2051, US frag., photo and drawing; Mickel 1044, NY).

Selaginella cuneata is very closely allied to S. oaxacana, differing most noticeably in the cuneate bases of the lateral and median leaves, the relative lack of idioblasts on lateral leaves, and the narrow strobili.

19. SELAGINELLA DELICATISSIMA Selaginella delicatissima Linden ex A. Braun, Index Sem. Hort. Bot. Berol. App. 13. 1857. Type. Cultivated plant at Berlin (ex herb. Braun) (B; isotype US!). Figs. 268A–F. Selaginella saccharata A. Braun, Ann. Sci. Nat. Bot., se´r 5, 3: 272. 1865. Syntypes. Mexico. Veracruz: Sierra de San Cristo´bal, 1853, Sartorius s.n. (B; frag. NY!, frag. US!), Schaffner s.n. (B).

Stems prostrate with slightly ascending branches to suberect, stramineous or commonly with reddish patches, to 35 cm long, 0.6–0.9 mm diam., not articulate, not flagelliform, not stoloniferous, 3–5 times branched, leafy branches concave on upper surface, their leaves curving upward at their tips; rhizophores throughout or along only the length of the main stems, lateral branches sometimes lacking rhizophores, filiform, 0.15–0.2 mm diam.; leaves of two kinds (anisophyllous) throughout stem length; lateral leaves obliquely oblong, 2–3 mm long, bases subcordate with the upper lobe overlapping the stem, margins longciliate, especially on the acroscopic edges, denticulate distally, curving upward (appearing concave from above), old leaves often orange or red, apices acute; median leaves lanceolate, uniauriculate to sub-biauriculate, margins short- to long-ciliate at bases, denticulate distally, apices acuminate to short-aristate; axillary leaves oblong, bases biauriculate with short, blunt, subparallel auricles, margins with a few long cilia, especially near the bases, apices acute to obtuse; strobili quadrangular, 6–9 mm long; sporophylls monomorphic; megasporangia several, in lower 1 ⁄2 of ventral two rows; microsporangia in two dorsal rows and

568

SELAGINELLA

distally in ventral rows; megaspores yellow, with low papillae merging and forming short ridges, 350–400 ␮m diam.; microspores orange, with many short, blunt papillae.

underside leaves lanceolate to lanceolate-elliptic (on central rows) or falcate (on marginal rows), 2–2.7 ⫻ 0.5–0.7 mm, bases decurrent, glabrous, margins ciliate, cilia transparent to opaque, spreading or ascending; strobili quadrangular, 3–8 mm long; sporophylls monomorphic; megasporangia scattered in ventral rows; microsporangia in two dorsal rows and scattered on ventral rows; megaspores pale yellow, reticulate, 230–340 ␮m diam.; microspores bright orange, 23–38 ␮m diam.

Distribution. Terrestrial on moist banks in exposed or lightly wooded areas, and rocky slopes; 900–2500 m. Mexico; Guat, Hond. Selected Specimens Examined. Chih (Be´nitez 1904, MEXU). Col (Jones 486, US). Dgo (Knobloch 2139, MSC). Gro (Rzedowski 48085, NY). Gto (Rzedowski 41570, NY). Hgo (Moore & Wood 4428, US). Jal (Pringle 2634, US). Me´x (Mickel 719, NY). Mich (King & Soderstrom 5036, MEXU, NY, TEX, US). Mor (Rose & Painter 7250bis, US). NL (Pringle 1978, ARIZ, NY, UC). Oax (Mickel 1596, NY, US). Qro (Rzedowski 48085, NY). SLP (Pringle 3296, NY, UC). Sin (Correll & Correll 28870, US). Tam (Bartlett 11046, ENCB, LL, NY, US). Ver (Purpus 2388, ARIZ, NY, US). Unverified, Doubtful, or Mistaken Reports. Ags (De la Cerda & Garcı´a 995, Garcı´a 2358, HUAA, cited by Siqueiros-Delgado & Gonza´lez-Adame, 2004, but not verified).

Distribution. Terrestrial or epipetric on rocky and sandy slopes, in open rock crevices or in soil; 300–2050 m. USA; Mexico. Selected Specimens Examined. BCN (Moran 15695, MEXU, UC, 24860, UC, 8524, DS, UC; Moran & Thorne 14429, ENCB, UC; Thorne & Charlton 60172, MEXU; Wiggins 9981, DS; Wiggins & Thomas 272, DS, ENCB). Coah (Shreve & Tinkham 9851, ARIZ). Son (Felger & Broyles 89-28, MEXU, TEX; Moser et al. 18135, ENCB).

The non-aristate to short-aristate median leaves that are short-ciliate at the bases to denticulate at the tips, yellow megaspores, and auriculate axillary leaves with long-ciliate margins distinguish S. delicatissima from S. chiapensis, S. flexuosa, S. guatemalensis, and S. martensii.

Selaginella eremophila is most closely related to S. carnerosana, S. landii, and S. parishii. In S. arizonica, S. basipilosa, S. carnerosana, S. eremophila, S. landii, S. parishii, and S. peruviana, the leaves are arranged in eight conspicuous rows: 3 underside (2 marginal, 1 central), 2 lateral, and 3 upperside (2 marginal, 1 central).

20. SELAGINELLA EREMOPHILA

21. SELAGINELLA EURYNOTA

Selaginella eremophila Maxon, Smithsonian Misc. Collect. 72: 3–5. 1920. Figs. 274N–T.

Selaginella eurynota A. Braun, Ann. Sci. Nat. Bot., se´r. 5, 3: 293. 1865. Type. Costa Rica. Aguacate: Hoffmann 907 (presumably B; isotype K). Figs. 277P–S.

Bryodesma eremophila (Maxon) Soja´k, Preslia 64: 154. 1992. Type. U.S.A. California: Riverside Co., Johnston 1047 (US!).

Stems prostrate, forming dense mats, not readily fragmenting, dorsiventral, tips upturned, 3–8 cm long, 0.4–0.6 mm diam., not articulate, not flagelliform, not stoloniferous, irregularly 2– 3 times branched; rhizophores borne throughout stem length, 0.2 mm diam.; leaves of one kind (isophyllous) throughout stem length, tightly appressed, green, apices with caducous, twisted, hyaline aristae, aristae Ⳳ 0.3 mm, deciduous, the leaf apices becoming acute to slightly mucronate in oldest branches; upperside leaves lanceolate, 1.3–1.4 ⫻ 0.3–0.4 mm, bases abruptly adnate, pubescent, margins ciliate, cilia transparent to opaque;

Stems suberect, greenish to stramineous, 20–35 cm long, ca. 1 mm diam., articulate, not flagelliform, not stoloniferous, 2–3 times branched; rhizophores in proximal 1⁄4 of stems, stout, straight, 0.4–0.5 mm diam.; leaves of two kinds (anisophyllous) throughout stem length; lateral leaves obliquely oblong, slightly to highly curled, 4–4.5 mm long near stem tips, 1–1.5 mm long near base of plants, bases truncate to subcordate, centrally attached, margins short-ciliate at bases, denticulate distally, apices acute; median leaves obliquely ovate-lanceolate, bases with a single peltate auricle, margins ciliate, apices long-acuminate; axillary leaves linear-oblong, auriculate with incurved auricles, ciliate

SELAGINELLA FINITIMA

in lower 1⁄2, denticulate distally; strobili quadrangular, 3–6 mm long; sporophylls monomorphic; megasporangia single at base of strobili; microsporangia occupying most of strobili; megaspores beige, reticulate with white, high ridges, 450–540 ␮m diam.; microspores beige.

569

teeth hyaline, apices acuminate to short-aristate, aristae greenish hyaline, 0.1–0.2 mm long; strobili frequently in pairs at branch tips, quadrangular, 0.5–2(–2.5) cm long; sporophylls monomorphic; megasporangia scattered on ventral rows; microsporangia in two dorsal rows and scattered on ventral rows; megaspores deep yellow, rugose-reticulate, with a prominent equatorial flange, the flange often appearing to have tuberculate to papillate projections, 300–360 ␮m diam. (–400 ␮m diam. including equatorial flange); microspores orange.

Distribution. Terrestrial on shaded roadsides and streambanks in wet tropical forests on Pacific slope; 50–900 m. Mexico; Guat, Nic, CR, Pan. Specimens Examined. Chis (Breedlove 28620, NY; Matuda 18036, US).

Selaginella eurynota is part of an incompletely resolved complex that includes S. schizobasis and S. sertata. Selaginella eurynota is characterized by having axillary leaves on the main stems with two incurved, ciliate auricles (vs. auricles divergent in S. schizobasis and absent on mature, non-stoloniferous stems of S. sertata) and lateral leaves with an acroscopic auricle clasping and directed downward along the stem (vs. only clasping, but not downward-directed in S. schizobasis and absent on mature, nonstoloniferous stems in S. sertata). Specimens cited by Mickel and Beitel (1988) from Oaxaca as S. eurynota have been assigned to S. sertata (which see for discussion), as have some collections from Guerrero and Jalisco. These specimens all show their main stems to have arisen from flagelliform branches, as is characteristic of S. sertata.

22. SELAGINELLA EXTENSA Selaginella extensa Underw., Bull. Torrey Bot. Club 25: 131.1898. Figs. 276A–E. Bryodesma extensa (Underw.) Soja´k, Preslia 64: 154. 1992. Type. Mexico. San Luis Potosı´: Las Canoas, Pringle 3900 (NY!; isotype DS!, MEXU, NY!, UC!, YU!).

Stems long-creeping, forming loose mats to 35 cm long, radially symmetric, 0.4–0.6 mm diam., not articulate, not flagelliform, not stoloniferous, 2–3 times branched; rhizophores throughout, 0.2–0.3 mm diam.; leaves of one kind (isophyllous) throughout stem length, lustrous, dark green or brownish to brownish with red spots when old, linear-lanceolate, 1.8–2.2 ⫻ 0.2–0.3 mm, bases truncate or cuneate, abruptly adnate to the stems, glabrous or pubescent, margins greenish, denticulate,

Distribution. Epipetric in limestone crevices and cliffs or epiphytic on tree trunks and branches; 700–2100 m. Mexico. Selected Specimens Examined. Hgo (Frye 2537, NY, UC). Jal (Dı´az 14295, UC). Oax (Lorence et al. 3415, MEXU). Qro (Rzedowski 48490, NY; Zamudio & Carranza 7184, NY). SLP (Mickel 566, NY; Rzedowski 9648, DS, NY). Tam (Breedlove 63686, CAS). Ver (Nee et al. 26819, NY; Purpus 6052, NY, UC). Unverified, Doubtful, or Mistaken Reports. Pue (Sharp 45375, US, cited by R. Tryon, 1955, but not verified).

Selaginella extensa is one of the most distinct spikemosses in Mexico. It is characterized by forming long, loose mats that frequently are epiphytic or on rocks, by lustrous linear-lanceolate leaves that frequently have reflexed apices, and by lanceolate sporophylls with long-acuminate, green apices. Megaspores of this species are remarkable by their discoid-ovoid shape and prominent equatorial flange, which is erose and appears to have tuberculate to mamillate projections. This species is most closely related to the western North American S. oregana D.C. Eaton, but it can be distinguished from that by its appressed (vs. mostly spreading) leaves with truncate to cuneate (vs. decurrent) bases.

23. SELAGINELLA FINITIMA Selaginella finitima Mickel & Beitel, Mem. New York Bot. Gard. 46: 338. 1988. Type. Mexico. Veracruz: Minantitla´n, 6.6 km N de la terracerı´a La Laguna-Rı´o Grande, sobre el camino nuevo a Ejido Belisario Domı´nguez, 14.7 km al E de La Laguna, 17⬚ 20' N, 94⬚ 22' W, Wendt & Villalobos 2543 (NY!; isotypes CAS!, MEXU!, MO!, NY!, TEX). Figs. 287A–F.

570

SELAGINELLA

Stems suberect to erect, stramineous or greenish, 6–32 cm long, 0.4–0.9 mm diam., not articulate, occasionally with flagelliform stem and branch apices, stoloniferous, 2–3 times branched; rhizophores restricted to proximal portion of stems or throughout stem length on flagelliform plants, 0.1–0.4 mm diam.; leaves of two kinds (anisophyllous) throughout stem length; lateral leaves broadly ovate to ovate-oblong, 1.5–3.2 ⫻ 1.2– 2.1 mm, bases rounded to subcordate, acroscopic bases strongly overlapping the stems, margins narrowly to widely hyaline or pale green, short- to long-ciliate proximally, denticulate distally, apices rounded, obtuse, or broadly acute, both surfaces glabrous, without idioblasts; median leaves broadly elliptical or suborbicular, 1.2–1.6 ⫻ 0.7–1 mm, bases with a single auricle, margins ciliate-denticulate or denticulate, apices gradually or abruptly long-aristate, aristae less than 1⁄2 the lamina length, 0.2–0.4 mm long, both surfaces glabrous, without idioblasts; axillary leaves similar to lateral leaves except more narrowly ovate and shortciliate to ciliate-denticulate proximally, bases rounded; strobili lax and flattened, dorsiventral, 2–10 mm long; sporophylls slightly dimorphic, the dorsal ones green, spreading, ventral ones hyaline to pale green, ascending; megasporangia generally occupying proximal 1⁄2 of two ventral rows or occasionally absent; microsporangia in two dorsal rows and along distal 1⁄2 of ventral rows, or strobili wholly microsporangiate; megaspores cream, with an equatorial flange, rugose to rugose-reticulate, 150–250 ␮m diam.; microspores orange.

Distribution. Terrestrial and epipetric on limestone rocks and stream banks in wet lowland forests; 100–800 m. Mexico. Selected Specimens Examined. Chis (Breedlove 35378, DS, 35379a, DS, ENCB, NY, 57707, CAS; Hoover 160, MO; Martı´nez et al. 25909, 26230, MEXU; Mu¨nch s.n., DS). Oax (Wendt et al. 2758-A, MEXU, NY). Tab (Magan˜a & Lorea 2166, ENCB). Ver (Dorantes 3970, XAL).

Selaginella finitima has a distribution similar to that of S. prolifera and S. lineolata and all three species have median leaves with an outer auricle that is frequently ciliate, suberect to erect habit, stoloniferous stem bases, and frequently ciliate lateral leaves. Selaginella finitima differs chiefly by lacking idioblasts on the upper surfaces of the leaves and sporophylls, and lateral leaves with rounded, obtuse, or broadly acute apices. Some specimens of S. finitima (e.g., Wendt & Villalobos 2543

(the type), and Breedlove 35379a) develop vegetative growth from the tips of strobili. In addition, Wendt & Villalobos 2543 possesses leafy rhizophores. This feature lends support to the view that rhizophores are modified shoots capable of developing leaves, but that usually bear roots at the tip.

24. SELAGINELLA FLAGELLATA Selaginella flagellata Spring, Bull. Acad. Roy. Sci. Bruxelles 10: 228. 1843. Lectotype (chosen here). “Guyana Gallica” [French Guiana]. Ad rivulus Oyapok superioris [upper Oyapok river], [Jun,] Leprieur s.n. (G!; isolectotypes B!, P!; probable isolectotypes GH!, P!, US!). Figs. 269T–Y. Selaginella ambigua A. Braun, Ann. Sci. Nat. Bot., se´r 5, 3: 286. 1865. Syntypes. Venezuela. Aragua: Colonia Tovar, Fendler [324] p.p. (B!, GH!, MO!, YU!); Sucre: Quebrada del Imposible, Cumana´, Moritz 221 (B!). Selaginella rhizophora Baker, J. Bot. 22: 244. 1884. Syntypes. Panama. Panama´: in sylvis prope Villa San Juan, Oct 1846, Seemann 29 (K!); in sylvis prope urbe Panama, Nov 1846, Seemann 32 (frag. B!, K!). Selaginella ambigua A. Braun var. fallax Hieron., Hedwigia 58: 317. 1917. Syntypes. Trinidad. Jenman 7 (3456) (B!, NY!); 1877–1878, Fendler 42 (B!, BM!, GH!, K!, M, NY!, S!, UC!, US!, YU!), 108 (M!).

Stems suberect to erect, stramineous, 6–15 cm long, 0.2–1 mm diam., not articulate, frequently with flagelliform stem apices, stoloniferous or not, 2–4 times branched; rhizophores restricted to proximal 1⁄3 or throughout stems on flagelliform plants, filiform to moderately stout, 0.1–0.5 mm diam.; leaves of two kinds (anisophyllous) throughout stem length; lateral leaves broadly ovate to ovate-oblong, 2–5 ⫻ 1–3 mm, bases rounded to subcordate, acroscopic bases strongly overlapping stems, basiscopic bases rounded, not overlapping stems, margins narrowly hyaline or pale green, denticulate, apices acute, both surfaces glabrous, without idioblasts; median leaves elliptic or broadly ovate-elliptic, 1–2 ⫻ 0.5–1.7 mm, bases subcordate, margins hyaline, denticulate, apices long-acuminate to long-aristate, aristae 1⁄2 the length to nearly as long as the lamina, 0.4–1 mm long, both surfaces glabrous, without idioblasts; axillary leaves similar to lateral leaves, bases rounded; strobili lax and flattened, dorsiventral, 0.2– 5 cm long; sporophylls dimorphic, the dorsal ones green, spreading, the ventral ones hyaline to pale green, ascending; megasporangia in two ventral rows or frequently absent; microsporangia in two dorsal rows, frequently in the distal region of ventral side of strobili, or strobili wholly microsporangiate; megaspores snowto bone-white, or cream, with an equatorial flange, rugosegranulate to granulate-reticulate, 230–240 ␮m diam.; microspores tan, the proximal face slightly rugose to smooth, the distal face baculate to clavate, 20–23 ␮m diam.

SELAGINELLA GUATEMALENSIS

571

250–350 ␮m diam.; microspores light orange, with slender, often capitate papillae.

Distribution. Terrestrial, epipetric, or epiphytic in lowland to montane forests, on open to shady roadside or stream banks, or forested slopes; 120–1000 m. Mexico; Guat, Hond, Nic, CR, Pan; Martinique?; Col, Ven, Trin, Tob, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol. Selected Specimens Examined. Chis (Breedlove 31585, 33958, DS, 37789, DS, ENCB 38478, DS; Fisher 35398, ARIZ, US; Martı´nez 13643, ENCB, MO; Mu¨nch 192, DS). Col (Gadow & Gadow s.n., BM). Oax (Mickel 4193, 5133A, NY, UC).

25. SELAGINELLA FLEXUOSA Selaginella flexuosa Spring, Flora 21: 197. 1838. Syntypes. Brazil. Swainson (K); Corcovado, near Rio de Janeiro, Langsdorff s.n. (BR, M!, P!), Raddi s.n. (BR); without precise locality: Desvaux s.n. (B); Sellow s.n. (B). Figs. 268G, H, J–M. Selaginella tarapotensis Baker, J. Bot. 21: 98. 1883. Type. Peru. Prope Tarapoto, Spruce 4625 (K!; isotypes LE!, US!, frag. NY!). Selaginella faucium Liebm. ex Baker, J. Bot. 21: 333. 1883. Syntypes. Mexico. Veracruz: Huitamolco, Liebmann s.n. [Pl. Mex. 2047] (K; isotype H, frag. US!), Hacienda de Jovo, Liebmann s.n. [Pl. Mex. 2048] (K; isotype H).

Stems prostrate, stramineous to pale green, ca. 0.6–0.8 mm diam., not articulate, not flagelliform, not stoloniferous; rhizophores borne throughout stem length, stout, straight, (0.3–)0.5– 0.8 mm diam.; leaves of two kinds (anisophyllous) throughout stem length; lateral leaves on main stem obliquely oblong to subovate, 2–3 mm long, bases rounded, margins minutely denticulate to entire, apices acute, green above, silvery below, sometimes with short idioblasts below, wrinkled on upper surface; median leaves ovate, with prominent midrib and white border, striate to weakly verruculose, exauriculate, margins denticulate to entire, apices long-aristate, aristae 0.8 mm long, 1⁄3 of lamina length; axillary leaves ovate to lanceolate, exauriculate, denticulate at bases; strobili quadrangular, 3–15 mm long; sporophylls monomorphic and of two colors, dorsal two rows green, ventral two rows hyaline, ascending; megasporangia in two ventral rows; microsporangia in two dorsal rows, smaller strobili often completely microsporangiate; megaspores white with faint markings,

Distribution. Terrestrial, frequent in wet montane forests; 200– 1900 m. Mexico; Guat, Nic, CR, Pan; Col, Ec, Peru, Bol. Selected Specimens Examined. Oax (McAlpin 1023, F; Mexia 9270a, UC, US; Mickel 946, MICH, US, 1063, MICH, NY, UC, US, 1445, MICH, NY, US; Rzedowski 30661, NY; Schultes & Reko 677, US). Pue (Garcı´a 72, ENCB, MICH, NY; Martin 4676, US). Ver (Lira 196, ENCB).

Selaginella corrugis, S. flexuosa, and S. subrugosa are characterized by their non-stoloniferous, non-flagellate, prostrate habit, stilt-like rhizophores, denticulate leaves, and narrow strobili. Selaginella flexuosa differs from S. corrugis by the long-aristate white-bordered median leaves (very short-cuspidate with narrow white border in S. corrugis), larger lateral leaves 2–3 mm long, each with a minutely denticulate apex and base (leaves 1–1.2 mm long, denticulate in S. corrugis). It has been collected in Oaxaca with S. corrugis and S. subrugosa.

26. SELAGINELLA GUATEMALENSIS Selaginella guatemalensis Baker, J. Bot. 21: 243. 1883. Lectotype (chosen by Alston 1955: 245). Guatemala. Alta Verapaz: Coba´n, Salvin & Godman s.n. (K!; isotype B!). Figs. 283F–H, J–L. Selaginella lehmannii Hieron., Hedwigia 43: 33. 1904. Type. Guatemala. Baja Verapaz: Prope Puxula´, Lehmann 1329 (B; isotype BM, K!, US!). Selaginella pansamalensis Hieron., Hedwigia 43: 35. 1904. Lectotype (chosen by Alston, 1955: 244). Guatemala. Alta Verapaz: Pansamala´, Tu¨rckheim s.n. [Donn. Sm. 653] (B; isolectotypes NY!, P, US!).

Stems mostly suberect, stramineous or pale green, to 18 cm tall and 1.5 mm diam., not articulate, not flagelliform, not stoloniferous, 2–4 times branched; rhizophores borne throughout stem length, stout, 0.4–0.5 mm diam.; leaves of two kinds (anisophyllous) throughout stem length; lateral leaves obliquely oblong, 2.5–3 mm long, upper surfaces corrugate, bases rounded acroscopically, truncate basiscopically, margins long-ciliate only at base on large acroscopic lobe, otherwise denticulate, apices obtuse or acute; median leaves ovate, bases scarcely auriculate, midribs prominent, margins short-ciliate, apices acuminate; ax-

572

SELAGINELLA

illary leaves ovate, long-ciliate in basal 1⁄4, bases rounded; strobili quadrangular, 5–14 mm long; sporophylls monomorphic and of two colors, dorsal rows green, ventral rows pale green; megasporangia basal or subbasal in ventral two rows; microsporangia in dorsal two rows and distal in ventral rows; megaspores white to beige, reticulate to smooth, 350–400 ␮m diam.; microspores pale orange.

Distribution. Terrestrial in wet montane forests and wet pine forests on Atlantic slopes; 200–1600 m. Mexico; Guat, Bel, Hond. Selected Specimens Examined. Chis (Breedlove 22329, MICH, NY, 32161, ENCB, MICH; Carlson 1728, US; Dressler 1634, MICH, NY, US; Hellwig & Whitaker 1360, NY; Rzedowski 33624, ENCB; Zamudio 1536, NY). Oax (Geith 95-MO52, NY; Hallberg 1418, NY, UC; Liebmann s.n. [Pl. Mex. 2060], frag. US!, drawing NY!).

Selaginella guatemalensis is distinguished from S. martensii by having acuminate median leaves, basally ciliate lateral leaves, long-ciliate axillary leaves, and more slender stems, whereas S. martensii has long-aristate median leaves, denticulate lateral and axillary leaves, and thick stems. Selaginella chiapensis, S. guatemalensis, S. huehuetenangensis, and S. idiospora are closely related and need more detailed study. Selaginella chiapensis has minutely denticulate lateral leaves that curl on the basiscopic margin, are long-ciliate on the proximal 1 ⁄2 of the acroscopic side and spread at about a 90⬚ angle from the stems. The median leaves are ovate with acuminate tips. The axillary leaves are ciliate 1⁄2–2⁄3 their length. Selaginella guatemalensis has minutely denticulate lateral leaves that are long-ciliate only on the proximal 1⁄4 of the acroscopic side, 2.5–3 mm long, and spread at about a 45⬚ angle. The median leaves are ovate with acuminate tips. The axillary leaves are ciliate only in the proximal 1⁄4. The lateral leaves of S. huehuetenangensis are long-ciliate proximally and ciliolate, denticulate, or entire distally. The whitemargined median leaves are long-aristate and ovate-lanceolate to broadly ovate, the axillary leaves are long-ciliate, and the sporophylls each have a distinct keel. Selaginella idiospora Alston, from Guatemala south to Costa Rica, has non-denticulate lateral leaves that curl on the longciliate acroscopic side and spread at about a 90⬚ angle. The median leaves are suborbicular with short aristae (1⁄4 length of

blade), and the axillary leaves are long-ciliate for 3⁄4 the length with the lateral margins often curling.

27. SELAGINELLA GYPSOPHILA Selaginella gypsophila A. R. Sm. & T. Reeves, Sida 10: 211. 1984. Type. Mexico. Nuevo Leo´n: Mpio. Galeana, S of Cerro Potosı´, ca. 2 mi N of Ejido Santo Domingo, and ca. 7 mi NE of San Roberto Junction (Hwys 57 & 58), 24⬚ 45'N, 100⬚ 12'W, Dorr et al. 2553 (UC!; isotypes GH, MEXU, MICH, MO, NY!, TEX, US). Figs. 279M–Q. Stems densely caespitose, rosette-forming, inrolled when dry, green above, yellowish or whitish green below, older leaves drying uniformly tan, branches strongly ascending, 3–5 cm long, 0.4–0.6 mm diam., not articulate, not flagelliform, not stoloniferous; rhizophores restricted to stem bases, ca. 0.4 mm diam.; leaves of two kinds (anisophyllous) throughout stem length; median and lateral leaves closely imbricate, oblong to oblonglanceolate, acutish or narrowly rounded at apex, margins green or inconspicuously hyaline, entire, sparingly denticulate, or long-ciliate proximally and denticulate distally; lateral leaves 1.4– 2 mm ⫻ 0.7–0.9 mm, ciliolate along acroscopic margins, mostly entire along basiscopic margins (except at bases), lower surfaces with broad whitish opaque margins to ca 0.3 mm wide and a green central band equal to or less than the width of the whitish margins, upper surfaces with green central band much broader than whitish margins; median leaves lanceolate, 1–1.3 mm ⫻ 0.4– 0.5 mm, bases pubescent, margins broad, whitish, 0.15–0.2 mm wide, color pattern reversed from that of lateral leaves, upper surfaces with narrow dark green median band, lower surfaces with broad green median band, apices acute to narrowly rounded; axillary leaves similar to lateral leaves but more nearly equilateral and ciliolate along both margins, bases with two minute, ciliolate lobes; strobili quadrangular, to 1.5 cm long; sporophylls monomorphic; megasporangia few, basal, in a vertical row in strobilus, or scattered in one or two vertical rows; microsporangia in vertical rows or scattered; megaspores yellow, with fine reticulate ridges, but at low magnification (30⫻) appearing smooth, ca. 300 ␮m diam.; microspores bright orangish red, capitate-papillate or tuberculate, shed in tetrads and singly.

SELAGINELLA HIRTIFOLIA

Distribution. Epipetric in rock crevices, dry gypsum hillsides; 2100 m. Mexico. Specimens Examined. NL (Dorr 2550, MEXU, TEX, UC; Hansen et al. 3859, MEXU; Hinton 18751, MEXU).

Selaginella gypsophila is distinguished from S. pilifera by its rounded (vs. long-aristate) leaf apices.

28. SELAGINELLA HARRISII Selaginella harrisii Underw. & Hieron. in Urban, Symb. Antill. 7: 162. 1912. Lectotype (chosen by Proctor, 1985: 37). Jamaica. Harris 7587 (BM!; isolectotypes IJ, K, NY!, US). Figs. 267G, H, 282A–F. Selaginella millspaughii Hieron., Hedwidia 58: 285. 1917. Type. Mexico. Yucata´n: Near Chichen Itza´, Millspaugh 1620 (B; isotypes BM, NY!).

Stems erect, green, 4–30 cm tall, ca. 1 mm diam., not articulate, not flagelliform, not stoloniferous, 3(–4) times branched; rhizophores only at base, 0.2–0.5 mm diam.; leaves uniform on main stems below first branches, anisophyllous otherwise, appressed on main stems below first branch, ovate, bases rounded to subauriculate, margins long-ciliate, bases often with branched hairs, apices acuminate; lateral leaves ovate, bases rounded, margins narrowly white-bordered, long-ciliate acroscopically, shortciliate to denticulate distally, old leaves green, apices sharply acute; median leaves ovate, bases with single short auricle, margins short-ciliate with narrow white borders, apices aristate; axillary leaves lanceolate, bases rounded, long-ciliate proximally, short-ciliate to denticulate distally; strobili quadrangular, 5–13 mm long; sporophylls monomorphic; megasporangia basal or subbasal in ventral rows; microsporangia in dorsal rows and distally in ventral rows; megaspores lemon-yellow to cream, 300–360 ␮m diam.; microspores bright orange.

573

Distribution. Terrestrial on dry soil in shady ravines, steep slopes, limestone ridges; 600–1550 m. Mexico; Bel, Hond; Jam. Selected Specimens Examined. Chis (Gilly & Herna´ndez X. 397, MSC; Riba et al. 1393, ENCB). Hgo (Chase 7433, ARIZ). Oax (Hallberg 1375, NY; Mickel 6153, NY). Qro (Ferna´ndez 3440, ENCB, NY, TEX). SLP (Hellwig & Whitaker 1220, MEXU; Pennell 17948, PH). Tab (Gilly & Herna´ndez 397, MSC). Ver (Copeland herb. 171, MEXU). Yuc (Montgomery 87-04, NY). Unverified, Doubtful, or Mistaken Reports. Camp (Palacios-Rios et al. 3609, XAL, cited by Palacios-Rios, 2002a., but not verified).

Selaginella harrisii differs from S. pallescens in having long stems that are not branched at the bases and have clasping leaves on most of the main stems. The only specimens of S. pallescens with those conditions occur in western Mexico (Jalisco and Nayarit) whereas S. harrisii is limited to eastern Mexico. Furthermore, S. harrisii has green older leaves (vs. tan), long ciliate stem leaves with slightly auriculate bases (vs. rounded), lateral leaves short-ciliate or denticulate with narrow white borders on acroscopic margins, entire to denticulate on basiscopic margins, and median leaves with white margins narrow to absent. It is related to S. polyptera and S. pulcherrima, which can be separated by characters given in the key. Another possible relative is S. microdendron Baker, known only from Cuba, which is a smaller frondose plant with biauriculate stem and axillary leaves with longciliate bases and short-ciliate margins.

29. SELAGINELLA HIRTIFOLIA Selaginella hirtifolia Valdespino, sp. nov. Type. Mexico. Chiapas: Solosuchiapa, 2–4 km below Ixhuata´n along road to Pichucalco, 1200 m, 20 Feb 1972, Breedlove 24183 (holotype NY!; isotypes DS!, ENCB!, F!, MICH!). Figs. 269A–G, 271V–Z, AA–BB. A S. revoluta foliis medianis acutis vel breviter acuminatis et foliis lateralibus acutis differt. (L., hirtus, hairy, folium, leaf, referring to the hairy leaf surfaces.)

Stems creeping, greenish or stramineous, 3–17 cm long, 0.4– 0.6 mm diam., not articulate, not flagelliform, not stoloniferous, 2–3 times branched; rhizophores throughout, 0.1–0.3 mm diam.;

574

SELAGINELLA

leaves of two kinds (anisophyllous) throughout stem length; lateral leaves broadly ovate-oblong, 1.5–2.5 ⫻ 0.8–1.3 mm, bases rounded to subcordate, margins whitish to hyaline or greenish, long-ciliate acroscopically, denticulate to entire distally and basiscopically, apices acute, upper surfaces corrugate and/or verruculose, hispid along the basiscopic half (particularly near margins), without idioblasts; median leaves ovate to broadly elliptic, 0.9–1.3 ⫻ 0.6–0.9 mm, bases subcordate, margins hyaline, longciliate to entire distally, apices acute to short-acuminate, upper surfaces corrugate and frequently verruculose, lower surfaces smooth, both surfaces without idioblasts; axillary leaves similar to lateral leaves, except margins ciliate in proximal 1⁄2 to 2⁄3, denticulate apically, bases rounded; strobili quadrangular, 3–10 mm long; sporophylls monomorphic; megasporangia few, ventral, at the base or center of the strobili and intermixed with microsporangia; microsporangia occupying most of the strobilus; megaspores snow-white, reticulate, with a low equatorial flange, 300–320 ␮m diam.; microspores orange, ca. 30 ␮m diam.

acuminate to aristate) median leaves and by its acute (vs. truncate to broadly acute) lateral leaf apices.

30. SELAGINELLA HOFFMANNII Selaginella hoffmannii Hieron., Hedwigia 41: 184. 1902. Syntypes. 18 specimens cited from Mexico (Veracruz), Guatemala, and Costa Rica, including Hoffmann 905 (B!), 905a (B!), 909 (B!), from Costa Rica. Figs. 267J–O, 269N–S. Stems erect, mostly 10–50 cm tall, 0.8–2 mm diam., much branched, frond-like, not articulate, usually not flagelliform (except some specimens from Chiapas), stolons red, extremely slender (2–3 mm diam.), main stems simple, red to pink, especially at base, 2–3 times branched; rhizophores only at stem bases or in proximal 5 cm of main stems, 0.2–0.3 mm diam.; leaves of two kinds (anisophyllous) throughout stem length; lateral leaves 1–2 mm long, to 3 mm on main stems, obliquely narrowly oblong, bases rounded to subcordate, margins denticulate and narrowly white-bordered (ciliate-denticulate on main stems), apices acute, often with idioblasts; median leaves ovate, slightly curved inward, bases exauriculate or with two very short, subequal auricles, margins denticulate, with narrow white borders, apices long-aristate; axillary leaves narrowly lanceolate, with rounded, exauriculate bases, margins denticulate and whitish; strobili quadrangular, 3–5 mm long; sporophylls monomorphic and of two colors, dorsal rows green, ventral rows nearly colorless; megasporangia in ventral two rows; microsporangia in dorsal two rows; megaspores white, finely rugose-reticulate, ca. 250 ␮m diam.; microspores orange, with slender projections.

Distribution. Terrestrial on wet banks in montane forests; 50– 2200 m. Mexico. Specimens Examined. Chis (Breedlove 56803, CAS, ENCB; Riba et al. 1383, ENCB). Oax (Herna´ndez 1160, ENCB, MEXU, MO, NY; Mickel 1477, NY 4761, NY, UC, 4790, NY). Tab (Cowan 3455, ENCB, MO, NY, UC).

Selaginella hirtifolia is characterized by the almost transparent to faintly green color of the ventral sporophylls. The presence of short hairs on the upper surfaces of the basiscopic half of the lateral leaves, particularly near the margins, distinguish S. hirtifolia from all other Mexican species except S. bernoullii, S. popayanensis, and S. porphyrospora. Selaginella hirtifolia is distinguished from these last three species by its ciliate (vs. denticulate) lateral and median leaves and sporophylls and its white (not orange) megaspores. Specimens of S. hirtifolia were misinterpreted by Alston (1955), Smith (1981), Mickel and Beitel (1988), and Fraile (in Davidse et al., 1995) as S. lindenii. Differences between these two taxa are discussed under S. lindenii. Selaginella hirtifolia is a member of the S. revoluta group. It differs from S. revoluta, the only other Central American member of this group, by its acute to short-acuminate (vs. long-

Distribution. Terrestrial and on rocks along streams in lowland woods and lower montane rain forests; 0–1950 m. Mexico; Bel, Guat, Hond, Nic, CR. Selected Specimens Examined. Chis (Purpus 7222, UC). Col (Orcutt 4640, DS, MEXU, NY p.p., US). Gro (Mexia 8860, ARIZ, UC, US). Jal (McVaugh 15943, US). Me´x (Matuda 27194, UC, US). Mich (Hinton 15223, LL, UC, US). Nay (Breedlove 44315, CAS). Oax (Hallberg 1275, US; Mickel 5201, UC, US). Sin (Sanders 21036, UC). Ver (Sharp 45797, US). Unverified, Doubtful, or Mistaken Reports. Qro (Rzedowski 45244a, QMEX, and Gutie´rrez 70, ENCB, cited by Arreguı´n et al., 2001, but not verified).

SELAGINELLA ILLECEBROSA

The erect stems with reddish bases and red stolons, and denticulate lateral, median, and sporophyllous leaves distinguish this taxon. As pointed out by Smith (1981), several specimens of S. hoffmannii in Chiapas have flagelliform lateral branches, similar to those of S. flagellata and S. tenella.

31. SELAGINELLA HUEHUETENANGENSIS Selaginella huehuetenangensis Hieron., Hedwigia 43: 32. 1904. Lectotype (selected by Reed, Mem. Soc. Brot.18: 128. 1965–66). Guatemala. Huehuetenango: “Inter Santa Cruz Almor ex Ixca´n,” Bernoulli & Cario 178 (presumably B). Figs. 283A–E. Stems creeping, stramineous, to 35 cm long, 0.5–0.8 mm diam., not articulate, not flagelliform, not stoloniferous, 2–4 times branched; rhizophores borne throughout stem length, 0.2– 0.3 mm diam.; leaves of two kinds (anisophyllous) throughout stem length; lateral leaves ovate-oblong, 1.7–2.2 ⫻ 0.8–1.2 mm, bases rounded, strongly overlapping stems acroscopically, margins hyaline, long-ciliate proximally, denticulate or entire distally, apices acute, upper surfaces corrugate with papilla-like projections, without idioblasts, lower surfaces glabrous, with obscure idioblasts; median leaves ovate-lanceolate or broadly ovate, 0.8–1.1 ⫻ 0.5–0.7 mm, bases subcordate, outer bases with ciliate auricles, margins broadly hyaline, long-ciliate, apices longaristate, aristae 0.3–0.6 mm long, both surfaces glabrous, upper surfaces corrugate with papilla-like projections, without idioblasts; axillary leaves ovate-lanceolate, bases rounded, margins long-ciliate, apices acute; strobili quadrangular, 0.3–2 mm long; sporophylls monomorphic; megasporangia few at base of ventral rows, intermixed with the microsporangia; microsporangia occupying most of strobilus; megaspores white, rugose, without an equatorial flange, 250–300 ␮m; microspores orange.

Distribution. Terrestrial on talus slopes or riverbanks of tropical rain forests; 160–400 m. Mexico; Guat, Bel, Hond, Nic, CR, Pan. Selected Specimens Examined. Chis (Breedlove 33161, DS, MICH, NY, 49201, 49985, ENCB; Martı´nez 6939, ENCB, NY, 7026, CAS, ENCB, 8147, 14884, NY; Matuda 2782, US, 3702, F, MICH, US; Sohns 1612, NY, US ).

575

Selaginella huehuetenangensis is part of a species group that also includes S. corrugis, S. flexuosa, S. guatemalensis, S. subrugosa, S. barnebyana, and S. idiospora. Selaginella denudata (Willd.) Spring and S. krugii Hieron., from the West Indies, also belong to this group, which is in need of revision. In Mexico S. huehuetenangensis seems to be restricted to low elevations. It can be distinguished chiefly by its long-aristate median leaves, with the aristae hyaline, 1⁄2 or more the length of the lamina, and by broadly hyaline leaf margins. Because of the long-aristate median leaves, S. huehuetenangensis can also be confused with S. barnebyana (which see for comparison).

32. SELAGINELLA ILLECEBROSA Selaginella illecebrosa Alston, Bull. Brit. Mus. (Nat. Hist.), Bot. 1: 239. 1955. Type. Guatemala. Izabal: stream tributary to Rı´o Frio, Steyermark 41579 (BM!; isotypes F!, US!). Figs. 272A–G. Selaginella wendtii Mickel & Beitel, Mem. New York Bot. Gard. 46: 355. 1988. Type. Mexico. Oaxaca: Distrito Juchita´n, Sta. Marı´a Chimalapa, Congregacio´n Nicolas Bravo, Ⳳ 3–5 km S of Pob. de Rı´o Alegre, Arroyo Chocolin, cerca del Rancho Rutt, Wendt, Ishiki I. & Vera C. 4749 (NY!; isotypes CAS!, MEXU!).

Stems erect, stramineous to pale green, 40–60 cm tall, to 3 mm diam., not articulate, not flagelliform, stoloniferous, (2–)3 times branched; rhizophores restricted to stem bases, to 1 mm diam.; leaves uniform on main stems below first branches, anisophyllous otherwise, glabrous, lacking idioblasts, appressed on stem below first branch, ovate, auriculate at bases, margins finely denticulate except for short- to long-ciliate bases; lateral leaves ovate, 4–6 mm long on main stems, obliquely oblong, 2.5–3 mm long on branchlets, bases rounded, short-ciliate proximally, entire distally, apices acute; median leaves ovate, bases cuneate, margins entire except for few long cilia at base of basiscopic margins, apices long-aristate; axillary leaves narrowly ovate to lanceolate, exauriculate, long-ciliate at bases, entire distally, apices acute; strobili quadrandular, 3–8 mm long; sporophylls monomorphic and of two colors, dorsal rows green, ventral rows colorless; megasporangia in ventral rows; microsporangia in dorsal rows; megaspores cream-white, with low, narrow ridges, 210–260 ␮m diam.; microspores orange-yellow.

576

SELAGINELLA

Distribution. Terrestrial in lowland wet forests; ca. 150 m. Mexico; Guat, Hond.

Distribution. Terrestrial or epipetric on dry, rocky, scrubby slopes; 1000–3700 m. Mexico.

Specimens Examined. Oax (only the type collection of S. wendtii). Ver (Wendt et al. 2523, ENCB, LL).

Specimens Examined. Jal (Bravo & Flores 30, IBUG; Barnes & Land 153, F; McVaugh 25825, CAS, MEXU, MICH, NY, UC; Pringle 10823, ARIZ, CAS, MICH, MSC). Me´x (Kenoyer s.n., ARIZ; Tejero-Dı´ez 3066, IZTA). Nay (Norris & Taranto 14156, ENCB, MICH, 14712, MICH).

Resembling S. oaxacana and S. cuneata in general habit, S. illecebrosa differs in leaves that are ciliate at their bases, lack idioblasts, and are silvery beneath, and in long-aristate median leaves. The orange-yellow microspores, exauriculate and entiremargined median leaves (except for a few long cilia), and appressed lateral stem leaves of S. illecebrosa distinguish it from S. mosorongensis. Selaginella mickelii differs in having leaves ciliate along their entire margin, ciliate sporophylls, and stems often flagelliform.

33. SELAGINELLA LANDII Selaginella landii Greenm. & N. Pfeiff., Ann. Missouri Bot. Gard. 5: 205. 1918. Type. Mexico. Jalisco: San Esteban Mts., 32 km from Guadalajara, Barnes & Land 2024 (MO!, frag. US!; isotype GH). Figs. 274A–F. Stems prostrate, branches ca. 5 mm apart, strongly dorsiventral, not articulate, not flagelliform, not stoloniferous; rhizophores borne throughout stem length; leaves of one kind (isophyllous) throughout stem length, spreading and bright green above, appressed and buff below, linear-lanceolate, apices acuminate on upperside leaves, underside leaves longer than upperside leaves, acute to short-aristate, leaf bases pubescent, margins short-ciliate; strobili quadrangular, ca. 4–6 mm long; sporophylls monomorphic; megasporangia in two ventral rows; microsporangia in two dorsal rows; megaspores yellow-orange, rugose, 340– 380 ␮m diam.; microspores orange, granulose, some in tetrads.

Unverified, Doubtful, or Mistaken Reports. Mor (Matuda 26352, MO, US, cited by R. Tryon, 1955, but not verified). Pue (Kenoyer 25, US, cited by R. Tryon, 1955, but not verified).

This species is closely related to S. basipilosa (see for discussion) and to S. parishii, from which it can be separated by the characters given in the key. It may be confused with S. wrightii, which has non-aristate underside leaves and acuminate sporophylls.

34. SELAGINELLA LEPIDOPHYLLA Selaginella lepidophylla (Hook. & Grev.) Spring in Martius, Fl. Bras. 1(2): 126. 1840. Figs. 266A–C. Lycopodium lepidophyllum Hook. & Grev., Icon. Filic. 2: t. 162. 1830. Type. Mexico. Nayarit: Tepic, near San Blas, Dundas s.n. (K!).

Plants in tight rosettes, flat when moist, curled upward when dry; stems not articulate, not flagelliform, not stoloniferous, 2–3 times branched, pubescent, green, mostly 8–10 cm long, ca. 1 mm diam.; rhizophores borne only at stem bases, 0.2–0.5 mm diam.; leaves of two kinds (anisophyllous) throughout, but lateral and median leaves nearly alike, broadly ovate, subcordate at bases, tightly imbricate, ca. 1 mm long and broad, pubescent at bases, broadly white-bordered, the margins entire to irregularly incised to short-ciliate, old lateral leaves often dark reddish to red-brown, apices short-cuspidate; lateral leaves pubescent particularly at basiscopic base, apices acute but becoming roundish with age; median leaf bases pubescent; strobili quadrangular, ca. 6 mm long; sporophylls monomorphic; megasporangia in two ventral rows; microsporangia in two dorsal rows; megaspores yellow, with high irregular ridges, 245–413 ␮m diam.; microspores orange, often shed in tetrads.

SELAGINELLA LINEOLATA

577

exauriculate, lanceolate; strobili quadrangular, 1–2.5 mm long; sporophylls monomorphic; megasporangia basal in ventral rows; microsporangia in dorsal rows and distal in ventral rows; spores immature.

Distribution. Terrestrial or epipetric among rocks in dry areas, dry rocky slopes, and exposed grassy hills; 100–2200 m. USA; Mexico. Selected Specimens Examined. BCN (Brandegee s.n., UC). BCS (Thomas 7859, DS). Chih (Pringle 887, NY). Coah (Wynd & Mueller 45, NY). Col (Alana & Cook 1691, UC). Dgo (Torke et al. 154 NY). Gro (Viveros & Casas 373, MEXU). Jal (McVaugh 17371, NY). Me´x (Mickel 721, NY). Nay (Jones 23493, CAS, NY, UC). NL (Dorr et al. 2602, UC). Oax (Mickel 3843, NY). Pue (Purpus 3155, NY). Qro (Rose & Rose 11198, NY). SLP (Rzedowski s.n., NY). Sin (Gentry & Arguelles 18367, NY). Son (Gentry 11372, UC). Tam (Rozynski 42, NY, UC). Zac (Lloyd & Kirkwood 21, UC, US). Unverified, Doubtful, or Mistaken Reports. Ags (Siqueiros 4241, HUAA, and eight other collections, cited by Siqueiros-Delgado & Gonza´lez-Adame, 2004, but not verified).

Selaginella lepidophylla is at the center of a species group that includes S. nothohybrida and S. novoleonensis, and S. ribae (which see for discussion). Selaginella convoluta is another rosetteforming species that curls inward when dry, but it has median leaves with non-whitened and short-ciliate margins.

35. SELAGINELLA LINDENII Selaginella lindenii Spring, Bull. Acad. Roy. Sci. Bruxelles 10: 142. 1843. Lectotype (chosen by Alston, 1955: 240). Mexico. Tabasco: Teapa, Oct, Linden s.n. (P!; isolectotypes BM!, LG!). Figs. 283O–T. Stems creeping, stramineous, 16.5 cm long, 0.3–0.4 mm diam., not articulate, not flagelliform, not stoloniferous, 2–3 times branched; rhizophores throughout, filiform, 0.1–0.2 mm diam.; leaves of two kinds (anisophyllous) throughout stem length; lateral leaves ovate-oblong, 1.5–2.2 ⫻ 0.7–1.2 mm, bases subcordate, strongly overlapping stems acroscopically, margins whitish to hyaline acroscopically, short- to long-ciliate proximally, denticulate to entire distally, apices obtuse, upper surfaces hispid throughout, without idioblasts, lower surfaces glabrous, striate, with or without obscure idioblasts; median leaves ovate to ovate-elliptic, 0.7–1.2 ⫻ 0.4–0.7 mm, bases with small outer auricle, margins hyaline (especially the outer margins), longciliate, apices acute to short-acuminate, both surfaces glabrous, without idioblasts; axillary leaves similar to lateral leaves or more

Distribution. Terrestrial on shaded, wet rocks; elevation unknown, undoubtedly low. Mexico. Known only from the type collection. In Mexico this species has been misinterpreted by Alston (1955), Smith (1981), Mickel and Beitel (1988), and Fraile (in Davidse et al., 1995), who all considered S. lindenii to include specimens here newly named S. hirtifolia. Selaginella lindenii can be easily distinguished from the latter by the base of the median leaves with (vs. without) outer auricles, lateral leaves with obtuse (vs. acute) apices, and leaves dark to bluish green (vs. light to yellow-green) above. Because of similarities in general habit and leaf shape, S. lindenii may be confused with S. arsiclada, which lacks an outer auricle on the median leaves and hairs on the upper surface of the lateral leaves. Selaginella lindenii seems to be more closely related to the South American species S. jungermannioides (Gaudich.) Spring and relatives (S. homaliae A. Braun, S. schultesii Alston ex Crabbe & Jermy, S. applanata A. Braun, and S. truncata H. Karst. ex A. Braun). It differs from them by the lateral leaves being hispid above.

36. SELAGINELLA LINEOLATA Selaginella lineolata Mickel & Beitel, Mem. New York Bot. Gard. 46: 343. 1988. Type. Mexico. Oaxaca: Putla, 24 km N of Putla, Mickel 4014 (NY!; isotype NY!). Figs. 286G, H, J–M. Stems suberect to erect, stramineous, 4.5–33 cm long, 0.2–0.9 mm diam., not articulate, with flagelliform branch and stem apices, stoloniferous, 2–3 times branched; rhizophores restricted to proximal 1⁄4 of stems or throughout stems on flagelliform plants, 0.1–0.3 mm diam.; leaves of two kinds (anisophyllous) throughout stem length; lateral leaves broadly ovate or oblong, 1.6–2.9 ⫻ 0.9–1.7 mm, bases rounded to subcordate, overlapping stems acroscopically, basiscopic bases auriculate to rounded, with the auricle overlapping the dorsal side of stems, margins nar-

578

SELAGINELLA

rowly to widely hyaline or pale green, long-ciliate to denticulate proximally, denticulate or entire distally, apices acute, surfaces glabrous, with idioblasts; median leaves broadly ovate-elliptic to ovate-oblong, 0.8–1.8 ⫻ 0.5–1 mm, bases asymmetrically biauriculate, with inner bases rounded or auriculate, frequently ciliate, ciliate-denticulate, or entire, outer bases with a well developed and long-ciliate or entire auricle, margins hyaline, ciliatedenticulate, apices gradually long-acuminate to long-aristate, aristae 1⁄2 or less the lamina length, 0.1–0.5 mm long, both surfaces glabrous, with idioblasts above; axillary leaves similar to lateral leaves, exauriculate; strobili lax and flattened, dorsiventral, 2–7 mm long; sporophylls dimorphic, the dorsal ones green, spreading, the ventral ones hyaline to pale green, ascending; megasporangia generally in two ventral rows or at the base of ventral rows, or infrequently absent; microsporangia in two dorsal rows, frequently also at the distal part of ventral side of strobili, or strobili wholly microsporangiate; megaspores bone-white, with a slightly to moderately developed equatorial flange, striatereticulate, 275–340 ␮m diam.; microspores reddish orange.

Distribution. Terrestrial or epipetric on shaded slopes, limestone outcrops, sandy soils, or banks of waterways; 900–2500 m. Mexico; Guat. Selected Specimens Examined. Chis (Breedlove 48159, CAS p.p., ENCB p.p.). Col (McVaugh 16097, CAS, MICH, US). Gro (Rzedowski & McVaugh 26, MICH). Hgo (Sharp 45865, MEXU p.p., US). Jal (Pringle 2593a, BR, E, GH, M, NY, P, US). Me´x (Hinton et al. 8212, DS, GH, MEXU p.p., NY p.p., U, US). Nay (McVaugh 18756, CAS, MICH, NY). Oax (Mickel 7331, NY). Qro (Rzedowski 46613, NY). Sin (Breedlove 35733, CAS, MICH, MO). Ver (Pringle 10806, ARIZ, CAS, F, GH, LL, MICH, MSC, TEX, UC, US).

Selaginella lineolata has glabrous lateral and median leaves with conspicuous idioblasts and white megaspores, which distinguish it from some other erect to suberect taxa with stoloniferous stems and flagelliform branches, e.g., S. bernoullii and S. porphyrospora. Selaginella lineolata is closely related to S. prolifera, S. finitima (which see for discussion), and S. tenella. It can be distinguished from S. finitima by having idioblasts on the upper surfaces of both median and lateral leaves, flagelliform stems and branch apices, and conspicuous idioblasts and stomata on upper surfaces of dorsal sporophylls. Selaginella lineolata differs from S. prolifera

in having flagelliform stem and branch apices, small, round auricles at the basiscopic bases of lateral leaves, and by short ciliatedenticulate projections on the acroscopic margins of the lateral leaves, which are less than 1⁄4 the width of the lamina (vs. more than 1⁄3 or even 1⁄2 in S. prolifera). Selaginella tenella differs from S. lineolata by the median leaves elliptic to ovate-elliptic, with bases oblique and lacking definite outer auricles, leaves generally with broad hyaline margins, and sporophylls slightly dimorphic. In addition, S. lineolata has basal stolons from which the upright stems develop, whereas these are absent in S. tenella. In Mexico, S. lineolata is more robust than S. tenella. The ciliate-denticulate leaves with conspicuous idioblasts of S. lineolata resemble leaves of S. stenophylla, which can be distinguished by suberect habit, stout rhizophores, absence of stolons, and more elongate lateral leaves that are 2–2.5 mm long and broadest at the bases. Differences in the lateral leaf size and shape are especially conspicuous when large stem leaves are compared. In S. lineolata, lateral leaves of the main stems are 2–2.5 mm long, whereas in S. stenophylla they are 3–3.5 mm long. Pringle 10806, from Veracruz, is slightly different from typical S. lineolata by its more compact habit, lateral leaves perpendicular to the stem for most of the stem length, and obscure idioblasts; in general the acroscopic margins of the lateral leaves are more obviously long-ciliate. Specimens from Chiapas were among the tallest observed. One of them, Breedlove 48159, CAS, is atypical by its almost quadrangular to weakly flattened strobili, dorsal sporophylls not distinctly larger than the ventral sporophylls, lateral leaves with few short to long hairs at the basalmost portion of the basiscopic margin, and median leaf bases with two rather well developed and long-ciliate auricles. In these last characters it resembles S. hoffmannii Hieron. Red stem color was reported in living plants of S. lineolata (Hellwig & Whitaker 1373, NY), but this color fades on drying. Reddish to pink stems are typical of S. hoffmannii. That specimen differs from S. hoffmannii by its conspicuous to obscure idioblasts on the upper surfaces of the lateral leaves (vs. idioblasts absent), stems stramineous (vs. usually reddish to pink or occasionally stramineous), and stoloniferous leaves non-green or hyaline (vs. red-green). In general the median and lateral leaves of this collection fall within the morphological range of S. lineolata.

37. SELAGINELLA MACRATHERA Selaginella macrathera Weath., J. Arnold Arbor. 24: 326. 1943. Type. Mexico. Chihuahua: Sierra del Virulento, 2–3 miles E of Rancho Virulento, ca. 65 miles S of Ojinaga, Johnston 8067 (GH; isotypes CAS!, MO!, UC!). Figs. 276J–L. Stems prostrate or creeping, forming loose mats, radially symmetric, 0.3–0.5 mm diam., not articulate, not flagelliform, not stoloniferous, 1–2 times branched; rhizophores borne throughout stem length, filiform, ca. 0.2 mm diam.; leaves of one kind (isophyllous) throughout stem length, grayish green, linear to linearlanceolate, 2.7–5 ⫻ 0.3–0.5 mm, bases truncate, abruptly

SELAGINELLA MARTENSII

adnate to the stems, glabrous, margins green, short-ciliate proximally, denticulate distally, cilia and teeth hyaline; apices longaristate, aristae hyaline, 1.4–2.6 mm (as long as lamina length or more); strobili quadrangular, 0.5–1.1 cm long; sporophylls monomorphic; megasporangia few, scattered in ventral rows; microsporangia in two dorsal rows and scattered in ventral rows; megaspores deep yellow, reticulate or reticulate on proximal face and rugose to smooth on distal face, 300–360 ␮m diam.; microspores orange, 40–50 ␮m diam.

579

500–600 ␮m diam.; microspores buff to tan, echinulate, 24–26 ␮m diam.

Distribution. Terrestrial on exposed or disturbed pasture lands, thorn forests; 0–700 m. Mexico; Ven, Braz, Bol, Parag, Arg, Uru.

Distribution. Terrestrial or epipetric on cliffs; 1100–1400 m. Mexico. Specimens Examined. Chih (Martin et al. s.n., ARIZ). Son (Sanders et al. 13502, UC).

This species is characterized by having the longest aristae, as long as the laminae or longer, of all isophyllous species treated here.

38. SELAGINELLA MARGINATA Selaginella marginata (Humb. & Bonpl. ex Willd.) Spring, Flora 21: 194. 1838. Figs. 277T–X. Lycopodium marginatum Humb. & Bonpl. ex Willd. Sp. Pl., ed. 4, 5(1): 41. 1810. Type. Venezuela. Apure: Mouth of Rı´o Meta, Humboldt s.n. (B).

Stems long-creeping, stramineous, 40 or more cm long, 0.7–1 mm diam., articulate, not flagelliform, not stoloniferous, 3–4 times branched; rhizophores borne throughout stem length, 0.3– 0.5 mm diam.; leaves of two kinds (anisophyllous) throughout stem length; lateral leaves ovate-oblong to oblong, 1–3.5 ⫻ 0.7– 1.5 mm, bases peltate, with a long downward-pointing ciliate auricle, margins hyaline to greenish, short- to long-ciliate proximally, denticulate distally, apices acute, upper surfaces glabrous, without idioblasts; median leaves broadly ovate to ovate-oblong, 0.7–2.5 ⫻ 0.4–1.2 mm, bases peltate, with a long-decurrent ciliate auricle, margins hyaline (especially the outer margin) to whitish, denticulate, apices acuminate to short-aristate; axillary leaves similar to lateral leaves or more lanceolate; strobili quadrangular, 4–10 mm long; sporophylls monomorphic; megasporangia single at base of strobili; microsporangia occupying nearly the entire strobilus; megaspores white, reticulate with high muri,

Selected Specimens Examined. Dgo (Ortega 4465, US). Gro (Langlasse´ 371, MEXU, US; Verduzco 417, NY). Nay (Jones 23494, NY, UC; Rose 3148, NY, US). Sin (Breedlove 35630, MEXU, MICH; Mexia 222, CAS, UC, US; Molseed 6, LL, MEXU, NY, UC; Pennell 20187, PH, US; Stork & Horton 8601, LL, UC, US).

Selaginella marginata is readily distinguished among articulate species by having the base of its lateral, median, and axillary leaves peltate, each with a long, ciliate auricle extending lengthwise along the stem.

39. SELAGINELLA MARTENSII Selaginella martensii Spring, Me´m. Acad. Roy. Sci. Belgique 24: 129. 1849. Lectotype. Gregory & Riba (1979: 16) cited Galeotti 6606-bis, LV (intending LG?), as “type” (i.e., lectotype). Mickel and Beitel (1988: 344) pointed out that of the four syntypes of S. martensii (Galeotti 6606, 6608, 6618, and Jurgensen s.n., no herbarium indicated), Alston (1955: 257) had previously selected Galeotti 6606 as lectotype of S. galeottii (⫽ S. stellata ). For this reason (and others, discussed in 1988), Mickel and Beitel lectotypified S. martensii with Galeotti 6618. This specimen is not at LG, said to house Spring’s personal herbarium, but has been seen at BR (3 sheets) and P (2 sheets), herbaria consulted by Spring. One of the BR specimens should be selected as “second-stage” lectotype. Figs. 270F–H, J, K. Selaginella solmsii Baker, Handb. Fern-Allies 56. 1887. Type. Guatemala. Costa Grande, Bernoulli & Cario 181 (K).

Stems suberect, branching from proximal 2⁄3, stramineous to pale green, 2–2.5 mm diam., not articulate, not flagelliform, not stoloniferous; rhizophores in proximal 1⁄3–2⁄3 of stems, stout, mostly 0.8–1.2 mm diam.; leaves of two kinds (anisophyllous) throughout stem length; lateral leaves obliquely oblong, 3.5–5 mm long, bases rounded, margins denticulate to entire, rarely with a few long cilia at bases, apices obtuse; median leaves ovate,

580

SELAGINELLA

midribs prominent, aristae 0.8–1 mm long, bases each with one basiscopic auricle, margins slightly white-bordered, long-ciliate to denticulate distally, apices long-aristate; axillary leaves ovate, bases cordate, exauriculate, with a few cilia at bases, otherwise denticulate to entire, apices obtuse; strobili quadrangular, 3–15 mm long; sporophylls monomorphic; megasporangia in two ventral rows; microsporangia in two dorsal rows; megaspores white, 200–300 ␮m diam.; microspores orange-tan, with low broad papillae coalescing into ridges, shed mostly in tetrads.

Distribution. Terrestrial in montane rain forests and cloud forests; 130–2300 m. Mexico; Guat, Hond, Sal, Nic, CR, Pan. Specimens Examined. Chis (Breedlove 33634, NY; Martı´nez et al. 19402, NY). Me´x (Schaffner 6, NY). Oax (Mickel 6151 *, 6352 *, NY, US; Torres & Martı´nez 5979, NY). Qro (Dı´az-Barriga 4890, QMEX; Dı´azBarriga & Carranza 6510, QMEX). SLP (Pringle 3293, NY p.p., US). Ver (Nee & Taylor 29467, NY; Ortega 244, MEXU, NY).

Selaginella martensii is distinguishable by its thick, straight rhizophores, long-aristate median leaves with ciliate (to denticulate) margins and basiscopic auricles, large lateral leaves 3.5–5 mm long with only a few short hairs at bases, and broad stems 2–2.5 mm wide. As pointed out by Mickel and Beitel (1988), some specimens of S. martensii from Oaxaca (two of which are cited here and marked by an asterisk) have long-ciliate median leaves with prominent idioblasts and lateral leaves with ciliate bases; they may represent a distinct taxon restricted to Oaxaca. Selaginella stenophylla is similar to S. martensii in being suberect with stout rhizophores, with ciliate-denticulate to shortciliate, long-aristate median leaves, and denticulate lateral axillary leaves. Selaginella stenophylla differs in having idioblasts on the median and lateral leaves, which are smaller (1.8–3 mm long), strongly winged, dimorphic sporophylls and cream-colored megaspores. See S. lineolata for comparison of it with S. stenophylla.

40. SELAGINELLA MICKELII Selaginella mickelii Valdespino, Brittonia 44: 316. 1992. Type. Mexico. Oaxaca: Dist. Tuxtepec: 4–9 km S of Valle Nacional on Rte. 175, Mickel 5871 (NY!; isotypes MEXU!, MICH!, MO!, NY!, PMA!, UC!, US!). Figs. 268T–Y.

Stems erect from a creeping or decumbent rhizomatous and stoloniferous base, light-green to stramineous, to 45 cm tall, 1– 2 mm diam. (on erect stems) or 0.2–0.9 mm diam. (on creeping stems), not articulate, 2–4 times branched, frond-like, stolons restricted to proximal part of stems, apices of both stem and branches often flagelliform, flagelliform tips usually longcreeping, touching ground and sending up erect frond-like branches; rhizophores restricted to bases of erect stems or throughout on creeping stems and flagelliform tips, 0.5–1 mm diam. (on erect stems) or 0.2–0.4 mm diam. (on creeping stems and flagelliform tips); leaves of two kinds (anisophyllous) throughout stem length; lateral leaves conspicuously corrugate or wrinkled, mostly dimorphic on creeping stems, strongly appressed and curling inward to first branch (or beyond), dimorphic on erect stems above first branch, sometimes curling inward, ovate, 2–3.5 ⫻ 1.2–2.1 mm (1⁄4 to 1⁄3 less so on branches and creeping stems), acroscopic bases rounded, strongly overlapping stems, basiscopic bases truncate, margins green, longciliate proximally, denticulate distally, apices acute to obtuse; median leaves on erect stems imbricate, appressed, ovate, 1.5–3.3 ⫻ 1–2.3 mm, bases cordate with unequal auricles, margins hyaline to obscurely so, long-ciliate throughout, or denticulate distally, apices acuminate to short-aristate; axillary leaves ovatelanceolate to lanceolate, bases cordate to rounded, usually auriculate, auricles ciliate, margins ciliate in proximal 3⁄4, distal 1⁄4 obscurely denticulate, apices acute to narrowly-obtuse; strobili quadrangular, 0.3–2 cm long; sporophylls monomorphic or ventral ones sometimes lighter green; megasporangia usually in two ventral rows; microsporangia usually in two dorsal rows; megaspores white, sometimes with a low equatorial flange, granulose to low-reticulate, 230–320 ␮m diam.; microspores light-orange, rugose-granular to baculate, 19–25 ␮m diam.

Distribution. Terrestrial and epipetric on rocky stream banks, steep slopes and ravines, limestone cliffs, and roadsides in lower montane wet forests; 0–1200 m. Mexico. Selected Specimens Examined. Chis (Breedlove 56801, ENCB, MICH, NY; Matuda 3691, LL, MICH, NY). Oax (Mexia 9190, ARIZ, LL, MICH, NY, UC; Mickel 7198, NY). Tab (Gilly & Herna´ndez 282, MICH, MSC, TEX; Martı´nez et al. 8540, NY; Rovirosa 615, NY, PH; Zamudio 207, NY). Ver (Martı´nez Caldero´n 1744, MEXU; Nee & Diggs 24744, F, MICH, MO, NY, TEX, XAL).

SELAGINELLA MIXTECA

581

Specimens of S. mickelii have previously been referred to S. mollis A. Braun of Colombia and Venezuela, but S. mickelii is distinct by the flagelliform stem apices. Within Mexico S. mickelii mostly closely resembles S. illecebrosa, the former being distinct by the leaves on the main stems short-ciliate along margins (vs. ciliate at leaf bases), median and axillary leaves long-ciliate along margins (vs. entire except at leaf bases), and the stems often flagelliform (vs. non-flagelliform).

41. SELAGINELLA MINIMA Selaginella minima Spring, Bull. Acad. Roy. Sci. Bruxelles 10: 139. 1843. Lectotype (chosen here). French Guiana. Leprieur s.n. (LG!; isolectotype P!) Figs. 271P–U. Selaginella cruegeri Jenman, Gard. Chron. III, 22: 378. 1897 [“crugerii”]. Type. Trinidad. St. George Co.: Arouca [?], 15 Dec 1851, Herb. Trin. No. 194 (TRIN!; isotypes B!, US!). Selaginella tropidophora Mickel & Beitel, Mem. New York Bot. Gard. 46: 354. 1988. Type. Mexico. Oaxaca: Distrito Tuxtepec, 1–3 km S of Usila, Mickel 7392 (NY!). The collection data of the holotype were cited incorrectly by Mickel and Beitel (1988), being those of the paratype, Mickel 4204 (NY).

Stems ascending to suberect, stramineous to green, 1–9 cm long, 0.2–0.5(–0.8) mm diam., not articulate, not flagelliform, not stoloniferous, 1–3 times branched; rhizophores restricted to proximal 1⁄2 of stems, filiform, 0.1–0.2 mm diam.; leaves of two kinds (anisophyllous) throughout stem length, frequently with sparse short hairs or glabrescent, without idioblasts; lateral leaves ovate to broadly ovate, 1.5–1.9 ⫻ 0.8–1.1 mm, bases rounded, the acroscopic bases strongly overlapping stems, margins hyaline or greenish, acroscopically long-ciliate to denticulate distally, basiscopically entire to denticulate, apices acute; median leaves ovate, 1–1.3 ⫻ 0.45–0.7(–0.8) mm, inner bases truncate, outer bases oblique, margins conspicuously hyaline, denticulate, or the outer margins short-ciliate proximally and denticulate distally and the inner margins denticulate, apices acute to longacuminate; axillary leaves similar to lateral leaves, except for longciliate margins along the proximal 1⁄4–1⁄2, exauriculate; strobili lax, flattened, dorsiventral, 2–5(–12) mm long; sporophylls dimorphic, the dorsal ones green, spreading, the ventral ones pale green to hyaline, ascending to spreading; megasporangia in two ventral rows or sometimes mixed in one of these rows with microsporangia; microsporangia usually in two dorsal rows, sometimes mixed with megasporangia in a ventral row; megaspores snow-white, with a slightly developed equatorial flange, rugulate, reticulate, 270–350 ␮m diam.; microspores tan, verruculose with small points, 45–50 ␮m diam.

Distribution. Terrestrial on dry and open wooded slopes, on clay banks in savannas and on shaded stream banks; 300 m. Mexico; Hond, Nic, Pan; Ven, Trin, Fr Gui, Braz, Bol. Specimens Examined. Oax (Mickel 4204, NY, UC; Tryon & Tryon 7389, 7389a, GH).

Because of its small size, S. minima has been very often confused with S. simplex (Valdespino, 1993). See discussion under S. simplex) for differences.

42. SELAGINELLA MIXTECA Selaginella mixteca Mickel & Beitel, Mem. New York Bot. Gard. 46: 345. 1988. Type. Mexico. Oaxaca: Sola de Vega, 23–29 km S of Sola de Vega, 27–33 km N of bridge at Juchatengo, Mickel 6321 (NY!; isotype UC!). Figs. 271H, J–O. Stems suberect to erect, stramineous, 4.5–13 cm long, 0.3–0.8 mm diam., not articulate, not flagelliform, stoloniferous, 2–3 times branched; rhizophores restricted to stem bases, mostly on underground stolons, filiform, 0.2–0.3 mm diam.; leaves of two kinds (anisophyllous) throughout stem length; lateral leaves broadly ovate to ovate-oblong, 2.3–3 ⫻ 1.3–1.7 mm, acroscopic bases rounded, strongly overlapping the stems, the basiscopic bases truncate to rounded, margins pale green, long-ciliate acroscopically, entire to denticulate distally and basiscopically, apices acute; median leaves broadly ovate to ovate-lanceolate, 1.2– 1.9 ⫻ 0.7–1 mm, bases rounded or subcordate, margins hyaline, long-ciliate along proximal part of the stem, short-ciliate to denticulate above first branch, apices acute on leaves along proximal part of the stems, gradually long-acuminate to long-aristate on leaves above first branch, aristae half or less the lamina length, 0.1–0.5 mm long, both surfaces glabrous, without idioblasts; axillary leaves similar to lateral leaves or more oblong, bases exauriculate; strobili slightly lax and flattened, dorsiventral, 2–4 mm long; sporophylls weakly dimorphic, the dorsal ones green, spreading, the ventral ones pale green, ascending; megasporangia single at ventral base; microsporangia occupying most of the strobilus; megaspores pale yellow to lemon-yellow, striate-reticulate with close reticulum of low ridges, 320–360 ␮m diam.; microspores orange.

582

SELAGINELLA

dorsal rows or absent; megaspores beige or orange, reticulatestriate to corrugate-reticulate with moderately high ridges, 270– 367 ␮m diam.; microspores orange.

Distribution. Terrestrial or epipetric along waterway banks, roadcuts, or on hillsides with limestone outcrops; 1350–2100 m. Mexico. Specimens Examined. Oax (Hellwig & Whitaker 1269, 1272a, MEXU, NY, 1277, CAS, MICH, US; Mickel 6300, NY; Norris & Taranto 16526a, NY). Pue (Orcutt 1425, DS).

Because of its suberect to erect and stoloniferous habit, S. mixteca may be confused with S. prolifera, S. lineolata, S. finitima, and S. tuberosa. It differs from them chiefly by having longciliate to fimbriate, whitish or reddish scale-like leaves on the proximal and underground parts of the stems, and pale yellow to lemon-yellow megaspores.

43. SELAGINELLA MORITZIANA Selaginella moritziana Spring, Me´m. Acad. Roy. Sci. Belgique 24: 249. 1849. Type. Venezuela. Me´rida: Moritz 377 (B; isotypes BM!, G!, K!, LG!, P!). Figs. 287G, H, J–M. Stems creeping or ascending to suberect, stramineous to greenish, 8–22 cm long, 0.4–0.8 mm diam., not articulate, not flagelliform, stoloniferous or not, 2–3 times branched; rhizophores borne throughout stem length, or restricted to proximal 1 ⁄4–1⁄2 of stems, filiform to stout, 0.1–0.2 mm diam.; leaves of two kinds (anisophyllous) throughout stem length; lateral leaves ovate to ovate-oblong, (1.7–)2.5–3.3 ⫻ (0.7–)1.2–1.9 mm, bases rounded, usually strongly or slightly overlapping stems acroscopically, denticulate to entire, apices acute to long-acuminate, upper surfaces occasionally hispid near the basiscopic margins, with or without idioblasts, lower surfaces glabrous, with conspicuous idioblasts, median leaves broadly ovate, 1–2 ⫻ 0.7–1.2 mm, bases subcordate, outer bases auriculate, margins hyaline, denticulate, apices gradually acuminate to aristate, aristae 1⁄4 or less the lamina length, 0.1–0.3 mm long, upper surfaces with conspicuous or inconspicuous idioblasts; axillary leaves similar to lateral leaves, except the margins minutely denticulate proximally, entire distally, exauriculate; strobili lax and flattened, dorsiventral, 2–8 mm long; sporophylls dimorphic, the dorsal ones green, spreading, the ventral ones hyaline to pale green, ascending; megasporangia in two ventral rows; microsporangia in two

Distribution. Terrestrial or epipetric in mossy cloud forests, in marshy meadows, on steep forested slopes, pastures, along waterways and road banks, on wet mossy rocks, exposed to wind and fog on Sphagnum beds; 1400–2800 m. Mexico; Guat, Hond, CR, Pan; Col, Ven, Ec, Peru. Specimens Examined. Chis (Breedlove 24378, DS, 29437, LL, NY, 31859, DS, F, MICH, NY, 51445, 53366, 55973, CAS). Oax (Wendt et al. 6811, NY).

Selaginella moritziana comprises three varieties. In Mexico only var. moritziana is present; the other two are limited to northern South America. Selaginella moritziana is usually characterized by the presence of obscure to conspicuous idioblasts on the upper surface of median leaves, dorsal sporophylls, and occasionally on lateral leaves, by often conspicuous, short hairs on the upper surface of the lateral leaves near the basiscopic margins, by the median leaves with the inner margin hyaline throughout, composed of elongate cells, denticulate proximally and distally with the outer margin hyaline along distal half, composed of elongate cells, greenish along the proximal half, composed of Ⳳ polygonal to rounded cells, with stomata throughout. Selaginella moritziana varies in the following characteristics: habit, shape of lateral and median leaves, presence of idioblasts on the upper suface of the leaves and sporophylls, presence of submarginal hairs on the upper surface of the lateral leaves, and on the length of leaf apices. Wendt et al. 6811 is unusual in its lateral leaves long-ciliate along proximal half, median leaves with the outer base with 2– 8 long cilia, beige megaspores, sporophylls seemingly monomorphic, and quadrangular strobili. This specimen was found growing on very steep, almost vertical slopes, exposed to wind and fog on Sphagnum beds.

44. SELAGINELLA MOSORONGENSIS Selaginella mosorongensis Hieron., Hedwigia 43: 4. 1904. Type. Mexico. Veracruz: Near Motzorongo, between Tierra Blanca and

SELAGINELLA NOTHOHYBRIDA

583

Co´rdoba, Schumann 1905 (B; photo BM, US!). Figs. 268AA–EE.

Bryodesma mutica (D. C. Eaton) Soja´k, Preslia 64: 154. 1992. Lectotype (chosen by Weatherby, 1944: 413). U.S.A. Colorado: crevices of rocks, mountains of Colorado, 1871, Meehan s.n. (YU!, frag. NY!; isolectotypes NY!, MO).

Stems erect from decumbent base, greenish to stramineous, 25–50 cm long, ca. 2 mm diam., 4-angled, not articulate, not flagelliform, stoloniferous, 3–4 times branched; rhizophores restricted to stem bases; leaves monomorphic on main stems, of two kinds (anisophyllous) otherwise; lateral leaves on main stems below first branch reflexed, ovate, median leaves appressed, biauriculate with large, long-ciliate auricles, margins denticulate; lateral leaves ovate-deltate, 3–4 mm long on main stems, 1–2 mm long on branchlets, short-ciliate at bases, minutely denticulate distally, apices acute; median leaves ovate, biauriculate with one auricle much larger, margins long-ciliate, auricles ciliate, apices long-aristate; axillary leaves lanceolate, exauriculate, ciliatedenticulate in proximal half; strobili quadrangular, 3–4 mm long; sporophylls monomorphic and of two colors, dorsal rows green, ventral rows colorless; megasporangia in ventral two rows; microsporangia in dorsal two rows; megaspores cream, with low reticulum, 220–280 ␮m diam.; microspores tan, with rounded or capitate papillae, 19–23 ␮m diam.

Stems long- to short-creeping, forming loose mats, radially symmetric, 6–20 cm long, 0.2–0.3 mm diam., not articulate, not flagelliform, not stoloniferous, 1–2 times branched; rhizophores borne throughout stem length, filiform, 0.1–0.2 mm diam.; leaves of one kind (isophyllous) throughout stem length, tightly appressed, linear-lanceolate to lanceolate, 1–1.5 ⫻ 0.3–0.5 mm, bases rounded, glabrous, green, long-ciliate, denticulate to entire distally, cilia and teeth hyaline, apices blunt or acuminate to mucronate, mucro ca. 0.05 mm; strobili quadrangular, 3–5 mm long; sporophylls monomorphic; megasporangia few, scattered in ventral rows; microsporangia in two dorsal rows and scattered in ventral rows; megaspores yellow, rugose on proximal face, smooth on distal face, 300–320 ␮m diam.; microspores orange.

Distribution. Terrestrial or epipetric on limestone rocks; 1800 m. W USA; Mexico. Specimens Examined. Chih (Henrickson 12873, LL; Knobloch 2110, GH, MSC).

Distribution. Terrestrial in wet forests at low elevation; 0–350 m. Mexico. Specimens Examined. Oax (Breedlove 56694, CAS, NY; Hallberg 1647b, NY; Martı´nez-C. 84, CAS, F, NY, 331, UC, US, 830, US; Mickel 5812, NY, UC; Thurm et al. 228, NY). Ver (Leeds 147, PH).

Selaginella mosorongensis is distinguished from S. oaxacana by having long-aristate median leaves, each with a short-ciliate, basiscopic auricle, reflexed lateral stem leaves with large, long-ciliate auricles, axillary leaves short-ciliate at bases, and tan microspores. The auriculate median leaves, reflexed lateral stem leaves, tan microspores, and denticulate sporophylls of S. mosorongensis distinguish it from S. mickelii. See S. illecebrosa for differences from that species.

45. SELAGINELLA MUTICA Selaginella mutica D. C. Eaton ex Underw., Bull. Torrey. Bot. Club 25: 128. 1898. Figs. 278F–H.

Selaginella mutica var. mutica is a very distinct taxon characterized by its tightly appressed, ciliate, mucronate leaves. The tightly appressed leaves give the stems the appearance of a woven rope. Mexican material belongs to var. mutica; var. limitanea Weath., with short-ciliate, long-aristate leaves, occurs in southwestern United States (Arizona to Texas).

46. SELAGINELLA NOTHOHYBRIDA Selaginella nothohybrida Valdespino, Brittonia 44: 319. 1992. Type. Mexico. Guerrero. Mpio. Zupango del Rı´o: 38 km al S de Iguala, ca. 0.5 km al N de Xilitla, por la carretera a Acapulco, Koch et al. 7989 (NY!; isotypes MEXU!, MO!, NY!, PMA!). Figs. 280N–T. Stems rosette-forming, glabrous, not articulate, not flagelliform, not stoloniferous, 9–15 cm long, 0.5–1 mm diam., 1–4 times branched, branches flat and spreading when moist, curling inwards when dry (ball-like); rhizophores on proximal half of branches, 0.2–0.3 mm diam.; leaves of two kinds (anisophyllous)

584

SELAGINELLA

throughout stem length, green above (in new growth), to yellowgreen, yellow, or brown (when old), without reddish patches; lateral leaves (on primary branches) imbricate, ovate, 1.4–2.3 ⫻ 0.8–1.4 mm, bases cordate, acroscopic bases strongly overlapping stems, margins broadly hyaline or whitish, abundantly ciliate, apices long-cuspidate to aristate, hyaline; median leaves (on primary branches) imbricate, ovate-lanceolate or lanceolate, 1–1.7 ⫻ (0.5–)0.7–1.2 mm, bases rounded, margins hyaline, apices short- to long-aristate, tips as in lateral leaves; axillary leaves ovate-lanceolate to lanceolate, bases cordate with two incurved, ciliate auricles, margins hyaline, ciliate, apices short- to longaristate; strobili quadrangular, 4–10 mm long; sporophylls monomorphic; megasporangia in two ventral rows; microsporangia in two dorsal rows and ventral tip of strobili; megaspores deep yellow, reticulate-rugose to long-papillate to short-baculate, 300– 375 ␮m diam.; microspores orange, rugose to baculate, 25–32 ␮m diam.

47. SELAGINELLA NOVOLEONENSIS Selaginella novoleonensis Hieron. in Engler & Prantl, Nat. Pflanzenfam. 1(4): 676. 1901. Type. Mexico. Nuevo Leo´n: Sierra de la Silla, near Monterrey, Pringle 2489 (presumably B; isotypes ARIZ!, DS!, MEXU!, MSC!, NY!, UC!, US!). Figs. 279G, H, J–L. Stems rosette-forming, green, 15 cm long, 0.5–1.5 mm diam., not articulate, not flagelliform, not stoloniferous, brownish or reddish, 2–3 times branched; rhizophores restricted to stem bases, 0.3–0.5 mm diam.; leaves of two kinds (anisophyllous) throughout stem length, green above or reddish at tips, green, golden, or brownish to reddish below; lateral leaves ovate-deltate to suborbicular, 1.5–3.5 ⫻ 1.7–2.3 mm, bases rounded, margins green, short-ciliate proximally, denticulate distally, apices cuspidate to short-aristate; median leaves ovate, 1.2–2.5 ⫻ 0.8–2 mm, bases rounded, margins hyaline, ciliate-denticulate, to denticulate or entire distally, apices cuspidate to short-aristate; axillary leaves similar to lateral leaves, exauriculate; strobili quadrangular, 5–10 mm long; sporophylls monomorphic, ventral ones sometimes paler green; megasporangia in two ventral rows; microsporangia in two dorsal rows; megaspores yellow to tan, reticulate with high ridges, 300–450 ␮m diam.; microspores orange, shed in tetrads.

Distribution. Terrestrial or epipetric in rock crevices on dry rock faces, or shady places on deciduous forest slopes; 450– 1800 m. Mexico. Selected Specimens Examined. Gro (Bruce et al. 1495, MEXU, MICH; Calzada 15715, 16369, NY; Lozano 33, NY; Matuda s.n., MEXU; McVaugh 22170, MICH; Orcutt 2559, DS). Oax (Mickel 4097, NY). Pue (Bo´ege 1924, MEXU). Ver (Oro s.n., MEXU).

Selaginella nothohybrida is easily distinguished from S. lepidophylla by its median leaves that have margins short-ciliate (vs. serrate to very short-ciliate), leaf apices long-acuminate to aristate (vs. acuminate to short-cuspidate), lateral leaves with acroscopic margins ciliate to fimbriate (vs. serrate to very shortciliate), lower leaf surfaces lacking reddish patches (vs. with reddish patches), sporophylls ovate-lanceolate (vs. deltate-ovate), and sporophyll bases strongly pubescent (vs. weakly puberulent). Selaginella nothohybrida can be distinguished from S. pallescens by its rosette-like habit (vs. mostly erect to sometimes rosettelike), median leaf bases slightly peltate and truncate to roundish, without auricles (vs. non-peltate, cordate, with two incurved auricles or sometimes only an outer one on main stem), sporophyll bases strongly pubescent (vs. glabrous), and sporopyll apices abruptly long-cuspidate to short-aristate (vs. gradually tapering, long-acuminate to long-aristate).

Distribution. Terrestrial or epipetric in rock crevices, dry rock faces, limestone slopes; 300–1900 m. Mexico. Selected Specimens Examined. BCS (Turner & Lowe 59-129, NY). Chih (Knobloch 2106, MSC). Coah (Sa´nchez 779, US). Gro (Clark 7243, NY). Me´x (Hinton et al. 6859, NY). NL (Copeland herb. 173, US; Pringle 2038, NY). Son (Lloyd 506, NY, US). Tam (Bartlett 10177, DS, ENCB, NY; King 4559, NY, US).

Selaginella novoleonensis is distinguished from S. lepidophylla by the former having median leaves long-acuminate to aristate (vs. acute to mucronate), leaf margins narrowly hyaline (ca. 0.05 mm wide vs. broadly, ca. 0.1 mm), and the median leaf bases and stems with hairs short, frequently inconspicuous (vs. long, conspicuous).

SELAGINELLA ORIZABENSIS

48. SELAGINELLA OAXACANA Selaginella oaxacana Spring, Nouv. Me´m. Acad. Roy. Sci. Bruxelles 24: 177. 1850. Type. Mexico. Oaxaca: Chinantla, Galeotti 6608 bis (presumably LG; isotypes P!, BR! as 6608, same no. as cited for S. martensii). Figs. 267U–Z. Lycopodium flabellatum L. var. strictum M. Martens & Galeotti, Me´m. Foug. Mexique 12. 1842. Type. Mexico. Oaxaca: Galeotti 6608 (BR!; isotype P!; see above).

Stems erect, bases decumbent with age, stramineous to pale green, to 90 cm tall, 2–3 mm diam., not articulate, not flagelliform, stoloniferous, mostly 3-times branched, frond-like; rhizophores restricted to stem bases, to 1 mm diam.; leaves on main stem monomorphic, of two kinds (anisophyllous) above first branch, glabrous, with idioblasts (especially visible on upper surface on lateral and median leaves), leaf margins often curling on drying, appressed on main stems below first branch, ovate, bases rounded, margins minutely denticulate, apices acute; lateral leaves obliquely oblong, ca. 4–5 mm long on main stems, ca. 3 mm long on branches, 2 mm wide, bases truncate to rounded along basiscopic margins, rounded along acroscopic margins, margins minutely denticulate at bases, entire distally, apices obtuse; median leaves ovate, denticulate, oblique at bases with small subequal auricles, apices acuminate; axillary leaves ovate, bases rounded to subcordate, exauriculate, margins minutely denticulate at bases; strobili quadrangular, 5–10 mm long; sporophylls monomorphic and of two colors, dorsal rows green, ventral rows colorless, ascending; megasporangia in two ventral rows; microsporangia in two dorsal rows; megaspores cream-white, with low narrow ridges to nearly smooth, 260–420 ␮m diam.; microspores orange-yellow, nearly smooth to short-papillate.

585

rongensis and S. cuneata, but S. mosorongensis can be distinguished by its reflexed stem leaves with large, long-ciliate auricles, more ciliate, long-aristate median leaves with unequal auricles, ciliate axillary leaves, and tan megaspores. Selaginella cuneata is distinct in its cuneate-based lateral and axillary leaves and relative lack of idioblasts.

49. SELAGINELLA ORIZABENSIS Selaginella orizabensis Hieron., Hedwigia 41: 193. 1902. Syntypes. Mexico. Veracruz: Orizaba, 1853, Mu¨ller 268 (isosyntype NY!); near Eugenio, mountains of Sierra de San Cristo´bal, Sartorius? s.n. [ex herb. Schultz-Bipont.] (presumably B). Figs. 285N–S. Stems suberect, stramineous, 11 cm long, 0.2–0.5 mm diam., not articulate, not flagelliform, stoloniferous, 2–3 times branched; rhizophores in proximal 1⁄4 of stems, filiform, 0.1–0.2 mm diam.; leaves of two kinds (anisophyllous) throughout stem length; lateral leaves broadly-ovate to ovate-oblong, (1.4–)1.7–2.3 ⫻ (0.7–)0.9–1.6 mm, bases rounded, strongly overlapping stems acroscopically, margins hyaline, long-ciliate to denticulate or entire acroscopically, entire or denticulate basiscopically, apices acute to obtuse, both surfaces glabrous, without idioblasts; median leaves ovate to ovate-lanceolate, 0.9–1.2 ⫻ 0.5–0.7 mm, bases subcordate, margins hyaline, long- to short-ciliate, apices long-acuminate to aristate, aristae 0.2–0.5 mm, both surfaces glabrous, without idioblasts; axillary leaves similar to lateral leaves or more ovate-lanceolate, exauriculate; strobili quadrangular to slightly dorsiventral, 2–2.6 mm long; sporophylls monomorphic; megasporangia in two ventral rows or only two at base; microsporangia occupying most of strobilus; megaspores yellow, with a low equatorial flange, reticulate to rugose-reticulate, 300– 350 ␮m diam.; microspores deep orange.

Distribution. Terrestrial in montane rain forests; 250–1800 m. Mexico; Guat, Hond, CR, Pan; Col, Ec. Selected Specimens Examined. Chis (Breedlove 32599, DS, F, NY; Ghiesbreght 602, NY; Rovirosa 868, NY). Oax (Gonza´lez & Conzatti 734, US; Hallberg 1556, NY, UC, US; Martı´nez 258, CAS, ENCB, F, NY; Mickel 5677, NY, UC). Ver (Lira 136u, ENCB; Nee et al. 24688, CAS, NY; Pe´rez 168, ENCB).

This is the most common of the tall, caulescent Mexican selaginellas. In general, S. oaxacana closely resembles S. moso-

Distribution. Terrestrial, habitat unknown; elevation unknown. Mexico. Specimen Examined. Ver (only an isosyntype).

Selaginella orizabensis is a poorly known species, apparently closely related to S. mixteca. They can be separated by the characters given in the key.

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SELAGINELLA

50. SELAGINELLA PALLESCENS Selaginella pallescens (C. Presl) Spring in Martius, Fl. Bras. 1(2): 132. 1840. Figs. 267A–F.

Unverified, Doubtful, or Mistaken Reports. Ags (De la Cerda & Garcı´a 1232, HUAA, and 16 other collections, cited by SiqueirosDelgado & Gonza´lez-Adame, 2004, but not verified). Qro (Ferna´ndez 3385, 3440, 3463, ENCB, and Rzedowski 44092, 46449, QMEX, all cited by Arreguı´n et al., 2001, but not verified).

Lycopodium pallescens C. Presl, Reliq. Haenk. 1(1): 79. 1825. Type. Mexico. Haenke s.n. (PR; photo BM ). Lycopodium cuspidatum Link, Hort. Berol. 2: 161. 1833. Selaginella cuspidata (Link) Link, Fil. Spec. 158. 1841. Type. A cultivated plant, said to be from Mexico (presumably B). Selaginella cuspidata (Link) Link var. elongata Spring, Nouv. Me´m. Acad. Roy. Sci. Bruxelles 24: 67. 1850. Type. Guatemala. Collector not named (presumably LG).

This species exhibits extreme variation in habit and habitat, from caespitose to erect. See S. harrisii, S. polyptera, and S. pulcherrima for comparison with those species.

Stems erect or rosette-forming, caespitose, glabrous, lustrous, whitish, 4–25 cm tall, ca. 1 mm diam., angled when dry, not articulate, not flagelliform, not stoloniferous, 2–3 times branched; rhizophores limited to stem bases; leaves on main stems monomorphic, tan, ovate, bases with two long, almost overlapping auricles, margins long-ciliate, apices cuspidate, of two kinds (anisophyllous) above first branch; lateral leaves ovate, 1– 2 mm long, bases rounded to cordate, margins long- to shortciliate, lustrous, old leaves light tan, apices aristate; median leaves ovate, bases with a single short auricle, margins short-ciliate with a broad white border, apices aristate, aristae 0.3–0.5 mm; axillary leaves lanceolate, exauriculate, long-ciliate at bases to shortciliate; strobili quadrangular, 3–6 mm long; sporophylls monomorphic; megasporangia basal or subbasal; microsporangia in distal portions of strobili; megaspores white or cream to light yellow, with ridges very low and anastomosing, 387–448 ␮m diam.; microspores bright orange to scarlet, capitate-papillate, usually shed singly but sometimes in tetrads.

Selaginella parishii Underw., Bull. Torrey Bot. Club 33: 202. 1906. Figs. 274G, H, J–M.

51. SELAGINELLA PARISHII

Bryodesma parishii (Underw.) Soja´k, Preslia 64: 154. 1992. Type. Mexico. Zacatecas: Palmer 306 (NY!; isotypes F!, UC!)

Stems prostrate, forming rather loose mats, dorsiventral, 3–14 cm long, 0.3–0.5 mm diam., not articulate, not flagelliform, not stoloniferous, 2–3 times branched, branches horizontal or curving upward and curling (in dry conditions); rhizophores throughout stem length, 0.2–0.3 mm diam.; leaves of one kind (isophyllous) throughout stem length; upperside leaves spreading, green to grayish green or light brown (in old leaves), narrowly lanceolate to linear-lanceolate, slightly falcate on lateral rows, 1.2–1.8 ⫻ 0.4–0.6 mm (widest at base), bases abruptly adnate (distinct from stem), truncate, pubescent, hyaline, margins ciliate, cilia hyaline, apices broadly acute to obtuse; underside leaves appressed, imbricate, buff to light brown or golden-brown, lanceolate, 2–3 ⫻ 0.5–1 mm (widest at middle), bases truncate to decurrent, glabrescent, margins ciliate, cilia hyaline, apices acute to obtuse, 0.1–0.4 mm; strobili quadrangular, 5–10 mm; sporophylls monomorphic; megasporangia in two ventral rows; microsporangia in two dorsal rows; megaspores lemon-yellow, reticulate with low ridges, 250–300 ␮m diam.; microspores light orange, granulose, shed mostly in tetrads.

Distribution. Terrestrial or on rocks in pine-oak forests, mesic woods; 50–2700 m. Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Col, Ven, Sur, Braz. Selected Specimens Examined. BCS (Carter et al. 2335, DS, UC). Chih (Knobloch 553, UC). Chis (King 2755, UC). Coah (Arse`ne 3453, MEXU). Col (Orcutt 4617, NY p.p.). DF (Matuda 22002, UC). Dgo (Palmer 886, NY, UC). Gro (Hinton et al. 10282, LL). Gto (Morley 652, UC). Jal (Pringle 2592, UC). Me´x (Tejero-Dı´ez 2319, IZTA). Mich (Zu´n˜iga G. 18290, UC). Mor (Harris 23, UC). Nay (Mexia 653, UC). NL (Pringle 1979, UC). Oax (Hellwig 1310, LL, NY). Pue (Arse`ne 7084, MEXU). Sin (Mexia 221, UC). Son (Pennell 19513, PH). Tam (Bartlett 10099, DS). Ver (Purpus 8925, UC). Zac (Breedlove 63225, CAS).

Distribution. Terrestrial or epipetric on sandstone crevices, calcareous gravelly soil; 1600–2900 m. Mexico. Specimens Examined. Coah (Arse`ne 3453, MEXU; Johnston et al. 11006, LL, MEXU; Pennell 17272, MEXU, MICH, PH, US). NL (Hinton 21112, ANSM). Pue (Arse`ne 7084, MEXU). Zac (only the type).

SELAGINELLA PILIFERA

52. SELAGINELLA PERUVIANA Selaginella peruviana (Milde) Hieron., Hedwigia 39: 307.1900. Figs. 273F–H, J, K. Selaginella rupestris (L.) Spring f. peruviana Milde, Fil. Eur. 263. 1867; Bryodesma peruviana (Milde) Soja´k, Preslia 64: 155. 1992. Type. Peru. Huanuco: Ruı´z s.n. (B; isotype NY!). Selaginella sheldonii Maxon, Proc. Biol. Soc. Wash. 31: 171. 1918. Type. U.S.A. Oklahoma: Quanah Mountain, Indian Territory, Sheldon 233 (US!).

Stems prostrate, forming loose mats, not readily fragmenting, dorsiventral, irregularly forked, branch tips upturned; rhizophores borne throughout stem length, 0.2–0.3 mm diam.; leaves of one kind (isophyllous) throughout stem length, tightly appressed, green, apices aristate, aristae whitish, 0.3–0.8 mm long; upperside leaves linear-lanceolate on central rows to falcate on marginal rows, 2.3–2.8 ⫻ 0.5–0.65 mm, bases abruptly adnate, pubescent, margins ciliate, cilia transparent to opaque, ascending or spreading, 0.1–0.2 mm; underside leaves linear-lanceolate on central rows to falcate on marginal rows, 2.5–4 ⫻ 0.4–0.6 mm, bases decurrent to oblique on marginal rows, pubescent or sometimes glabrous, margins ciliate, cilia hyaline to opaque, spreading at base, ascending toward apex, 0.1–0.2 mm; strobili quadrangular, 0.5–2 cm long; sporophylls monomorphic; megaspores yellow to orange, rugose-reticulate; microspores orange.

587

Selaginella peruviana can be distinguished from S. arizonica by its underside leaves widest at base (vs. at middle), leaf apices each with a terete arista 0.3—0.8 mm long (vs. flattened, 0.1– 0.3 mm long), and sporophylls aristate (vs. acute to acuminate).

53. SELAGINELLA PILIFERA Selaginella pilifera A. Braun, Index Seminum (Berlin), App. 20. 1857. Type. U.S.A. Texas: El Paso Co., El Paso, 1849, Wright s.n. (B; US! frag ex B). Figs. 280A–F. Selaginella pringlei Baker, Handb. Fern-Allies 88. 1897. Selaginella pilifera A. Braun var. pringlei (Baker) C. V. Morton, Amer. Fern J. 29: 15. 1939. Type. Mexico. Chihuahua: Santa Eulalia Mts., Pringle 271 (BM; isotypes CAS, NY!, US!).

Stems rosette-forming, flat when moist, almost flat when dry, glabrous, not articulate, not flagelliform, not stoloniferous, 2–3 times branched; rhizophores restricted to bases of rosette, 4–5 mm diam.; leaves of two kinds (anisophyllous) throughout stem length, thick and stiff; lateral leaves imbricate, elliptic to ellipticovate, (2–)3–3.5 ⫻ 0.8–1 mm, bases cordate, margins hyaline, short-ciliate proximally, denticulate distally, apices aristate, aristae 1⁄3–1⁄2 the lamina length; median leaves oblique-lanceolate, 2– 3 ⫻ 0.7–1 mm, bases peltate, rounded to truncate, margins green to slightly hyaline, denticulate, short-ciliate at bases, apices aristate, aristae 1⁄3 the lamina length; axillary leaves lanceolate, bases cordate, exauriculate, margins short-ciliate at bases, mostly denticulate, apices long-acuminate; strobili quadrangular, 5–10 mm; sporophylls monomorphic; megasporangia basal in ventral rows; microsporangia in dorsal rows, also in ventral rows distally; megaspores yellow, ca. 250 ␮m diam.; microspores orange; 2n⫽20.

Distribution. Terrestrial or epipetric on rocky slopes, rock crevices, ledges of sandstone or igneous cliffs, less often on sandy or clay soil; 1600–3200 m. USA; Mexico; Peru, Bol, Arg. Selected Specimens Examined. Ags (Rzedowski 25004, ENCB). Chih (Harvey 1454, MICH). Coah (Stanford et al. 118, ARIZ). DF (Knobloch 1060, MICH, UC). Dgo (Pennell 18126, PH). Gto (Rzedowski 4788a, NY). Hgo (Steyermark 52276, F). Jal (Rollins & Tryon 58320, MICH). Mich (Dı´az-Barriga 4745, NY). NL (Hinton et al. 20938, TEX). Oax (Mickel 4496, NY). Pue (Arse`ne 7087, MEXU). Qro (Pe´rez & Zamudio 3583, NY). SLP (Lundell 56, MICH). Son (Phillips 800, MICH). Tlax (Vibrans 385, MEXU). Ver (Barrington 450, MEXU). Zac (Pennell 17476, PH). Unverified, Doubtful, or Mistaken Reports. Me´x (Schaffner 933, GH, YU, cited by R. Tryon, 1955, but not verified).

Distribution. Terrestrial or epipetric on dry rocky soil, rock crevices, limestone rocks, and cliff faces; 475–2000 m. USA; Mexico. Selected Specimens Examined. Chih (Pringle 886, NY). Coah (Arse`ne 10677, US; Rollins & Tryon 58156, US). Dgo (Torke et al. 155, NY). Hgo (Tejero-Diez 4057, IZTA). NL (Frye & Frye 2466, NY, UC, US; Pringle 13959, ARIZ). Qro (Dı´az-Barriga & Carranza 7477, NY; Sohmer

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SELAGINELLA

9235, MEXU, NY). SLP (King 3873, NY, US; Rzedowski 11493, ENCB, NY). Tam (Johnston 12784, UC; Sharp et al. 52211, US).

The long-bristled leaf apices of Selaginella pilifera are unique among New World xerophytic members of series Circinatae Spring. The closest relative of S. pilifera is S. gypsophila, which differs by having obtuse leaf species. Further studies are needed to determine whether S. gypsophila represents a well differentiated species or an environmental variant of S. pilifera.

54. SELAGINELLA POLYPTERA Selaginella polyptera Valdespino, sp. nov. Type. Mexico. Oaxaca: 4–5 mi E of Xilitla, near natural bridge, Mickel 599 (holotype NY!; isotype MICH!). Figs. 282G, H, J–N. A. S. pulcherrima foliis conspicue dimorphis in caulibus principalibus infra ramos primos, foliis lateralibus patentibus, foliis medianis longiaristatis, aristis longitudine ⬎1⁄3 foliorum aequantibus dignoscenda; a S. harrisii primo charactere differt. (Gr., poly, many, pteron, wing, alluding to the spreading, imbricate lateral leaves that give the apprearance of wings.)

Stems erect, fasciculate, to 45 cm tall, 1–1.5 mm diam., light green to stramineous, not articulate, not flagelliform, not stoloniferous, 2–4 times branched; rhizophores restricted to stem bases, stout, 0.3–0.8 mm diam.; leaves of two kinds (anisophyllous) throughout stem length; lateral leaves imbricate below first branch to distant, ovate to ovate-deltate, 2.5–4 ⫻ 1.5–2.6 mm, bases rounded, margins hyaline, long-ciliate proximally, denticulate distally, apices long-acuminate; median leaves appressed, broadly ovate to ovate-deltate, 1.5–2.5 ⫻ 1–1.5 mm, bases rounded or cordate, margins hyaline, long-ciliate proximally, denticulate distally, apices long-acuminate; axillary leaves broadly ovate to ovate, bases shortly biauriculate, margins ciliate proximally, denticulate distally, apices acuminate; strobili quadrangular, 3–10 mm long; sporophylls monomorphic (ventral ones sometimes lighter green); megasporangia scattered on two ventral rows; microsporangia in two dorsal rows and mixed on ventral rows; megaspores pale yellow, reticulate with a low equatorial flange, 250– 300 ␮m diam.; microspores orange.

Distribution. Terrestrial or epipetric on and among limestone rocks, in seasonal cloud forests; 180–1750 m. Mexico; Bel.

Selected Specimens Examined. Chis (Breedlove 56384, CAS). Hgo (Fraile & Pe´rez 12a, ENCB; Hellwig 1235, NY; Moore 5080, BH, UC). Oax (Geith 95-M121, NY; Hallberg 1375, NY). SLP (King 4328, NY, TEX, UC, US; Mickel 599, NY). Ver (Bourgeau 2024, US; Nee & Taylor 26657, NY, UC). Bel (Bartlett 11321, UC).

Selaginella polyptera is part of a taxonomically confusing species group including S. acutifolia, S. harrisii, S. pallescens, and S. pulcherrima. It seems intermediate between S. pallescens and S. pulcherrima. Selaginella polyptera can be distinguished from S. pallescens by its axillary leaves broadly ovate to ovate (vs. ovate to ovate-lanceolate), with the bases subcordate (vs. cordate), with only an obscure outer auricle (vs. with inner and outer conspicuous auricles). It differs from S. harrisii and S. pulcherrima mainly by its obviously dimorphic, ascending to spreading (vs. leaves seemingly monomorphic, strongly clasping the stems) leaves on the main stems below the first branch. Selaginella polyptera is remarkable by having median leaves on main stems with wide hyaline margins to ca. 4–6 cells wide and long-aristate apices, the aristae hyaline and ca. 1⁄3 or more as long as the lamina length.

55. SELAGINELLA POPAYANENSIS Selaginella popayanensis Hieron., Hedwigia 43: 9. 1904. Type. Colombia. Cauca: near Popaya´n, Lehmann 6968 (B!; isotype K, US!). Figs. 271A–G. Selaginella anisoclada Alston ex Crabbe & Jermy, Fern Gaz. 11: 255. 1976. Type. Venezuala. Aragua: above Guamitas, Parque Nacional, Alston 5814 (BM!; isotype US!). Selaginella disticha Mickel & Beitel, Mem. New York Bot. Gard. 46: 337. 1988. Type. Mexico. Oaxaca: Dist. Ixtla´n, 7 km S of Vista Hermosa, Mickel 6735 (NY!; isotype UC!).

Stems creeping or ascending to suberect, stramineous to greenish, 4–12 cm long, 0.3–0.6 mm diam., not articulate, not or infrequently flagelliform, stoloniferous or not, 2–3 times branched; rhizophores throughout or restricted to proximal 1⁄2 of stems, filiform, 0.1–0.2 mm diam.; leaves of two kinds (anisophyllous) throughout stem length; lateral leaves ovate to ovateoblong, 1.5–3.1 ⫻ 1–1.8 mm, bases rounded, acroscopic bases strongly overlapping stems, margins pale green, entire, apices acute, upper surfaces hispid near the basiscopic margins, without idioblasts, lower surfaces glabrous, with obscure idioblasts; median leaves ovate-lanceolate, 0.8–1.5 ⫻ 0.4–0.9 mm, bases rounded, margins hyaline, denticulate, apices acute to acuminate, the tips 1⁄4 or less the lamina length, 0.1–0.2 mm long, both surfaces glabrous, without idioblasts; axillary leaves ovatelanceolate, bases rounded, exauriculate, margins entire, apices acute; strobili lax and flattened, dorsiventral, (2–)4–15 mm long; sporophylls dimorphic, the dorsal ones green, spreading, the ventral ones hyaline to pale green, ascending; megasporangia in two ventral rows; microsporangia in two dorsal rows or absent; megaspores orange-buff or light to deep orange-yellow, frequently with a prominent equatorial flange, reticulate to corrugate-reticulate with moderately high ridges, 240–295 ␮m diam.; microspores orange.

SELAGINELLA PORPHYROSPORA

Distribution. Terrestrial or epipetric on ground or on wet mossy rocky banks in montane forests; 1400–1900 m; Mexico; Guat, CR, Pan; Col, Ven, Bol.

589

usually with one long idioblast along each side of midribs, or occasionally idioblasts absent; median leaves ovate to ovatelanceolate, 0.9–1.5 ⫻ 0.5–0.9 mm, bases rounded to subcordate, margins hyaline, sparingly denticulate, apices abruptly aristate, aristae half to nearly equalling the expanded lamina length; axillary leaves ovate-lanceolate, bases rounded, exauriculate, margins entire, apices acute; strobili lax and flattened, dorsiventral, or occasionally weakly quadrangular, 3–20 mm long; sporophylls dimorphic, the dorsal ones green, spreading, the ventral ones pale green to hyaline, ascending; megasporangia in two ventral rows, or strobili wholly megasporangiate; microsporangia in two dorsal rows, or strobili wholly microsporangiate; megaspores orange, with a well developed equatorial flange, reticulate to reticulate-rugulate with high, dense ridges, 250–265 ␮m diam.; microspores orange.

Specimens Examined. Chis (Breedlove 32239, MICH, 32680, DS, NY; Gittins 4238-B, NY). Gro (Lorea 2998, F, NY). Oax (McAlpin 1022, CAS p.p., F; Mickel 1110a, MICH, NY, US, 1560, 4295, NY).

This species has often been misinterpreted. Alston (1952), Fraile (in Davidse et al., 1995), and Stolze (1983) referred many of the specimens of S. popayanensis cited here to S. porphyrospora A. Braun, mostly because the lateral leaves are usually hispid on the upper surface near the basiscopic margin. Selaginella popayanensis is easily distinguished from S. porphyrospora by lacking “false veins” (idioblasts) on the lateral leaves, deep orange microspores, and acute lateral leaves. Selaginella popayanensis is closely related to a species group including S. moritziana var. moritziana, also found in the flora area, and S. cavifolia A. Braun and S. flacca Alston in South America.

56. SELAGINELLA PORPHYROSPORA Selaginella porphyrospora A. Braun, Ann. Sci. Nat. Bot., se´r 5, 3: 286. 1865. Type. Mexico. Veracruz: [Mirador?], Sartorius s.n. (B!, frag. BM!). Figs. 269Z, AA–FF. Selaginella bulbifera Baker, Gard. Chron. 27: 783, 950. 1867. Type. From garden material of unknown origin (fide Alston et al. 1981: 214). Selaginella binervis Liebm. ex Baker, J. Bot. 22: 112. 1884. Type. Mexico. Veracruz: Dos Puentes, San Antonio Huatusco, Liebmann s.n. (K!; isotypes B, C, P, frag. US!).

Stems creeping, ascending to suberect, stramineous or greenish, (3–)5–12(–23) cm long, 0.2–0.5 mm diam., not articulate, frequently with flagelliform, gemmiferous apices, stoloniferous, 1–3 times branched; rhizophores restricted to proximal 1⁄3–1⁄2 of stems, filiform, 0.1–0.3 mm diam.; leaves of two kinds (anisophyllous) throughout stem length; lateral leaves ovate, 2–3.4 ⫻ 1–2 mm, bases rounded to subcordate, acroscopic bases strongly overlapping stems, margins hyaline, denticulate or ciliatedenticulate proximally, denticulate distally, pale green, entire to sparsely denticulate, apices acuminate, upper surfaces usually hispid, especially near basiscopic margins, one or both surfaces

Distribution. Terrestrial or epipetric on moist banks and on wet mossy rocks in or near waterfalls, streams, roadside banks, or forest clearings, mostly in humid forests; 750–2500 m. Mexico; Guat, Hond, Salv, Nic, CR, Pan; Col, Ven, Ec. Selected Specimens Examined. Chis (Purpus 6720, BM, F, GH, M, NY, UC). Col (McVaugh 16097A, US). Gro (Lorea 1656b, NY). Hgo (Breedlove 59427, CAS, NY). Jal (Pringle 2593, BM, BR, DS, E, F, K, LL, NY, P, UC, US, W). Me´x (Tejero Dı´ez 2233, TEX). Mich (King & Soderstrom 4746, NY, TEX, UC, US). Mor (Rose & Painter 6943, BM, US). Nay (Yatskievych & Beetle 81-347, ARIZ). Oax (Mickel 3748, NY, UC). Pue (Orcutt 4035, BM, DS, GH, K, MEXU). Sin (Breedlove 35686, CAS, MICH, MO). Son (Gentry et al. 19317, NY, US). Ver (Purpus 7438, GH, NY, UC, US).

Selaginella porphyrospora is distinguished by flagelliform stems, primary branches, and tips of strobili, which typically produce round buds distally. It is probably sister to S. bernoullii (which see for discussion). Selaginella popayanensis also has hispid lateral leaves, but can be distinguished from S. porphyrospora by its lack of conspicuous idioblasts on the upper surface of the lateral leaves, light orange microspores, and acute lateral leaves. The microsporangia of S. porphyrospora, with microspores included, have a distinct raspberry color, hence the species epithet. As previously reported (Alston, 1955), some specimens of S. porphyrospora lack conspicuous idioblasts and/or conspicuous hairs on the upper surface. For the most part, these specimens are easily distinguished by the other characteristics here mentioned.

590

SELAGINELLA

57. SELAGINELLA PROLIFERA

MEXU; Mickel 5128, 5144, 5972, 6912, NY, UC; Orcutt 5238, DS; Williams 9800, F). Nic (Hitchcock s.n., 1–8 Nov 1911, US; Maxon 7745, US, 7750a, US; Oersted 2078, US).

Selaginella prolifera Valdespino, sp. nov. Type. Mexico. Oaxaca: Juchita´n, Isthmus of Tehuantepec, 6 km S of Matı´as Romero, Mickel 6880 (holotype NY!; isotypes MEXU!, PMA!, UC!). Figs. 270A–E.

Mickel and Beitel (1988) treated most collections of S. prolifera from Oaxaca as S. rhizophora Baker. We now agree with other authors (e.g., Alston, 1955; Lellinger, 1989; Smith, 1981) in concluding that S. rhizophora is conspecific with S. flagellata. Thus, specimens representing S. rhizophora as defined by Mickel and Beitel (1988) are in need of a name, which is provided here. Selaginella prolifera differs from S. flagellata by having median and lateral leaves and sporophylls with ciliate (vs. denticulate) margins and upper surfaces with (vs. without) idioblasts. The upright stems in S. prolifera arise from underground or partially buried stolons. In addition to this characteristic, S. prolifera also agrees with other erect to suberect species such as S. lineolata, S. finitima, and S. tuberosa in having the median leaves with an outer auricle; this suggests a close relationship among these taxa.

A. S. flagellata foliis medianis, foliis lateralibus, et sporophyllis marginibus ciliatis, superne idioblastis in structis diversa. (L. proliferus, producing offshoots, referring to the upright stems that arise from stolons.)

Stems suberect to erect, stramineous, 4–20 cm long, 0.3–0.6 mm diam., not articulate, not flagelliform, stoloniferous, 2–3 times branched; rhizophores restricted to proximal 1⁄4–1⁄3(–1⁄2) of stems, filiform, ca. 0.2 mm diam.; leaves of two kinds (anisophyllous) throughout stem length; lateral leaves narrowly oblong to oblong-ovate, 2–3 ⫻ 0.9–1.4 mm, bases rounded to subcordate, strongly overlapping stems acroscopically, margins narrowly hyaline to pale green, long-ciliate proximally, minutely denticulate or entire distally, apices acute, both surfaces glabrous, with idioblasts; median leaves broadly ovate-elliptic or ovateoblong, 1.3–1.6 ⫻ 0.7–1.2 mm, outer base with a well developed auricle, margins hyaline, long-ciliate, apices long-aristate, aristae half or more the lamina length, 0.5–0.7 mm long, both surfaces glabrous, upper surfaces with idioblasts; axillary leaves lanceolate to lance-ovate, bases rounded, margins pale green to hyaline, long-ciliate proximally, denticulate distally, apices acute; strobili loosely quadrangular to weakly flattened, 2–9 mm long; sporophylls slightly dimorphic, the dorsal ones green, spreading to ascending, the ventral ones pale green, ascending; megasporangia generally in two ventral rows or at the base of ventral rows, or infrequently absent; microsporangia in two dorsal rows, frequently also on the apical region of ventral side of strobili, or strobili wholly microsporangiate; megaspores snow-white, without an equatorial flange or with a slightly developed one, granulose-striate to granulose-reticulate with high ridges, 220– 255 ␮m diam.; microspores orange.

Distribution. Terrestrial or epipetric on banks of waterways, shaded roadside banks, or on moist rocky banks; 50–900 m. Mexico; Nic. Selected Specimens Examined. Chis (Breedlove 37635A, DS). Gro (Langlasse´ 421, GH, MEXU, US). Oax (Hallberg 1655, NY; Hellwig 1319,

58. SELAGINELLA PULCHERRIMA Selaginella pulcherrima Liebm. ex E. Fourn., Mexic. Pl. 1: 147. 1872. Lectotype (chosen by Alston, 1955: 239). Mexico. Veracruz: near Hacienda de Jovo, Liebmann 2061 (C!; isolectotypes BM!, K!, NY!, US!). Figs. 281A–G. Stems erect, fasciculate from short-creeping rhizomatous bases, green, 15–75 cm long, 0.5–2.5 mm diam., not articulate, not flagelliform, not stoloniferous, 3–4 times branched, frondlike; rhizophores restricted to stem bases, stout, 0.5–1 mm diam.; leaves of two kinds (anisophyllous) throughout most of plant, monomorphic and strongly clasping the stems below the first to fifth well developed lateral branches; lateral leaves ovate, 2.5–4 ⫻ 2–3 mm, bases subcordate to subauriculate, acroscopic bases rounded, margins hyaline to greenish, short-ciliate proximally, denticulate to entire distally, apices long-acuminate to aristate, surfaces glabrous, without conspicuous idioblasts; median leaves ovate, 3–3.5 ⫻ 1.5–2 mm, bases subcordate, margins greenish to slightly hyaline, short-ciliate proximally, denticulate distally, apices acuminate to aristate; axillary leaves ovate to ovate-lanceolate, bases subcordate to subauriculate, margins greenish to hyaline, ciliate proximally, denticulate distally, apices long-acuminate to aristate; strobili quadrangular, 2–14 mm long; sporophylls monomorphic, ventral ones sometimes less chlorophyllous; megasporangia mostly basal on two ventral rows; microsporangia in two dorsal rows and distally on ventral rows; megaspores white to beige, rugose to smooth or reticulate with low ridges, 300–400 ␮m diam.; microspores orange.

SELAGINELLA RIBAE

591

yellow, with broad ridges, 320–370 ␮m in diam.; microspores redorange, with ridge-like papillae.

Distribution. Terrestrial or epipetric on shaded trail banks and road banks; 1200–2400 m. Mexico; Hond. Selected Specimens Examined. Chis (Breedlove 8969, US, 21115, ENCB, TEX; Me´ndez Ton 5310, ARIZ, ENCB, MO). Gro (Zamudio et al. 9164, NY). Hgo (Luna et al. 633, NY). Oax (Garcı´a Mendoza & Torres 1524 NY; Hellwig 1326, LL, US). Ver (Conzatti 807, US; Va´squez V-1862, ENCB).

Selaginella pulcherrima is characterized by having monomorphic leaves along the stems below the first to fifth well developed branches. It is closely related to S. harrisii. Luna et al. 633 is unusual by having reticulate megaspores. Selaginella pulcherrima is further characterized by median leaves on main stems with narrowly hyaline margins 1–2 cells wide and long-acuminate to short-aristate apices. The acuminate tips or aristae are green at the center, narrowly hyaline at margins, and less than 1⁄3 the laminar length. These characters separate S. pulcherrima from S. polyptera, which see for further comparison.

59. SELAGINELLA REFLEXA Selaginella reflexa Underw., Bull. Torrey Bot. Club 21: 268. 1894. Lectotype (chosen by Mickel, 1992: 425). Mexico. Jalisco: Barranca of Guadaliajara, Pringle 2635 (NY!; isolectotypes BM, MEXU!, NY!, UC!, US!). Figs. 284H, J–N. Stems prostrate, long-trailing, stramineous to pale green, to 8 cm long, 0.1–0.3 mm diam., not articulate, not flagelliform, not stoloniferous, 2–3 times branched; rhizophores borne throughout stem length, filiform, ca. 0.1 mm in diam.; leaves of two kinds (anisophyllous) throughout stem length; lateral leaves often strongly incurved and forming narrow cylinders, ovate, 1–1.5 ⫻ 1–1.5 mm, bases rounded to cordate, margins white, apices obtuse, often with an apical mucro; median leaves ovate, bases cordate, margins hyaline, long-ciliate, sometimes denticulate distally, apices aristate, aristae about 1⁄2 the lamina length; axillary leaves oblong to oblanceolate, exauriculate, margins long-ciliate at bases, with ciliate-denticulate keels; strobili quadrangular, 2– 3 mm long; sporophylls monomorphic, ventral ones colorless; megasporangia basal in two ventral rows; microsporangia in dorsal two rows and distal in two ventral rows; megaspores lemon-

Distribution. Terrestrial or epipetric on shaded, damp rocks and steep banks in dry deciduous forests; 900–2000 m. Mexico; Guat. Selected Specimens Examined. Dgo (Breedlove 24519, CAS). Gro (Pringle 10326, F, MICH, UC, US; Rose 9391, US). Hgo (Chase 7199, UC; Hellwig 1237, MEXU). Jal (McVaugh 17348, CAS, MICH, NY). NL (Kenoyer 583, F). Qro (Carranza & Zamudio 5257, NY). SLP (Pringle 3292, F, LL, MEXU, MICH, NY, UC; Purpus 5488, F, UC, US).

Selaginella reflexa is related to S. schiedeana, which differs in having larger and more lanceolate lateral leaves (1.5–2 mm long) that remain flat and do not curve around the stem, short-ciliate median leaves with narrower white margins, and lateral leaves long-ciliate proximally.

60. SELAGINELLA RIBAE Selaginella ribae Valdespino, sp. nov. Type. Mexico. Guerrero: Mpio. Zupango del Rı´o, 38 km S of Iguala, ca. 0.5 km N de Xilitla [Xalita] along the road to road Acapulco, Koch et al. 7987 (holotype NY!; isotypes CAS!, MEXU!, NY!). Figs. 278A–E. Inter species rosulares Selaginellae, S. ribae apicibus foliorum lateralium et medianorum rotundatis vel late obtusis distincta. (Dedicated to the late Ramo´n Riba, dean of Mexican pteridology.)

Stems rosette-forming, green to reddish, to 18 cm long, 0.5– 1 mm diam., not articulate, not flagelliform, not stoloniferous, 2–3 times branched; rhizophores restricted to stem bases, stout, 0.3–0.5 mm diam.; leaves of two kinds (anisophyllous) throughout stem length, coriaceous; lateral leaves ascending, tightly imbricate, broadly ovate to ovate-deltate, 1.3–2.2 ⫻ 1.2–1.4 mm, bases rounded to cordate, margins hyaline, densely short-ciliate, apices rounded to broadly obtuse; median leaves similar to lateral leaves, tightly imbricate, ovate-deltate to suborbicular, 1–1.7 ⫻ 1–1.7 mm, bases rounded, margins hyaline, densely short-ciliate, apices rounded to broadly obtuse; axillary leaves similar to median and lateral leaves, bases cordate, exauriculate; strobili quadrangular, 4–8 mm long; sporophylls monomorphic (ventral ones sometimes less green); megasporangia mostly basal on two

592

SELAGINELLA

ventral rows; microsporangia in two dorsal rows and on ventral rows; megaspores lemon-yellow, echinate to striate, rugose, 300– 360 ␮m diam.; microspores orange.

tightly appressed and equally developed on all sides, imbricate, blue or gray-green, linear-lanceolate, margins short-ciliate, the cilia white, leaf apices aristate, aristae 0.2–0.5 mm, making a white tuft at the ends of sterile branches; strobili quadrangular, ca. 5 mm long; sporophylls monomorphic; megasporangia several at base of strobili; microsporangia distal to megasporangia; megaspores pale orange, rugulose-reticulate, (190–)240–270(–340) ␮m diam.; microspores bright orange.

Distribution. Terrestrial or epipetric on rocky hillsides, rocky soil, steep slopes; 600–950 m. Mexico. Specimens Examined. Dgo (Torke et al. 154, NY). Gro (Lorea 737, ENCB, MEXU; Thomas et al. 2865, NY). NL (Hinton 19975, MEXU).

Selaginella ribae is a member of the S. lepidophylla group and closely resembles S. lepidophylla and S. novoleonensis in its rosette habit. It can be separated from these by its round to broadly obtuse leaves. All three species have pubescent stems, which are more visible when the median leaves are lifted or removed. According to the information on the type, S. ribae grows on clay soils and forms dense rosettes mixed with Koch et al. 7989, the type of S. nothohybrida. Selaginella ribae can be easily distinguished from S. nothohybrida by its distally rounded lateral and median leaves (vs. long-cuspidate to aristate). In addition, S. ribae has the proximal face of the megaspores echinate (vs. reticulate-rugose).

Distribution. Terrestrial or epipetric on dry, scrubby, rocky hillsides; (200–)1450–2800 m. USA (Tex, N Mex, Ariz); Mexico. Selected Specimens Examined. Ags (Rzedowski & McVaugh 1250, ENCB) Chis (Knobloch 921, MSC). Coah (Villarreal et al. 3457, ARIZ). Dgo (Pennell 18178, PH). Gto (Rzedowski & McVaugh 1179, ENCB, MICH). Hgo (Arreguı´n 641, ENCB). Jal (McVaugh 17007, CAS). Me´x (Rzedowski 29876a, ENCB). Oax (Yatskievych & Wollenweber 83-141, ARIZ). Pue (Purpus 3156, BM, F, GH, MO, NY, US). SLP (Rzedowski 10745, ENCB). Son (Gentry 1194, UC). Tam (Bartlett 10876, DS, NY). Unverified, Doubtful, or Mistaken Reports.. Chih (Palmer 92, GH, MO, US, cited by R. Tryon, 1955, but not verified). DF (Schaffner 11, B, GH, US, cited by R. Tryon, 1955, but not verified).

61. SELAGINELLA RUPINCOLA Selaginella rupincola Underw., Bull. Torrey Bot. Club 25: 129. 1898. Figs. 266M–P. Bryodesma rupincola (Underw.) Soja´k, Preslia 64: 155. 1992. Lectotype (chosen by R. Tryon, 1955: 20). U.S.A. New Mexico: Dona Ana Co., Organ Mts., Wooton 124 (NY!; isolectotypes K, MO, US!). Selaginella chrismarii Hieron., Hedwigia 39: 299. 1900. Syntypes. Mexico. Near Guanajuato, von Chrismar s.n. (B, frags. NY!); Purpus 3156 (F, GH, MO, US!). Selaginella chrismarii Hieron. var. karwinskyana Hieron., Hedwigia 39: 300. 1900. Lectotype (chosen by R. Tryon, 1955: 20). Mexico. Near Durango, 1896, Palmer 554 p.p. (B; isolectotypes NY!–3 sheets, UC!, US!). Selaginella chrismarii Hieron. var. neeana Hieron., Hedwigia 39: 300. 1900. Type. Mexico. Ne´e s.n. (B).

Stems suberect to erect, rarely decumbent, to 10 cm, radially symmetric, somewhat flexuous, branched, secondary branches short, generally less than 1 cm long, not articulate, not flagelliform, not stoloniferous; rhizophores borne mostly only near stem bases; leaves of one kind (isophyllous) throughout stem length,

62. SELAGINELLA RZEDOWSKII Selaginella rzedowskii Lorea-Hern., Bol. Soc. Bot. Me´xico 44: 24. 1983. Type. Mexico. Guerrero: Zumpango del Rı´o, 2 km al ENE de Xochipala, Lorea 834 (FCME; isotype FCME). Figs. 280G, H, J–M. Stems creeping, brown in older portions and green in younger tips and branches, 4–20 cm long, 0.4–0.7 mm diam., not articulate, not flagelliform, not stoloniferous, 2–3 times branched; rhizophores borne throughout stem length, dark brown, 0.2–0.3 mm diam.; leaves of two kinds (anisophyllous) throughout stem length, lustrous; lateral leaves ovate to ovate-oblong, 1.8–2.7 ⫻ 0.8–1.3 mm, bases rounded, strongly overlapping stems, margins obscurely differentiated, ciliate proximally, denticulate or entire distally, apices obtuse, surfaces glabrous, without idioblasts; median leaves lanceolate, 1.5–2.1 ⫻ (0.6–)0.7–0.9 mm, bases rounded, margins obscurely differentiated to green, short-ciliate

SELAGINELLA SCHAFFNERI

proximally, denticulate or entire distally, apices acute to obtuse; axillary leaves similar to lateral ones, bases cordate, exauriculate; strobili quadrangular, (4–)7–13(–17) mm long; sporophylls monomorphic; megasporangia in two ventral rows; microsprorangia in two dorsal rows; megaspores lemon-yellow, striate to reticulate with high muri, 400–470 ␮m; microspores pale orange to deep yellow, shed in tetrads, tetrads ca. 50 ␮m diam.

Distribution. Terrestrial or epipetric on limestone rocks in rocky soil in seasonally dry forests; 450–1000 m. Mexico. Specimens Examined. Gro (Barkley & D. J. Carr 36145, GH; Contreras 338, XAL, 609, NY; Frye & Frye 2578, NY, UC).

Selaginella rzedowskii is distinct from S. flexuosa by the former having median leaves acute with ciliate bases (vs. long-aristate with glabrous bases), rhizophores dark brown and borne on the upper side of stems (vs. stramineous to green and borne on the lower side of stems), and megaspores pale orange to deep yellow (vs. white).

63. SELAGINELLA SARTORII Selaginella sartorii Hieron., Hedwigia 39: 304. 1900. Figs. 275N–P. Selaginella rupestris (L.) Spring var. sartorii (Hieron.) Frye, Ferns Northwest 31. 1934. Bryodesma sartorii (Hieron.) Soja´k, Preslia 64: 155. 1992. Lectotype (chosen by Weatherby, J. Arnold Arbor. 25: 418. 1944). Mexico. Veracruz: Mirador, Sartorius s.n. (B!, frag. NY!). Selaginella porrecta Weath., J. Arnold Arbor. 25: 416. 1944. Type. Mexico. Nuevo Leo´n: “Alamar,” Pablillo, SE of Galena, Pennell 17198 (US!, photo UC!). Selaginella hintonii Weath., J. Arnold Arbor. 25: 418. 1944. Type. Mexico. Me´xico: Dist. Temascaltepec, Ypericones, Hinton 8423 (GH, frag. US; isotypes ENCB, NY!, UC!, photo UC!).

Stems prostrate, branches 2–10 mm apart, ultimate branches short, 3–8 mm long, ascending, dorsiventral, not articulate, not flagelliform, not stoloniferous; rhizophores borne throughout stem length; leaves of one kind (isophyllous) throughout stem length; blue- to gray-green, often some reddish, appressed to slightly spreading, imbricate, covering leaf bases and stems, linear-lanceolate, bases pubescent, margins ciliate, cilia white, usually fewer than 15 per side, leaf apices aristate, aristae white,

593

0.8–1 mm long; strobili short and nearly round, subquadrangular to quadrangular, 8–10 mm long; sporophylls monomorphic; megasporangia several at base of strobili; microsporangia distal to megasporangia; megaspores yellow to orange-yellow, rugosereticulate, 150–220 ␮m diam.; microspores reddish orange to orange.

Distribution. Terrestrial or epipetric on dry, rocky slopes, often with cacti, Agave, Acacia; 1500–2350 m. Mexico; Col, Ven. Selected Specimens Examined. Me´x (type of S. hintonii).NL (only the type of S. porrecta). Oax (Iltis et al. 1185, NY; Mickel 810, ISC, 4075, 4483, 4496, 4498, 4953, 4961, 6489, 6654, NY). Ver (Purpus 8464, UC). Unverified, Doubtful, or Mistaken Reports. Dgo (Ortega 5319, US, cited by R. Tryon, 1955, but not verified). NL (Orcutt 1098, US, cited by R. Tryon, 1955, but not verified). SLP (Pennell 17753, US, cited by R. Tryon, 1955, but not verified). Tam (Bartlett 10355, US, cited by R. Tryon, 1955, but not verified).

R. Tryon (1955) reported that S. sartorii and S. sellowii are not well separated morphologically. The principal character of piliform pubescence on leaf bases is best seen on leaves at branching points. The less imbricate leaves of S. sellowii appear to correlate with the lack of pubescence and shorter setae on leaf tips. Tryon also synonymized S. porrecta and S. hintonii, which Weatherby (1944) distinguished on the basis of minor spore characters.

64. SELAGINELLA SCHAFFNERI Selaginella schaffneri Hieron. in Engler & Prantl, Nat. Pflanzenfam. 1(4): 674. 1901. Lectotype (chosen by Alston, 1952: 240, and again by Mickel 1992: 427, neither author fixing the location of the lectotype. We here designate the specimen at B as lectotype). Mexico. San Luis Potosı´: San Rafael Mountains, 1879 (not 1878, as in publ.), Schaffner 8 (B!–image seen; isolectotypes BM, NY!, US!). Figs. 278J–R. Stems long-creeping, rooting at intervals of 10–15 cm, forming very loose mats, 0.4–0.6 mm diam., erect stems 5–8 cm tall produced at intervals of 1–3 cm, apices curling when dry, graytan, not articulate, not flagelliform, not stoloniferous, 2–3 times branched; leaves of two kinds (anisophyllous) on erect stems, monomorphic on arching stems, buff, coriaceous, tightly ap-

594

SELAGINELLA

pressed to wiry, brownish stems that are often visible between the leaves, leaves on the erect portion grading to grayish green distally, spreading, tightly imbricate, neither stems nor leaf bases visible; lateral and median leaves similar in size, shape, and texture, ovate, 1–1.5 mm long, apices rounded or cuspidate, peltately attached, each with downward-pointing, erose auricle, margins otherwise entire, acroscopic margins thin, subscarious; axillary leaves oblanceolate, apices rounded to cuspidate, peltately attached, with ciliate to erose auricles; strobili quadrangular, 2–5 mm long; sporophylls monomorphic; megaspores white to cream, with low, unequal, sparingly anastomosing ridges; microspores orange to orange-tan, with heavy broad ridges.

Distribution. Terrestrial or epipetric on rocks, in tropical deciduous forests, sometimes in oak-pine forests; 1100–2100 m. Mexico. Specimens Examined. Gro (Kenoyer 27, US). Jal (Pringle 2594, US, 15630, CAS, UC, US). Me´x (Hinton 8858, DS, NY, UC, US). Mor (Matuda 26349, UC, US). Nay (Breedlove 45441, CAS; Pringle 2594, NY). SLP (Rzedowski 11370, NY).

Selaginella schaffneri is distinguished by the peltate attachment of leaves and sporophylls, all of which have branched veins that occasionally anastomose. As pointed out by Wagner et al. (1982), leaves with complex venation are a radical departure from the usual condition (a single, unbranched vein) in the entire lycopod evolutionary line (including also Isoe¨tes). Unfortunately, because of the thick and opaque nature of the leaves, the complex veins are not visible unless the leaves are cleared and the vascular tissue stained. The name S. saccharata has been mistakenly applied to plants of S. schaffneri, but that name is a synonym of S. delicatissima.

65. SELAGINELLA SCHIEDEANA Selaginella schiedeana A. Braun, Index Sem. Hort. Berol. App. 14. 1857 [1858]; Ann. Sci. Nat. Bot., se´r. 4, 13: 62. 1860. Type. Mexico. Veracruz: Papantla, Schiede & Deppe [827] (B!; isotypes BM, LE!). Figs. 271CC–HH. Stems prostrate, becoming long-trailing, stramineous to pale green, to 8 cm long, ca. 0.3 mm diam., not articulate, not fla-

gelliform, not stoloniferous, 2–3 times branched; rhizophores borne throughout stem length, filiform, ca. 0.1 mm diam.; leaves of two kinds (anisophyllous) throughout stem length; lateral leaves ovate, less than 2 mm long, bases rounded, with a few long cilia at base, otherwise finely denticulate, apices acute; median leaves ovate, bases rounded, margins long-ciliate to tips, with narrow white borders, apices long-acuminate; axillary leaves oblong, exauriculate, with a few long cilia at bases, denticulate apically, apices acute; strobili quadrangular, 2–3 mm long; sporophylls monomorphic and of two colors, dorsal rows green, ventral rows colorless; megasporangia in two ventral rows; microsporangia in two dorsal rows; megaspores lemon-yellow, with low reticulum, 250–300 ␮m diam.; microspores red-orange, with low broad papillae of irregular size and shape.

Distribution. Terrestrial in wet lowland and lower montane forests on Atlantic slopes; 0–1100 m. Mexico. Selected Specimens Examined. Chis (Martı´nez et al. 25130, MEXU). Me´x (Abundiz Bonilla 299, ENCB). Oax (Mickel 1480, 5820, NY, UC). SLP (Fryxell & Magill 2281a, MEXU; Rzedowski 10475, ENCB). Ver (Copeland herb. 167, MEXU, UC; Nee & Hansen 18408, F, MEXU; Purpus 2938, F, UC).

The combination of creeping habit, ciliate median leaves, proximally ciliate axillary leaves, lateral leaves with glabrous surfaces, short-ciliate sporophylls, each with a denticulate midrib, and lemon-yellow megaspores distinguish S. schiedeana from similar species (such as S. mixteca) with prostrate habit and denticulate median leaves.

66. SELAGINELLA SCHIZOBASIS Selaginella schizobasis Baker, J. Bot. 21: 333. 1883. Type. Mexico. Chiapas: Ghiesbreght 605 (K!; isotype NY!). Figs. 277M–O. Stems prostrate (to suberect), greenish to stramineous, 30–50 cm long, ca. 1–2 mm diam., articulate, not flagelliform, not stoloniferous, 2–4 times branched; rhizophores borne throughout stem length, stout, straight, 0.3–0.6 mm diam.; leaves of two kinds (anisophyllous) throughout stem length; lateral leaves obliquely oblong, 3–4 ⫻ 1.3–1.7 mm, acroscopic bases auriculate,

SELAGINELLA SERTATA

the auricles curved and hooked over stems, basiscopic margins truncate to short-auriculate, margins short-ciliate at base of plants, otherwise minutely denticulate, apices acutes; median leaves ovate-lanceolate, bases peltate with one auricle, margins short-ciliate on auricles, otherwise denticulate, apices acuminate to short-aristate; axillary leaves linear-lanceolate, bases with two divergent, ciliate auricles, margins minutely denticulate; strobili quadrangular, 6–14 mm long; sporophylls monomorphic, 0.6–3 cm long; megasporangia single at base of strobili; microsporangia in most of strobilus; megaspores white, reticulate, ca. 500 ␮m diam.; microspores cream, echinate.

595

length, blue- to gray-green, slightly spreading (the leaf bases and stems often visible), linear-lanceolate, margins ciliatedenticulate, apices aristate, aristae white, 0.5–0.8 mm long; strobili quadrangular, 2–3 mm long, obscure; sporophylls monomorphic; megasporangia at base of strobili; microsporangia distal to megasporangia; megaspores yellow to yellow-orange, lowreticulate; microspores orange.

Distribution. Terrestrial or epipetric in dry, scrubby arroyos; 1650–1700 m. Mexico; Cuba; Col, Ven, Ec, Peru, Braz, Bol, Parag, Arg, Uru. Distribution. Terrestrial in montane rain forests on Atlantic slope; 0–1300 m. Mexico; Guat, Bel, Hond, Nic. Selected Specimens Examined. Chis (Breedlove 22444, NY, TEX; Matuda 3676, LL, NY). Oax (Hallberg 1662, NY; Martı´nez Caldero´n 116, MEXU, NY; Mickel 1786, NY). Tab (Gilly & Herna´ndez 76, MSC, TEX; Rovirosa 539, NY). Ver (Copeland herb. 170, MEXU, MSC, NY; Nee et al. 25110, NY; Rzedowski 20352, NY, TEX).

Selaginella schizobasis, S. eurynota, and S. sertata (series Articulatae) form a poorly resolved complex with peltate median leaves. Selaginella schizobasis bears rhizophores along most of its stem length and occasionally has flagelliform stem apices. It is distinguishable by its oblong-elliptic lateral leaves, which are widest at middle, and by its axillary leaves with divergent auricles.

67. SELAGINELLA SELLOWII Selaginella sellowii Hieron., Hedwigia 39: 306. 1900. Figs. 275Q, R. Bryodesma sellowii (Hieron.) Soja´k, Preslia 64: 155. 1992. Lectotype (chosen by R. Tryon, 1955: 34). Brazil. Praia de San Diego, 1821, Sellow s.n. (B, frag. NY!). Selaginella pauciciliata Hieron., Hedwigia 58: 284. 1916. Type. Cuba. Holguı´n: Mayari, Wright 20 [1820, fide R. Tryon 1955: 34] (B; frag. NY!).

Stems prostrate, 0.5–0.6 mm diam., not articulate, not flagelliform, not stoloniferous, not much branched, not strongly dorsiventral, branches thin, sparse, 5–10 mm apart, ultimate branches short, 3–10 mm long; rhizophores borne throughout stem length; leaves of one kind (isophyllous) throughout stem

Specimens Examined. Gto (Rzedowski 47120, NY p.p.). Me´x (Abundiz-Bonilla 315, IZTA). Oax (Mickel 6654b, NY). Pue (Arse`ne 891, F, UC, US; Nicolas 10053, UC). Unverified, Doubtful, or Mistaken Reports. DF (Rose & Painter 6835, US, cited by R. Tryon, 1955, but not verified). Ver (Liebmann 2062, US, cited by R. Tryon, 1955, but not verified).

Selaginella sellowii is distinguished from S. wrightii and S. landii by its glabrous leaf bases and long white awns, and from S. sartorii by the less imbricate, non-ciliate leaves.

68. SELAGINELLA SERTATA Selaginella sertata Spring, Nouv. Me´m. Acad. Roy. Sci. Bruxelles 24(1): 104. 1850. Type. “Panama” [Costa Rica, Nicoya]. Sinclair [49] (isotype K!, frag. B!). Figs. 277A–H, J–L. Selaginella nicaraguensis Baker, J. Bot. 21: 333. 1883. Type. “Guatemala, in forests of Grenada” [Nicaragua]. Levy 360 (K, frag. NY!).

Stems suberect, greenish to stramineous, generally 10–20 cm long, ca. 1 mm diam., articulate, flagelliform at apices, not stoloniferous; rhizophores in proximal 1⁄4 of plant and on flagelliform portion; leaves of two kinds (anisophyllous) above first stem branch, mostly 3 times branched; lateral leaves obliquely oblong, 3–4 mm long, bases truncate with central attachment, shortciliate at bases, denticulate, apices acute; median leaves obliquely ovate-lanceolate, bases with single peltate auricle, margins ciliate, apices long-acuminate; axillary leaves linear-oblong, truncate, ciliate in proximal half, denticulate distally; strobili quadrangular, 3–7 mm long; sporophylls monomorphic; megasporangia single at base of strobili; microsporangia in most of strobilus; megaspores

596

SELAGINELLA

gray, reticulate with low muri, ca. 500 ␮m diam.; microspores cream-colored, spiny.

lanceolate, bases with subequal auricles, margins minutely denticulate, apices long-acuminate to aristate; axillary leaves obovate-lanceolate, exauriculate, margins denticulate at bases and tips; strobili quadrangular, 5–15 mm long, with megasporophylls as large as lateral leaves, microsporophylls as small as median leaves; megasporangia single, at base of strobili; microsporangia in most strobili; megaspores cream to beige, reticulate; microspores beige to pale orange, spinulose.

Distribution. Terrestrial in lowland wet forests along rivers and grassy roadsides; 0–1000 m. Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan. Selected Specimens Examined. Gro (Hinton et al. 11369, NY; Rzedowski 22808, NY). Jal (Castillo et al. 11940, NY; Iltis et al. 31021, NY). Mich (King & Soderstrom 4951, NY). Nay (Feddema 795, NY; McVaugh 19008, NY). Oax (Hellwig 1321, NY; Mickel 5125, NY, UC; Rzedowski 31333, NY). Unverified, Doubtful, or Mistaken Reports. Camp (Chavelas et al. ES⫽357, ENCB, cited by Palacios-Rios, 2002, but not verified, doubtful).

Selaginella sertata frequently has main stems arising vegetatively from flagelliform branches. In those “juvenile” stems, it is common to find axillary leaves with two prominent, ciliate, overlapping auricles, as well as biauriculate lateral leaves with acroscopic auricles prominent and curving downward along the stems. Specimens collected in this stage tend to be confused with mature specimens of S. eurynota. Selaginella sertata can be identified in the juvenile stage from S. eurynota by its conspicuously crowded (vs. distant) leaves at the stem bases which are rooted only at the base (vs. throughout). Selaginella sertata differs from S. eurynota and S. schizobasis in its fully developed stage by the characters given in the key. Additional discussions are under S. eurynota and S. schizobasis.

69. SELAGINELLA SILVESTRIS Selaginella silvestris Asplund, Ark. Bot. 20A(7): 30, f. 3–5. 1926. Type. Bolivia. Sur Yungas: Ad El Chaco, Asplund 1140 (S). Figs. 270L–P. Stems suberect to trailing, stramineous, ca. 0.8–1.5 mm diam., articulate, not flagelliform, not stoloniferous, stramineous, 2–3 times branched; rhizophores borne throughout stem length, ca. 0.5 mm diam., leaves of two kinds (anisophyllous) throughout stem length; lateral leaves obliquely oblong, to 3.5 mm long, bases truncate, rounded acroscopically, margins denticulate proximally, entire distally, apices obtuse; median leaves ovate-

Distribution. Terrestrial in wet montane forests and cloud forests; (0–)1700–2800 m. Mexico; Guat, Bel, Hond, Salv, Nic, CR; Col, Ven, Ec, Peru, Bol. Selected Specimens Examined. Chis (Breedlove 15240, DS, US; Matuda 6023, LL, US). Hgo (Knobloch 668, UC; Moore 3213, UC, US). Me´x (Hinton et al. 7350, US, LL). Oax (Martı´nez C. 332, NY, UC, US; Mickel 3793, NY, UC). Ver (Balls 4737, UC, US; Copeland herb. 168, UC, US; Nee & Diggs 24910, CAS, LL).

The denticulate lateral and axillary leaves, both lacking auricles, and denticulate median leaves with small, subequal auricles distinguish S. silvestris from the two other articulate species (S. stellata and S. tarda) with non-peltate median leaves. Selaginella stellata has ciliate lateral and axillary leaves, both with auricles, and ciliate median leaves with one auricle larger than the other; S. tarda has median leaves with one large auricle. The other articulate species in Mexico, S. schizobasis, S. eurynota, and S. sertata, all have peltate median leaves. Most specimens in Mexico are from higher elevations than other articulate species (1700–2800 m). Two collections from Oaxaca (100 m, Mickel 6828, NY; 20 m, Martı´nez-C. 332, NY, UC, US) are from low elevations and appear similar to S. silvestris in most respects. However, they have peculiar branches, especially near the plant base, which lack lateral leaves, thus appearing flagelliform. It is not clear whether this is a genetic feature or the action of an herbivore.

70. SELAGINELLA SIMPLEX Selaginella simplex Baker, J. Bot. 23: 293. 1885. Type. Brazil. Para´: near Santare´m, Spruce 947 (K!; isotypes BM!, CGE, W!) Figs. 286A–F.

SELAGINELLA STENOPHYLLA

Stems ascending to suberect, stramineous to greenish, 0.6–5 cm long, 0.1–0.3 mm diam., not articulate, not flagelliform, not stoloniferous, 1–2 times branched; rhizophores restricted to the proximal 1⁄3 of stems, filiform, ca. 0.1 mm diam.; leaves of two kinds (anisophyllous) throughout stem length; lateral leaves ovate to ovate-elliptic, 1.3–1.6 ⫻ 0.8–1 mm, bases rounded, the acroscopic margins greenish, short-ciliate proximally, denticulate distally, apices acute to acuminate, both surfaces glabrous, without idioblasts; median leaves lanceolate to ovate-lanceolate, 0.6– 1.2 ⫻ 0.2–0.4 mm, bases rounded, margins greenish, denticulate, apices long-acuminate to aristate, aristae 1⁄3–1⁄2 the lamina length, 0.2–0.3 mm, both surfaces glabrous, without conspicuous idioblasts; axillary leaves similar to lateral leaves, bases rounded, exauriculate; strobili lax, flattened, dorsiventral, 1.8–6.4 mm long; sporophylls dimorphic, the dorsal ones green, spreading, the ventral ones hyaline, ascending; megasporangia in two ventral rows; microsporangia usually in two dorsal rows, or dorsal sporophylls sterile; megaspores pale yellow, frequently with an equatorial flange, the proximal face granulose-tuberculate or rugulate, the distal face reticulate with open to closed reticulum of low ridges, 260–300 ␮m diam.; microspores orange.

Distribution. Terrestrial or epipetric in wet ravines on shaded and wet rock ledges, on dry and open river banks, and on banks in hillside savannas; 600–1000 m. Mexico; CR; Ven, Trin, Braz, Bol. Specimens Examined. Gro (Lorea 4808, NY). Nay (Feddema 2660, MICH). Oax (Tryon & Tryon 7389, GH p.p.; Mickel 3996a, NY).

Selaginella simplex is very similar in size and general morphology to S. minima, and in the past they have been confused (Valdespino, 1993). Selaginella simplex can be distinguished from S. minima by having greenish (vs. hyaline) leaf and sporophyll margins, upper surfaces of lateral leaves and sporophylls without (vs. frequently with) conspicuous stomata and short hairs, and pale yellow (not white) megaspores with the proximal face granulate-tuberculate (not striate, reticulate, or rugulate).

71. SELAGINELLA STELLATA Selaginella stellata Spring, Flora 21: 194. 1838. Type. Mexico. Schiede s.n. (B). Figs. 270Q–U.

597

Selaginella galeottii Spring, Bull. Acad. Roy. Sci. Bruxelles 10(1): 230.1843. Lectotype (chosen by Alston, 1955: 258). Mexico. Veracruz: Jalapa, Mirador, Galeotti 6606 (LG!; isolectotypes BR!, photo B!, K!, P, photo P!, photo BM); the other syntype is from Bolivia, d’Orbigny s.n. and is S. diffusa (C. Presl) Spring.

Stems erect to suberect, stramineous, 3 mm diam., articulate, not flagelliform, not stoloniferous, 2–3 times branched; rhizophores borne in proximal third of stems, straight, ca. 0.4–0.5 mm diam.; leaves of two kinds (anisophyllous) throughout stem length; lateral leaves ovate to oblong-ovate, to 4.5 mm long on main stems, 3–4 mm long on branches, bases rounded, with a few long cilia at bases, margins otherwise entire, apices acute; median leaves ovate-lanceolate, acuminate, bases subequally auriculate, with a few cilia on the auricles, margins denticulate to entire; axillary leaves linear-oblong, slightly auriculate, with 1 or 2 hairs on each auricle; strobili quadrangular, 0.5–2.2 cm long; sporophylls monomorphic; megasporangia single at base of strobili; microsporangia in most of strobilus; megaspores white or light brown, with high, white reticulum, ca. 350 ␮m diam.; microspores cream to pale beige, with closely set spines.

Distribution. Terrestrial in montane rain forests; 300–2200 m. Mexico; Guat, Bel, Hond, CR, Pan. Selected Specimens Examined. Chis (Breedlove 24068, NY). Gro (Hinton et al. 11369, UC). Hgo (Gimate s.n., NY). Jal (Iltis et al. 317, UC). Mich (King & Soderstrom 4951, UC). Nay (Kimnach 249, UC). Oax (Martı´nez C. 494, ENCB, MEXU, NY). Pue (Rzedowski 23389, NY). Ver (Pringle 6107, NY; Purpus 8922, NY).

See S. silvestris for a comparison.

72. SELAGINELLA STENOPHYLLA Selaginella stenophylla A. Braun, Index Sem. Hort. Berol. App. 22. 1857. Lectotype (chosen here). Cultivated material at Horto Botanico Berolinensi, 1857, Braun s.n. (B!; isolectotypes B!, M!). Figs. 285T–Y. Selaginella incurvata Baker, J. Bot. 21: 99. 1883. Type. Mexico. Veracruz: Liebmann s.n. (K; isotype frag. NY!). Selaginella miradorensis Hieron., Hedwigia 43: 30. 1904. Lectotype (chosen here). Mexico. San Luis Potosı´: Tamasopo Canyon, Pringle 3293 (B!; isolectotypes BM!, BR!, CAS!, DS!, K!, L!, M!, MICH!, NY!, P!, UC!, US!, W!).

598

SELAGINELLA

Stems creeping, greenish or stramineous, to ca. 35 cm long, 0.5–1.4 mm diam., not articulate, not flagelliform, not stoloniferous, 3–4 times branched; rhizophores restricted to proximal half or throughout stems, stout, 0.5–0.8 mm diam.; leaves of two kinds (anisophyllous) throughout stem length; lateral leaves oblong to oblong-ovate, (1.5–)2–3.7 ⫻ (0.6–)1–1.5 mm, bases cordate, margins hyaline, acroscopic margins finely denticulate to entire, apices broadly acute to rounded, both surfaces glabrous, with conspicuous or obscure idioblasts; median leaves elliptic or elliptic-ovate, 1.3–2 ⫻ 0.6–1.2 mm, bases biauriculate, margins hyaline, long-ciliate along proximal half, especially along the inner margin, short-ciliate to denticulate distally, apices abruptly aristate, aristae 1⁄4 to 1⁄2 the lamina length, 0.3–0.8 mm long, both surfaces glabrous, upper surfaces with conspicuous idioblasts; axillary leaves lanceolate to ovate-lanceolate, bases cordate; strobili lax and flattened, dorsiventral, 3–10 mm long; sporophylls dimorphic, the dorsal ones green, spreading, the ventral ones hyaline to pale green, ascending; megasporangia in two ventral rows; microsporangia in two dorsal rows; megaspores white, infrequently cream to brownish, granular-striate or striate-reticulate, with low to high reticulum, 330–356 ␮m diam.; microspores orange.

Bryodesma steyermarkii (Alston) Soja´k, Preslia 64: 155. 1992. Type. Guatemala. Huehuetenango: between San Sebastia´n H. and large pen˜asco above town, Steyermark 50501 (BM; isotype F!).

Stems prostrate, creeping, forming loose mats to 30 cm across, radially symmetric, 0.3–0.6 mm diam., not articulate, not flagelliform, not stoloniferous, 3–4 times branched; rhizophores borne throughout stem length, 0.2–0.4 mm diam.; leaves of one kind (isophyllous) throughout stem length, light green with reddish spots or totally reddish when old, ascending, linearlanceolate, 1.3–2.5 ⫻ 0.2–0.3 mm, bases truncate to slightly subcordate, abruptly adnate to stems, glabrous or puberulent, margins greenish, long-ciliate, short-ciliate to denticulate distally, cilia and teeth hyaline, apices long-aristate, aristae hyaline, 0.5–0.8 mm long; strobili quadrangular, 0.4–1.2 cm long; sporophylls monomorphic; megasporangia few, scattered in ventral rows; microsporangia in two dorsal rows and scattered in ventral rows; megaspores yellow, rugose to reticulate, with a well defined equatorial flange; microspores orange.

Distribution. Terrestrial or epipetric on rocky soil and rock faces; 960–1700 m. Mexico; Guat.

Distribution. Terrestrial or epipetric in shaded, damp areas, on limestone slopes, and on mossy rocks; 150–1300 m. Mexico; Guat. Selected Specimens Examined. Chis (Ishiki et al. 2025, NY). Hgo (Wiggins 13336, DS). NL (Wright s.n., PH). Pue (Fro¨derstro¨m & Hulte´n 750, NY). Qro (Rzedowski 43867, NY). SLP (Pennell 17995, PH). Tam (Nee & Diggs 24511, NY). Ver (Purpus 128, PH).

The very long, creeping stems, mostly oblong lateral leaves with broadly acute to rounded apices, median leaves each with a well developed outer auricle, and stout rhizophores 0.5–0.8 mm distinguish S. stenophylla from all other Mexican species.

73. SELAGINELLA STEYERMARKII Selaginella steyermarkii Alston, Ann. Mag. Nat. Hist., ser. 12, 7: 638. 1954. Figs. 275J–M.

Specimens Examined. Chis (Breedlove 40588, ENCB, MICH, NY, TEX, 41805, CAS, 46325, CAS, ENCB, NY; Gittins 4251, 4252, NY, Matuda 1846, MICH; Stevens & Martı´nez S. 25724, NY, UC).

Selaginella steyermarkii is easily distinguished from most isophyllous species, except S. sartorii, by having leaves green with reddish spots or totally red when old and by its long-attenuate sporophylls. Selaginella steyermarkii may be confused with some specimens of S. sartorii from Tamaulipas, which occasionally have reddish leaves. Additional studies are needed to determine the relationship between the two species.

74. SELAGINELLA SUBRUGOSA Selaginella subrugosa Mickel & Beitel, Mem. New York Bot. Gard. 46: 352. 1988. Type. Mexico. Oaxaca: Distrito Ixtla´n, 23 km S of Valle Nacional, 5 km N of Vista Hermosa, Mickel 6686 (NY!; isotype UC!). Figs. 283U–X. Stems prostrate, stramineous to pale green, 5–12 cm long, 0.6– 0.8 mm diam., not articulate, not flagelliform, not stoloniferous,

SELAGINELLA TARDA

2–3 times branched; rhizophores borne throughout stem length, stout, ca. 0.3 mm diam.; leaves of two kinds (anisophyllous) throughout stem length; lateral leaves on main stems oblong, 1.8–2.5 mm long, bases rounded, margins minutely denticulate, apices acute, sometimes with short idioblasts below, verruculose on upper surfaces; median leaves ovate, bases rounded, margins white and denticulate, surfaces rugulose, apices short-aristate, aristae 0.3–0.5 mm long, 1⁄4 the lamina length; axillary leaves ovate to lanceolate, exauriculate, denticulate at bases; strobili quadrangular, 8–15 mm long; sporophylls monomorphic and of two colors, dorsal rows green, ventral rows pale green; megasporangia in two ventral rows, smaller strobili often completely microsporangiate; microsporangia in two dorsal rows; megaspores white, with faint markings, 250–350 ␮m diam.; microspores light orange, with slender, often capitate papillae.

Distribution. Terrestrial in wet montane forests; 800–1600 m. Mexico. Selected Specimens Examined. Oax (Mickel 1579, NY, US, 1589, 4795, NY, UC, 5682, NY, 5766, 5933, 7084, 7128, NY, UC; Rzedowski 33391, ENCB, 34057, IEB).

Selaginella subrugosa is intermediate between S. flexuosa (q.v.) and S. corrugis, and differs from the former in having thinner rhizophores (0.3 mm diam.), suborbicular median leaves with short aristae (0.3–0.5 mm) and somewhat verruculose upper surfaces, and lateral leaves smaller (1.8–2.5 mm long) and more ovate. Selaginella corrugis has thinner rhizophores, smaller, orbicular, verruculose, short-cuspidate median leaves, and ovate lateral leaves only 1–1.2 mm long. Selaginella subrugosa is known only in Oaxaca, where it grows with S. flexuosa (Mickel 5932, 7083, NY).

75. SELAGINELLA TARDA Selaginella tarda Mickel & Beitel, Mem. New York Bot. Gard. 46: 353. 1988. Type. Mexico. Oaxaca: Distrito Putla, 61 km S of Putla, 12 km S of Zacatepec, Mickel 6927 (NY!). Figs. 272H–N.

599

Stems suberect to erect, stramineous, 8–15 cm long, ca. 0.5– 1.5 mm diam., articulate, not flagelliform, not stoloniferous, 2 times branched; rhizophores limited to proximal 1⁄3 of plant, stout, 0.5–0.8 mm diam.; leaves of two kinds (anisophyllous) throughout stem length; lateral leaves narrowly oblong, 1.5–2 mm long, bases truncate to subcordate basiscopically, cuneate acroscopically, margins finely denticulate, with narrow white borders, apices acute; median leaves ovate-lanceolate, nonpeltate, bases with one large auricle, borders narrow, white, margins finely denticulate, apices short-aristate; axillary leaves linear-lanceolate, exauriculate, margins denticulate; strobili quadrangular, 2–4 mm long; sporophylls dimorphic, the dorsal ones green, spreading, the ventral ones colorless, ascending, megasporophylls as large as lateral leaves, microsporophylls as small as median leaves; megasporangia single at base of strobili; microsporangia throughout strobili; megaspores cream to beige, 410–550 ␮m diam.; microspores beige.

Distribution. Terrestrial in wet forests; 300–1400 m. Mexico. Selected Specimens Examined. Col (Orcutt 4690, US p.p.). Dgo (Ortega 4287, US). Gro (Lorea 2842, NY). Jal (Lo´pez 21289, NY). Me´x (Hinton 2491, K, LL, PH, 8462, ENCB, F, LL, MICH, NY, PH, US). Mich (Dı´az Barriga & Garcı´a 6404, CAS; King & Soderstrom 4911, MICH, NY, TEX, US). Nay (Pennell 19966, PH). Oax (only the type). Sin (Breedlove 35658, CAS).

Selaginella tarda differs from S. sertata, S. schizobasis, and S. eurynota in smaller size, basally attached median leaves, denticulate leaf margins, and lack of flagelliform shoots. Among species with basally attached median leaves, S. silvestris has median leaves with two small, subequal auricles and exauriculate lateral leaves, S. stellata has auriculate, ciliate, lateral, axillary, and median leaves, whereas S. tarda has denticulate lateral leaves with truncate bases, denticulate median leaves with one large auricle (like S. stellata), and exauriculate, denticulate axillary leaves. See S. silvestris for comparison with that species. Selaginella tarda is closely related to S. kunzeana A. Braun, which is distributed from Costa Rica to Peru. In S. kunzeana, the median leaves have two auricles with the inner auricle narrower than the outer one; the leaves are denticulate, as in S. tarda, although the basiscopic margin of the lateral leaves is less toothed and somewhat curled.

600

SELAGINELLA

76. SELAGINELLA TENELLA Selaginella tenella (P. Beauv.) Spring, Bull. Acad. Roy. Sci. Bruxelles 10: 234. 1843. Figs. 287N–T. Diplostachium tenellum P. Beauv., Mag. Encycl. 9(5): 481. 1804. Lycopodium tenellum (P. Beauv.) Desv. ex Poir., Encycl. Suppl. 3: 553. 1814. Neotype (chosen by Proctor, 1985: 45). Puerto Rico. India occidentalis in Portorico ex Herb. Vent. [Ventenat] (P-Desv., photo BM!). Selaginella cladorrhizans A. Braun, Ann. Sci. Nat. Bot., se´r. 5, 3: 282. 1865. Syntypes. Venezuela. Aragua: Colonia Tovar, Fendler 324 p.p. (B, BM!, E!, F!, G!, K!, NY!, US!, YU!); Falco´n: San Carlos, Moritz 448 (B, BM!, K!). Selaginella subcaulescens Baker, J. Bot. 22: 277. 1884. Type. Cuba. Wright 1822 (K!; isotypes B!, BM!, G!, GH!, MO!, NY!, P!, S!, YU!).

Stems ascending, suberect, or erect, stramineous, 3–15 cm long, 0.4–1 mm diam., not articulate, the stem apices infrequently flagelliform, stoloniferous, 2–3 times branched, branches frequently flagelliform; rhizophores restricted to proximal 1⁄4 or throughout stems on flagelliform plants, filiform, 0.1–0.5 mm diam.; leaves of two kinds (anisophyllous) throughout stem length; lateral leaves ovate to oblong, 1.2–3 ⫻ 0.5–2.5 mm, bases rounded to subcordate, overlapping the stems acroscopically, acroscopic margins hyaline or pale green, short-ciliate to denticulate or entire, apices acute, both surfaces glabrous, with idioblasts; median leaves distant, broadly ovate-elliptic to lanceolate, 0.7–2 ⫻ 0.4–1.4 mm, bases cuneate to subcordate or with one small auricle, margins hyaline, denticulate, apices acuminate, long-acuminate, or long-aristate, aristae 1⁄2 or more the lamina length, 0.2–0.7 mm long, upper surfaces glabrous with idioblasts; axillary leaves similar to lateral leaves, bases rounded; strobili lax and flattened, dorsiventral, 2–9 mm long; sporophylls dimorphic, the dorsal ones green, spreading, the ventral ones hyaline to pale green, ascending; megasporangia generally in two ventral rows or at the base of ventral rows, or infrequently absent; microsporangia in two dorsal rows, frequently also in the distal part of ventral side of strobili, or strobili wholly microsporangiate; megaspores white, sometimes yellowish with age, proximal faces granular to granular-reticulate, with reticulum of low ridges, distal faces reticulate with low to high ridges, 230– 270 ␮m diam.; microspores tan to orange, echinulate to baculate, 24–31 ␮m diam.

Distribution. Terrestrial to epipetric in lowland and montane forests, on open to shady roadsides or stream banks, on sandy or clayey soil, dry or moist, mossy rocks, or rocky slopes; 30– 1000 m. Mexico; Guat, Bel, Hond, Pan; Gr Ant; Col, Ven, Trin, Tob, Braz. Selected Specimens Examined. Col (Orcutt 4640, BM, GH). Gro (Mexia 8725, BM, CAS, F, GH, K, MICH, MO, NY, UC, US, 8755, BM, F, GH, K, MO, NY, UC, US; Hinton 8502, MICH, NY, US, 8602, TEX). Jal (McVaugh & Koelz 1727, MICH, NY; Mexia 1276, BM, GH, MICH, MO, NY, UC, US). Nay (Castillo 6069, XAL). Sin (Mexia 2682, MO). Ver (Wendt et al. 3087, NY).

Selaginella tenella is defined here in a broad sense. It is characterized by having conspicuous idioblasts on the upper surface of the leaves and sporophylls. Traditionally, S. subcaulescens has been separated because of its suberect to erect stems, whereas S. tenella and S. cladorrhizans were thought to be mostly creeping or rarely ascending. Specimens of S. cladorrhizans from Belize and Guatemala, however, also have suberect and erect stems. A thorough examination of specimens of these three taxa throughout their ranges reveals no consistent character that separates them. Megaspores examined from S. tenella, S. subcaulescens, and S. cladorrhizans show only minor variations in exospore sculpturing and microstructure. Several additional names are synonymous with S. tenella. They include S. albonitens Spring, S. mollis Fe´e (non A. Braun), and S. sintenisii Hieron., all of the West Indies. Two other species, S. humilis Jenman and S. maracasensis O. C. Schmidt, are probably also conspecific with S. tenella; however, we have not found type material of either of these.

77. SELAGINELLA TUBEROSA Selaginella tuberosa McAlpin & Lellinger, Brenesia 24: 409. 1985 [1986]. Type. Costa Rica. Puntarenas: Las Alturas de Coto´n, McAlpin 2505 (US!; isotypes CR, DUKE, F!, NY!). Figs. 286N–S. Stems suberect to erect, stramineous, 10–30 cm long, 0.5–1.4 mm diam., not articulate, not flagelliform, stoloniferous with tuber-like buds at the tip, 2–3 times branched; rhizophores restricted to proximal 1⁄4 of stems, filiform, 0.1–0.3 mm diam.;

SELAGINELLA VIRIDISSIMA

leaves of two kinds (anisophyllous) throughout stem length; lateral leaves broadly ovate to oblong, 2.8–4.5 ⫻ 1.6–2.5 mm, bases rounded, slightly or not overlapping stems, margins greenish to narrowly hyaline, long-ciliate at bases, denticulate distally, apices acute to obtuse, both surfaces glabrous, without conspicuous idioblasts; median leaves distant, elliptic, 1.8–3.5 ⫻ 1.3–1.6 mm, bases biauriculate, auricles long-ciliate, margins narrowly hyaline, minutely denticulate, apices short- to long-aristate, aristae half or less the lamina length, 0.3–0.9 mm long, both surfaces glabrous, with conspicuous idioblasts above, without idioblasts below; axillary leaves similar to lateral leaves or more oblong, bases rounded; strobili lax and flattened, dorsiventral, 2–4 mm long; sporophylls dimorphic, the dorsal ones green, spreading, the ventral ones pale green, ascending; megasporangia single at base of strobili; microsporangia occupying almost the entire strobilus; megaspores white to bone-white, with low ridges; microspores not seen.

601

Preslia 64: 155. 1992. Lectotype (chosen by Weatherby, 1944: 412). U.S.A. New Mexico: Fendler 1024 (NY!; isolectotype NY!). Selaginella underwoodii Hieron.var. dolichotricha Weath., J. Arnold Arbor. 25: 412. 1944. Type. U.S.A. New Mexico: Mogollon creek, Metcalfe 276 (GH; isotype NY!).

Stems usually long-creeping, sometimes short-creeping or pendent, forming loose festoon-like mats or rarely compact mats, radially symmetrical, dry stems not fragile, branching anisotomously, lateral branches well spaced, spreading, not ascending, 1–2 times forked; rhizophores borne throughout stem length, 0.2–0.3 mm diam.; leaves of one kind (isophyllous) throughout stem length, loosely appressed, linear to linear-lanceolate, 2–3.4 ⫻ 0.4–0.7 mm, bases mostly decurrent and cuneate, rarely adnate and rounded (on young branches), pubescent or glabrous, apices carinate, slightly attenuate, short- to long-aristate, aristae hyaline-greenish to greenish yellow or rarely white, 0.2–1 mm, sometimes breaking off, margins entire to denticulate or very short-ciliate, cilia hyaline, scattered; strobili sometimes paired, quadrangular, 0.5–3.5 cm long; sporophylls monomorphic; megasporangia basal, strobili mostly microsporangiate; megaspores yellow, reticulate, 350–475 ␮m diam.; microspores orange, 35–45 ␮m diam.

Distribution. Terrestrial or epipetric on soil in coffee plantations, banks of waterways, road banks, or rocky slopes in premontane forests with a pronounced dry season; 1100–1900 m. Mexico; CR, and to be expected throughout Central America. Specimen Examined. Oax (Mickel 6193, NY).

Selaginella tuberosa is easily distinguished from related species by having lateral leaves with raised and frequently hyaline midribs and median leaves each with a well developed, long-ciliate inner auricle. McAlpin and Lellinger speculated that S. tuberosa was closely related to S. silvestris, a member of series Articulatae. However, S. tuberosa has axillary or ventral rhizophores and lacks stem articulations, as well as lacking other characteristics that define series Articulatae (see Valdespino, 1993, for discussion).

78. SELAGINELLA UNDERWOODII Selaginella underwoodii Hieron. in Engler & Prantl, Nat. Pflanzenfam. 1(4): 714. 1901. Figs. 276F–H. Selaginella rupestris (L.) Spring var. fendleri Underw., Bull. Torrey Bot. Club 25: 127. 1898. Selaginella fendleri (Underw.) Hieron., Hedwigia 39: 303. 1900, hom. illeg., non Baker, 1883. Bryodesma underwoodii (Hieron.) Soja´k,

Distribution. Epipetric on moist or shaded cliffs, rocky slopes, rock crevices, granitic outcrops, hanging over granite cliffs, sandstone ledges or limestone; 1400–3200 m. USA; Mexico. Specimens Examined. Chih (Correll & Gentry 23163, UC; Correll & Johnston 21778, NY). Dgo (Gonza´lez & Acevedo 2143, MEXU). Tam (Nesom 6016, TEX). Unverified, Doubtful, or Mistaken Reports. Ags (Siqueiros 4354, HUAA, cited by Siqueiros-Delgado & Gonza´lez-Adame, 2004, but not verified).

Selaginella underwoodii is a rather distinct species; however, it shows some variation in the number and length of the marginal cilia (from none to many), leaf base pubescence, and arista length.

79. SELAGINELLA VIRIDISSIMA Selaginella viridissima Weath., J. Arnold Arbor. 24: 326. 1943. Figs. 275E–H.

602

SELAGINELLA

Bryodesma viridissima (Weath.) Soja´k, Preslia 64: 155. 1992. Type. Mexico. Coahuila: Can˜o´n de Calabaza, Sierra Mojada, Stewart 2204 (GH; isotype F!). Selaginella coryi Weath., Amer. Fern J. 36: 51. 1946. Type. U.S.A. Texas: Brewster Co., Chisos Mountains, Cory 44831 (GH, frag. US).

Stems forming clumps or mounds, radially symmetric, underground (rhizomatous) and aerial, not readily fragmenting, irregularly forked, rhizomatous and aerial stems often with 1 branch arrested, bud-like, tips straight, aerial stems mainly erect, seldom merely ascending, with bud-like arrested branches throughout stem length; rhizophores restricted to rhizomatous stems and proximal 1⁄4 of aerial stems, 0.2–0.3 mm diam.; leaves of one kind (isophyllous) throughout stem length; rhizomatous stem leaves loosely appressed, straight, scale-like, decurrent or long-decurrent; aerial stem leaves appressed, green, linearlanceolate to narrowly lanceolate, 1.8–2.1 ⫻ 0.5–0.6 mm, bases cuneate and decurrent to slightly rounded and adnate, glabrous, margins denticulate to very short-ciliate, apices acute to obtuse; strobili quadrangular, 0.5–1.2(–2.5) cm long; sporophylls monomorphic; megasporangia and microsporangia each in two rows; megaspores pale orange, rugose, 400–500 ␮m; microspores orange.

Stems prostrate, forming loose to dense mats, dorsiventral, branch tips curving upward; rhizophores borne throughout stem length, 0.3–0.4 mm diam.; leaves of one kind (isophyllous) throughout stem length; leaves crowded, slightly dimorphic, margins ciliate, cilia hyaline, apices aristate, aristae yellowish, 0.2–0.5 mm, often breaking off, usually more persistent on underside leaves; upperside leaves linear-lanceolate, 3.3–3.9 ⫻ 0.6– 0.8 mm, bases abruptly adnate, pubescent; underside leaves narrowly linear-lanceolate to falcate on marginal rows, 3.5–4.5(–5) ⫻ 0.6–0.7 mm, bases abruptly adnate or slightly decurrent, usually pubescent, sometimes glabrous; strobili quadrangular, (0.7– )1.5–2 (–2.6) cm long; sporophylls monomorphic; megasporangia few, basal, strobili nearly entirely microsporangiate; megaspores pale to bright orange, rugose, 270–380 ␮m diam.; microspores bright orange, 34–56 ␮m diam.

Distribution. Epipetric on exposed or shaded rocks, limestone cliffs; 375–2000 m. USA; Mexico.

Distribution. Epipetric on shaded cliffs, slopes, rock crevices, and igneous rocks; 1300–1850 m. USA; Mexico. Specimens Examined. Coah (Cowan & Rodrı´guez 3631, TEX; Johnston 8683, F, GH, LL, TEX; Wendt et al. 1949, TEX; Wendt & Kiskind 1616E, TEX).

Selaginella viridissima is distinct from S. bigelovii and S. rupincola in having its leaves acute to obtuse (vs. aristate), the leaf margins denticulate (vs. ciliate), and the bases of aerial stem leaves and rhizome scale-like leaves decurrent or long-decurrent (vs. abruptly adnate).

80. SELAGINELLA WRIGHTII Selaginella wrightii Hieron., Hedwigia 39: 298. 1900. Figs. 266H, J–L. Selaginella rupestris (L.) Spring f. wrightii (Hieron.) Clute, Fern Bull. 16: 52. 1908. Bryodesma wrightii (Hieron.) Soja´k, Preslia 54: 155. 1992. Type. U.S.A. Texas: Uvalde Co., Turkey Creek, near Cline, Wright 828 (B; isotypes GH!, NY!, US!).

Selected Specimens Examined. Chih (Weber & Charette 11626, UC). Coah (Johnston 8607, CAS, MEXU, 9106, CAS, UC). Gro(Lorea 1970, F, NY). NL (Dorr et al. 2494, UC; Muller 2638, UC). Oax (Mickel 4555, NY). Pue (Smith et al. 4125, F). Qro (Dı´az Barriga 3775, NY). Tam (Bartlett 10561, F; Harrison 11599, F). Zac (Lloyd & Kirkwood 20, UC). Unverified, Doubtful, or Mistaken Reports. Hgo (Kenoyer 1053, GH, cited by R. Tryon, 1955, but not verified). SLP (Lundell 96, US, cited by R. Tryon, 1955, but not verified).

Selaginella wrightii seems to be allied to the Californian S. hansenii Hieron., from which it can be distinguished by long (vs. short) strobili, strongly tapering, lanceolate (vs. shortattenuate, ovate-deltate) sporophylls, and green (vs. usually redspotted or wholly red) leaves. Some characters, such as the dorsiventral habit, the dimorphism of the leaves, and the upturned branch tips, suggest a closer alliance of these two species to series Eremophilae than to series Sartorii as suggested by R. Tryon (1955).

SPHENOMERIS

1 1 4. S PH A E R O PT E R I S Sphaeropteris Bernh., J. Bot. (Schrader) 1800(2): 122. 1801. Type: Sphaeropteris medullaris (G. Forst.) Bernh. [⬅ Polypodium medullare G. Forst.]. (Additional synonymy in R. Tryon, 1970.)

603

3 m long, bipinnate-pinnatifid; lamina glaucous beneath; pinnae stalked (1–3 cm); pinnules sessile (to 2 mm stalked), apices obtuse, slightly ascending, entire; veins 1–2 forked; sori medial at vein fork, indusia sphaeropteroid (globose), 0.5–0.8 mm wide, firm, splitting into 2–4 persistent segments.

Terrestrial; stems erect, massive; fronds 1–5 m; stipes smooth to muricate, scaly, the scales spreading or appressed abaxially, the marginal cells similar to those of the central part in shape, size, and orientation, with dark marginal hair teeth; blades pinnate to bipinnate-pinnatifid; surfaces with scales and hairs abaxially, costae and costules with small hairs curving adaxially; veins free; indusia hemitelioid to globose or absent; spores 64 per sporangium; x⫽69. Sphaeropteris has ca. 110 species in tropical regions of the New and Old Worlds, but only one in Mexico. The genus in its restricted sense in the New World (S. horrida and ca. five other species) is monophyletic and sister to Cyathea (Hasebe et al., 1995). References

Distribution. Wet montane forests; 400–2450 m. Mexico; Guat, Bel, Hond, Salv, Nic.

Tryon, R. M. 1971. The American tree ferns allied to Sphaeropteris horrida. Rhodora 73: 1–19; Windisch, P. G. 1977. Synopsis of the genus Sphaeropteris (Cyatheaceae) with a revision of the neotropical exindusiate species. Bot. Jahrb. Syst. 98: 176–198.

Selected Specimens Examined. Chis (Breedlove 32474, DS, NY, 48209, MEXU; Mu¨nch 64, DS). Gro (Gonza´lez Locra 332d, MEXU; Vela´zquez 379, MEXU). Oax (Mickel 4003, NY; Rzedowski 19580, NY; Tenorio L. 2800, MEXU). Ver (Barrington 396, MEXU; Pe´rez & Riba 176, MEXU).

1. SPHAEROPTERIS HORRIDA

Sphaeropteris horrida is distinct by its abundantly scaly stipes and rachises, the scales light golden brown with many dark setae, and the indusia completely enclosing the sporangia.

Sphaeropteris horrida (Liebm.) R. M. Tryon, Contr. Gray Herb. 200: 20. 1970, non Cyathea horrida (L.) Sm., 1793. Figs. 288A–E. Cibotium horridum Liebm., Mexic. Bregn. 279 (reprint 127). 1849. Lectotype (chosen by R. Tryon, 1971: 11). Mexico. Oaxaca: Chinantla, Teotalcingo, Liebmann s.n. [Pl. Mex. 2086, Fl. Mex. 873] (C!, photos GH, US, frag. US!). Cyathea princeps E. Mayer, Gartenflora 17: 10. 1868. Cibotium princeps Linden ex J. Smith, Ferns Brit. For. 291. 1866, nom. nud. in syn. Cyathea princeps J. Sm., Ferns Brit. For. 291. 1866, nom. nud. Syntypes. Linden s.n. ex Volca´n Tuxtla, Veracruz and Chiapas, Mexico, cult. in Brussels (BR). Cyathea bourgeaui E. Fourn., Mexic. Pl. 1: 135. 1872 [“bourgaei”]. Lectotype (chosen by R. Tryon, 1971: 11). Mexico. Valle de Cordoba, Bourgeau 2200 (P; isotypes GH, NY, P [several sheets but none from herb. Fournier (R. Tryon, 1971)], US!). Cyathea glauca E. Fourn., Mexic. Pl. 1: 135. 1872, hom. illeg., non Willd., 1810. Type. Mexico. Veracruz: Orizaba, Bourgeau 2794 (some specimens of this no. are C. mexicana (P!; isotypes BM!, GH, NY!) but some are C. princeps (P)). R. Tryon (1971) lectotypified C. glauca on the C. princeps material. Cyathea muenchii Christ, Bull. Herb. Boissier, se´r. 2, 7: 413. 1907 [“munchii”]. Syntypes. Mexico. Chiapas: San Cristobal, Mu¨nch 64 (herb. Christ, not seen at P fide R. Tryon (1971); frag. Mu¨nch s.n. at US!; isotype DS!).

Trunks 15–20 m tall, 25–40 cm diam.; stipes stramineous to tan, scabrous with stout scale bases but spines absent, scales conform, concolorous, tan, very narrow, with marginal 1-celled teeth generally darker than the scale itself, hairs absent on axes except in adaxial groove, some on adaxial segment midveins (usually 0 or 1); scales progressively narrower distally on frond until hairlike (1 cell wide) on abaxial costules and segment midveins; long hairs grading into narrow scales with bristle-like teeth; blades to

1 1 5 . SP H E N O M E R I S Sphenomeris Maxon, J. Wash. Acad. Sci. 3: 144. 1913, nom. cons. Type: Sphenomeris clavata (L.) Maxon [⬅ Adiantum clavatum L.]. Terrestrial or epipetric; rhizomes slender, long-creeping, protostelic or solenostelic, densely covered with dark, linear, hairlike scales 2 to few cells wide at bases, or septate hairs; fronds determinate, Ⳳ distichous; stipes adaxially sulcate; blades 3–4 times alternately pinnate, anadromous, ultimate segments linear or spatulate, cuneate at bases, truncate at tips; rachises smooth (not spiny); veins few, forking, free; sori borne at or near distal margins of segments, single or 2–4 joined; indusia flattish, pocket-like, attached at bases and sides, single at the clavate apices or borne along a common transverse receptacle; sporangia often few per sorus, sometimes mixed with paraphyses; spores tetrahedral or bilateral; x⫽38, 39, 48, and possibly others. According to Kramer (1971), a genus of 11 species; three occur in the Neotropics, two in Africa, and the rest in Asia and Polynesia. The affinities of Sphenomeris are apparently with Lindsaea and especially Odontosoria, and Sphenomeris is sometimes combined with the latter (e.g., by Kramer in Kubitzki, 1991, and Wolf, 1995).

604

STICHERUS References

Kramer, K. U. 1957. A revision of Lindsaea in the New World, with notes on allied genera. Acta Bot. Neerl. 6: 97–290; Kramer, K. U. 1971. Lindsaea group. Flora Malesiana, ser. II–Pteridophyta, vol. 1(pt. 3): 177–254.

1. SPHENOMERIS CLAVATA Sphenomeris clavata (L.) Maxon, J. Wash. Acad. Sci. 3: 144. 1913. Figs. 289A–D. Adiantum clavatum L., Sp. Pl. 2: 1096. 1753. Stenoloma clavatum (L.) Fe´e, Me´m. Foug. 5: 330. 1852. Odontosoria clavata (L.) J. Sm., Hist. Fil. 264. 1875. Type. Uncertain, based in part on Plumier, Traite´ Foug. Ame´r., pl. 101, f. B, 1705, and Descr. Pl. Ame´r., pl. 50, f. B. 1693. For additional synonymy, see Maxon, Pteridophyt. Porto Rico 6: 487. 1926.

Rhizomes short-creeping, 2 mm diam.; rhizome hairs dark brown, catenate, ca. 1 mm long; fronds 20–60 cm long, 2–3 mm distant; stipes 1⁄3–1⁄2 the frond length, stramineous, glabrous; blades 10–35 ⫻ 3–15 cm, 3–4 times pinnate, the rachises subflexuous, pinnae alternate; ultimate divisions distant, linear-cuneate, mostly 10–15 ⫻ 1–4 mm at apices, sides entire, the truncate apices sharply dentate if sterile; laminar tissue yellowish green, glabrous; indusia erose-dentate; spores globose with a trilete scar, brownish; 2n⫽78 (Fla), 76 (Jam).

vigatus (Willd.) C. Presl [⬅ Mertensia laevigata Willd.] ⫽ Sticherus truncatus (Willd.) Nakai. Terrestrial; rhizomes long-creeping, cord-like, often ca. 2–3 mm diam., protostelic, bearing peltately attached lanceolate brown to castaneous scales, often glabrescent proximally; fronds erect or scandent, monomorphic, often to several meters long; blades pseudodichotomous, with pinnae consistently and repeatedly Ⳳ equally forking, ending in forking pinnatifid laminar parts, the frond apices often becoming dormant; dormant buds concealed by lanceolate, often fringed scales, the proximal one (or several), especially on the main axis, often also with a pair of stipules; accessory pinnae lacking at the branching points of fronds; penultimate divisions pectinate, bearing narrowly lanceolate to linear segments; indument abaxially, and sometimes adaxially, of sparse to numerous scales and/or branched or arachnoid hairs; veins free, generally once-forked; sori abaxial, 3–6 per sorus; sporangia globose to pear-shaped, with oblique annuli; paraphyses absent or filamentous; indusia absent; spores ellipsoid, with perispores laevigate, faintly rugulate or minutely perforate; x⫽34. Sticherus is subsumed in the genus Gleichenia by some authors, but in this treatment we restrict the use of the latter to only those species having bead-like segments (e.g., G. circinnata Sw.) in Australasia to southern Africa; Gleichenia s.str. comprises about 10 species. Sticherus is thought to comprise about 90 species, with about 40 in tropical America. Sticherus is distinguished from Dicranopteris and Gleichenella by having veins 1-forked and by having scaly (vs. hairy) rhizomes. Affinities of Sticherus are clearly with the other gleichenioid fern genera, including also Gleichenia, Stromatopteris, and Diplopterygium (Hasebe et al., 1995; Pryer et al., 2003). In a recent unpublished thesis, Gonzalez (2003) has revised the taxonomy of New World Sticherus. Using her species concepts, the names of three of the four Mexican species will change. These name changes are not adopted herein but are discussed under the individual species that are affected. References

Distribution. Usually on limestone throughout its range, known in Chiapas by a single collection in rain forest, at edge of river; 300 m. USA (Fla); Mexico; Bah, Gr Ant. Specimens Examined. Chis (Breedlove 35362, DS, ENCB, NY).

The small plant size, narrow segments, and terminal pocketlike sori supplied by 1–4 veins serve to distinguish this taxon from Odontosoria.

1 1 6. S T I C H E R US

Ching, R. C. 1940. On the genus Gleichenia. Sunyatsenia 5: 269–288; Gonzales, J. 2003. A taxonomic revision of the genus Sticherus (Gleicheniaceae – Pteridophyta) in the Neotropics. Ph.D. dissertation, Univ. Go¨ttingen, Germany, 172 pp; Holttum, R. E. 1957. Morphology, growth-habit, and classification in the family Gleicheniaceae. Phytomorphology 7: 168–184; Maxon, W. R. 1909. Gleicheniaceae. N. Amer. Fl. 16: 53–63; Nakai, T. 1950. New classification of Gleicheniales, etc. Bull. Natl. Sci. Mus., Tokyo 29: 1–71; Østergaard Andersen, E. & B. Øllgaard. 1996. A note on some morphological terms of the leaf in the Gleicheniaceae. Amer. Fern J. 86: 52–57. 1996; Østergaard Andersen, E. & B. Øllgaard. 2001. 10. Gleicheniaceae. In: G. Harling & L. Andersson (eds.), Flora of Ecuador 66: 105–170.

Sticherus C. Presl, Suppl. Tent. Pterid. 51. 1836. Lectotype (chosen by Christensen, Index Filic. LIV. 1906): Sticherus laeKey to the Mexican Species of Sticherus 1. Segments densely and evenly arachnoid- to lanate-pubescent beneath; pinnae usually 1–2(–3)-forked. 2. “Bud” scales tan to whitish, long-ciliate; ciliate scales dense on penultimate costae (1.5–2 mm long), often running onto

STICHERUS BREVIPUBIS

605

ultimate segment midribs. .............................................................................................................................................................................. 1. S. bifidus. 2. “Bud” scales dark brown, short-ciliate to setose; ciliate scales rare on penultimate costae (0.5–1 mm long), absent on ultimate segment midribs. ........................................................................................................................................................................................ 2. S. brevipubis. 1. Segments sparsely hairy or scaly below, not arachnoid- or lanate-pubescent; pinnae usually 3–5-forked. 3. Pinnae mostly 3–5 cm wide; costules of ultimate segments with scattered deeply divided scales to simple hairs, each with basal reddish brown cell. ...................................................................................................................................................................................... 3. S. palmatus. 3. Pinnae 1.2–3(–3.5) cm wide; costules of ultimate segments with dense pectinate scales. ....................................................... 4. S. underwoodianus.

1. STICHERUS BIFIDUS Sticherus bifidus (Willd.) Ching, Sunyatsenia 5: 282. 1940. Figs. 290A–E. Mertensia bifida Willd., Kongl. Vetensk. Acad. Nya Handl. 25: 168. 1804. Gleichenia bifida (Willd.) Spreng., Syst. Veg., ed. 16, 4: 27. 1827. Dicranopteris bifida (Willd.) Maxon, N. Amer. Fl. 16: 60. 1909. Type. Venezuela. Caracas: Bredemeyer s.n. (B-Willd. 19468!). Mertensia fulva Desv., Me´m. Soc. Linn. Paris 6: 201. 1827. Dicranopteris fulva (Desv.) Underw., Bull. Torrey Bot. Club 34: 255. 1907. Type. Jamaica. Blue Mts., without collector (P; photo BM! ex P). Mertensia gleichenioides Liebm., Mexic. Bregn. 296 (reprint 144). 1849. Gleichenia liebmannii T. Moore, Index Fil. 379. 1862, nom. illeg. Type. Mexico. Veracruz: Cuapa, Liebmann s.n. [Pl. Mex. 2109; Fl. Mex. 604] (US drawing ex B?). This is a teratological form of S. bifidus.

Rhizomes long-creeping, with castaneous to dark brown ciliate scales 2–3 ⫻ 0.2–0.3 mm; stipes 2–3 mm diam.; main axes dormant or often continuing beyond first fork, scaly to glabrescent; dormant buds with tan to whitish, lax, long-ciliate scales 1.5–2 ⫻ 0.2–0.4 mm; pinnae 1–2 times pseudodichotomously forking; axes below ultimate forks pectinate on acroscopic side or lacking segments altogether; penultimate segments 3–8 cm wide, the costae bearing tan to whitish ciliate scales 1.5–2 ⫻ 0.2– 0.3 mm; ultimate segments chartaceous to subcoriaceous, linear, to ca. 4 cm ⫻ 2–3 mm at mid-segment, plane to usually revolute, abaxially densely tomentose with arachnoid hairs; sori inframedial to medial, with 3–4(–5) sporangia per sorus; 2n⫽68 (Chis, Jam, PR).

Selected Specimens Examined. Chis (Breedlove 34600, DS, ENCB). Gro (Cowan 4921, ENCB, NY, UC). Jal (McVaugh 21405, IEB, MEXU, MICH, NY). Me´x (Tejero-Dı´ez 3171, IZTA). Oax (Mickel 1026, ENCB, MEXU, NY, 5911, ENCB, MEXU, NY, UC). Pue (Marquez R. et al. 756, ENCB, MEXU, XAL; Ventura A. 374, ENCB, NY). Tab (Cowan 3171, MEXU, NY). Ver (Pringle 6130, BR, ENCB, K, MEXU, MO, NY, UC, US).

This is the commonest species of Sticherus in Mexico and is readily identified by its abaxially dense indument, which often hides the sori. It is most easily confused with S. brevipubis, which has similar dense arachnoid indument on the blades abaxially, but in that species (as well as in S. palmatus) the secondary axes (immediately proximal to the ultimate forks) are often pectinate on both sides of the axis, whereas in S. bifidus the secondary axes are seldom wholly pectinate; however, both species may be pectinate only on the acroscopic side. The dormant bud scales in S. brevipubis are dark brown, rigid, and mostly short-ciliate, and the costal scales on abaxial surfaces are nearly absent and less than 1 mm long. Gonzales (2003) has adopted the name Sticherus ferrugineus (Desv.) J. Gonzales, comb. ined., for this species, based on Mertensia ferruginea Desv., Ges. Naturf. Freunde Berlin Mag. Neusten Entdeck. Gesamten Naturk. 5: 307 (1811). Type. French Guiana, collector unknown (holotype: P, herb. A. N. Desvaux). Gonzales gave as distribution for S. ferrugineus: Mexico to Panama, Greater and Lesser Antilles, Venezuela, Colombia to Bolivia, and southeastern Brazil. According to Gonzales, S. bifidus s. str. co-occurs with S. ferrugineus in the southern part of this range (Panama, Cuba, Colombia to Bolivia, Venezuela, Trinidad, Guyana, French Guiana, and southeastern Brazil) and differs primarily by having narrower bud scales with truncate bases, lacking aphlebiae and proximal internal segments on the basal 1–2 cm of each axis, having a matted-whitish indument, and having laxer, more appressed axis scales.

2. STICHERUS BREVIPUBIS Sticherus brevipubis (Christ) A. R. Sm., Amer. Fern J. 70: 27. 1980. Figs. 291K–M. Gleichenia brevipubis Christ, Bull. Herb. Boissier, se´r 2, 6: 280. 1906. Lectotype (chosen by Lellinger, Proc. Biol. Soc. Wash. 89: 713. 1977). Costa Rica. 1905, Werckle´ s.n. (P photos UC!, US; isolectotypes UC!, US; frag. NY!).

Distribution. Common on roadside banks, clearings and forest margins, forming dense masses and thickets in lower montane and montane forests, oak forests, oak-beech forests; 150– 2100 m. Mexico; Guat, Hond, Salv, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol.

Rhizomes long-creeping, with dark brown ciliate scales 1.5–2.5 0.2–0.3 mm; stipes 2–3 mm diam.; main axes beyond first fork dormant or often eventually continuing, scaly to glabrescent; dormant buds with dark brown, stiff, short-ciliate scales 1.5–3 ⫻ 0.2–0.3 mm; pinnae 1–2(–3) times pseudodichotomously fork⫻

606

STICHERUS

ing; axes below ultimate forks wholly pectinate or pectinate on only the acroscopic side; penultimate segments 3–6 cm wide, the costae glabrescent or with very small dark brown, appressed, short-ciliate scales 0.5–1 ⫻ 0.1–0.2 mm; ultimate segments chartaceous, linear, to ca. 3 cm ⫻ 1.5–3 mm at mid-segment, plane to usually revolute, abaxially densely tomentose with arachnoid hairs; sori inframedial to medial, with 3–4 sporangia per sorus.

mata W. Schaffn. ex E. Fourn., Mexic. Pl. 1: 137. 1872. Gleichenia palmata (W. Schaffn. ex E. Fourn.) C. Chr., Index Filic. 323. 1905. Lectotype (chosen by Proctor, Brit. Fern Gaz. 9: 218. 1965). Mexico. Veracruz: Jalapa, Schaffner 229 (P, not found; isotype K!). Dicranopteris palmata J. H. Schaffn. ex Underw., Bull. Torrey Bot. Club 34: 259. 1907. Type. Mexico. Veracruz: Orizaba, Pringle 6129 (NY!; isotypes B!, BM!, BR!, GH!, IJ, K!, MEXU!, NY!, UC!, US!).

Rhizomes long-creeping, with castaneous ciliate scales 1.5–2.5 mm; stipes to 8 mm diam.; main axes beyond first fork dormant or eventually continuing, scaly to glabrescent; dormant buds with brownish or orange-brown, long-ciliate scales 1.5–3 ⫻ 0.2–0.5 mm; pinnae mostly 3–5 times pseudodichotomously forking; axes below ultimate forks usually pectinate on both sides or without segments altogether; penultimate segments 3–5 cm wide, the costae with scattered orangish tan, spreading, ciliate scales 0.8–1.5 ⫻ 0.1–0.3 mm; ultimate segments chartaceous, linear-lanceolate, to ca. 2.5 cm ⫻ 1.5–3 mm at mid-segment, plane to revolute, abaxially appearing “granulose” (due to whitish or pale stomata), lacking arachnoid hairs but with simple or branched white or yellowish hairs (reduced scales) 0.5–0.8 mm long, each with a reddish brown cell at base; sori medial to supramedial, with 3–5 sporangia per sorus; 2n⫽68 (Jam). ⫻ 0.2–0.3

Distribution. Infrequent at margins of montane forests, pinecypress forests, evergreen cloud forests, disturbed cloud forests, especially on the Atlantic slope but also in the southern Sierra Madre in Chiapas; 700–2400 m. Mexico; Guat, Hond, Salv, Nic, CR, Pan; Jam, Hisp; Col, Ec (Palacios & Tirado 13244, UC). Selected Specimens Examined. Chis (Breedlove 31966, DS, MEXU, NY, 40232, 41081, DS). Oax (Elliot 378, MEXU, XAL; Martı´nez C. 318, MEXU, NY; Mickel 5109, 6535, 7098, NY). Pue (Arreguı´n 545, ENCB, MEXU). Ver (Nee & Hansen 18661, F, MEXU, XAL).

This species is very similar to S. bifidus, especially in its tomentose indument, but can be distinguished by the darker scales of the dormant buds, the often pectinate secondary axes (proximal to ultimate forks), and the nearly scaleless costae on the penultimate segments. In Chiapas, S. brevipubis occurs at higher elevations than S. bifidus; this tendency exists elsewhere in the range, in both Mexico and Mesoamerica, but is less pronounced. Gonzales (2003) has adopted the name Sticherus fulvus (Desv.) Ching, Sunyatsenia 5: 283, 1940, for this species. This is based on a type from Jamaica, Blue Mts., collector unknown (holotype P). The basionym of this species is Mertensia fulva Desv., Me´m. Soc. Linn. Paris 6: 201, 1827; a synonym is Dicranopteris fulva Underw., Bull. Torrey Bot. Club 34: 255, 1907. According to Gonzales, Sticherus fulvus is mostly replaced in S. America by the closely related S. brevitomentosus Østergaard & B. Øllg.

3. STICHERUS PALMATUS Sticherus palmatus (W. Schaffn. ex E. Fourn.) Copel., Gen. Fil. 28. 1947. Figs. 291A–E. Mertensia palmata W. Schaffn. ex Fe´e, Me´m. Foug. 9: 40. 1857, nom. nud. Gleichenia palmata T. Moore, Index Fil. 380. 1862, nom. nud. Mertensia pal-

Distribution. Frequent in wet forests, in open forests and forest margins, especially on Atlantic slope, pine-oak forests; 450– 2450 m. Mexico; Guat, Bel, Hond, Salv, Nic, CR; Cuba, Jam. Selected Specimens Examined. Chis (Breedlove 32547, DS, MEXU, NY). Gro (Breedlove 65135, CAS; Rzedowski 18533, ENCB, NY). Hgo (Sa´nchez-Mejorada 877, MEXU). Oax (Hallberg 1529, NY, UC; Mickel 4717, 5926, ENCB, NY, UC). Pue (Moore 6301, UC, US; Sa´nchezMejorada 945, ENCB, MEXU, NY). Ver (Copeland herb. 9, MEXU, UC; Orcutt 2837, DS, MEXU, MO, NY). Unverified, Doubtful, or Mistaken Reports. DF (reported by Mickel & Beitel, 1988, but not verified). Mor (reported by Mickel & Beitel, 1988, but not verified; not listed by Riba et al., 1996). Tab (reported by Magan˜a, 1992, but not verified).

This species differs from S. bifidus in lacking a tomentose indument abaxially on the blade tissue. It more closely resembles S. underwoodianus, but that species has narrower penultimate divisions with broader and more abundant scales on the costae. One of the specimens cited by Smith (1981) as S. palmatus (Breedlove 23221, DS), is now judged to be S. underwoodianus. Gonzales (2003) has adopted the name Sticherus furcatus (L.)

STIGMATOPTERIS

Ching, Sunyatsenia 5: 283, 1940, for this species. This is based on a Plumier plate, Descr. Pl. Ame´r. t. 20 (1693). The basionym of this species is Acrostichum furcatum L., Syst. Nat., ed. 10, 2: 1321 (1759), synonyms Mertensia furcata (L.) Willd., Kongl. Vetensk. Acad. Nya Handl. 166 (1804) and Gleichenia furcata (L.) Spreng., Syst. Veg. 4: 26 (1827). Proctor (1977) treated this species as endemic to the Lesser Antilles.

4. STICHERUS UNDERWOODIANUS Sticherus underwoodianus (Maxon) Nakai, Bull. Natl. Sci. Mus. Tokyo 29: 31. 1950. Figs. 291F–H, J. Dicranopteris underwoodiana Maxon, N. Amer. Fl. 16: 59. 1909. Gleichenia underwoodiana (Maxon) C. Chr., Index Filic., Suppl. 1: 44. 1913. Type. Mexico. Chiapas: Ghiesbreght 271 (NY!; isotypes BM!, GH!, K!, NY!, PH!, US!).

Rhizomes long-creeping, with castaneous ciliate scales 1.5–2.5 0.2–0.3 mm; stipes 1.5–2 mm diam.; main axes beyond first fork dormant or eventually continuing, scaly to glabrescent, often also with a pair of stipules at the proximal fork; dormant buds with brown to orangish brown, long-ciliate scales 1.5–3 ⫻ 0.2–0.5 mm; pinnae mostly 3–5 times pseudodichotomously forking; axes below ultimate forks usually pectinate on both sides; penultimate segments 1.2–3(–3.5) cm wide, the costae with dense orange-tan, spreading, ciliate scales 1–2 ⫻ 0.3–0.5 mm; ultimate segments chartaceous to subcoriaceous, narrowly deltate, to ca. 1.5 cm ⫻ 2–3 mm at mid-segment, revolute, abaxially glabrous and lacking arachnoid hairs, but with simple or branched white or yellowish hairs (reduced scales) 0.5–0.8 mm long on veins, also with scales on costules similar to those of costae, but smaller; sori medial to supramedial, with 3–5 sporangia per sorus.

607

NY, UC, 6769, NY, UC). Pue (Clausen & Edwards 7477, MEXU, NY). Ver (Ventura A. 212, NY).

This species can generally be distinguished from congeners in Mexico by its narrower pinnae and occurrence at higher elevations. The type has pinnae 3–3.3 cm wide, whereas most Mexican material is 1.5–2.5 cm wide. Sticherus underwoodianus resembles S. jamaicensis (Underw.) Nakai of Cuba, Jamaica, and Hispaniola, and Smith (1981) suggested they might be conspecific. They do indeed appear to be nearest allies, but, as Underwood pointed out, S. jamaicensis is minutely glandular-puberulous (minute one-celled glandular hairs) on veins and leaf tissue and lacks stellate or branched hairs on the ultimate veins, whereas S. underwoodianus has glabrous leaf tissue and dissected scales on veins and costae. Both S. underwoodianus and S. palmatus differ from S. bifidus and S. brevipubis in having more highly forking pinnae, 3–5forked vs. 1–2(–3)-forked. See S. palmatus, the most closely related Mexican species, for further comparison.

⫻

Distribution. Frequent at higher elevations, on wet banks along roads and forest margins, cloud forests, elfin forests, alder forests, pine forests, pine-oak forests; 1500–2700(–3050) m. Mexico; Guat, Hond, Salv. Selected Specimens Examined. Chis (Breedlove 22009, DS, MEXU, NY; Ghiesbreght 271, BM, GH, NY, PH, US). Gro (Hinton et al. 14239, NY, US; Lorea 2334, IEB, FCME). Hgo (Gimate 803, ENCB, MEXU, NY; Herna´ndez Magan˜a 4294, MEXU, UC). Oax (Mickel 4243, ENCB,

1 1 7 . ST I GM A T O P T E R I S Stigmatopteris C. Chr., Bot. Tidsskr. 29: 292. 1909. Dryopteris Adans. subg. Stigmatopteris (C. Chr.) C. Chr., Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 7, 10: 73. 1913. Type: Polypodium flavopunctatum Kaulf. [⫽ Stigmatopteris rotundata (Willd.) C. Chr.]. Terrestrial; rhizomes short-creeping to suberect, bearing scales; fronds medium-sized to large, monomorphic; stipes with numerous vascular bundles arranged in an omega-shape; blades 1pinnate to 2-pinnate-pinnatifid, apices pinnatifid, in some species with proliferous buds along rachises; rachises and costae adaxially grooved, the grooves Ⳳ confluent, midribs abaxially usually with linear to ovate, tan scales, acicular hairs absent; pinnae gradually shortened distally, increasingly adnate and decurrent, blade apices pinnatifid and confluent with distal pinnae; blades pellucid-punctate with immersed yellowish glands (best observed with transmitted light), glabrous; veins free (ours) or irregularly netted, ending well short of margin in clavate tips (as seen adaxially); sori abaxial, round, exindusiate; paraphyses absent (ours) or present; spores oblong, bilateral, with brownish, winged perispores; x⫽41. Stigmatopteris is a neotropical genus of 23 species, mostly of South America. Affinities of the genus have generally been assumed to be with Cyclodium, also primarily South American, and other genera such as Polybotrya and Olfersia (Moran, 1991). However, unpublished molecular data (Cranfill, pers. comm.) suggests an alliance of Stigmatopteris with Ctenitis. In any case, Stigmatopteris is clearly a member of the dryopteroid clade, with base chromosome nnumber x⫽41. Stigmatopteris superficially resembles some species of Thelypteris, but stipes have more than three vascular bundles, veins end short of the margin, blades lack acicular hairs, and sori are exindusiate.

608

STIGMATOPTERIS References

Christensen, C. 1909. On Stigmatopteris, a new genus of ferns with a review of its species. Bot. Tidsskr. 29: 291–304; Christensen, C. 1913. A monograph of the genus Dryopteris. Part 1. The tropical American pinnatifid-bipinnatifid species. Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 7, 10: 55– 282; Moran, R. C. 1991. Monograph of the neotropical fern genus Stigmatopteris (Dryopteridaceae). Ann. Missouri Bot. Gard. 78: 857–914.

Key to the Mexican Species of Stigmatopteris 1. Blades pinnate; pinnae serrate or lobed less than 1⁄4 of distance to costae; all but the proximal 2–4 pinna pairs long-decurrent; costal scales lanceolate, hair-like, sparse. ........................................................................................ 1. S. longicaudata.

1. Blades pinnate-pinnatifid; pinnae deeply incised to a narrow wing to the costae, cut 1⁄2–3⁄4 of distance to costae; only the distalmost pinna pairs long-decurrent; costal scales ovate-lanceolate, 1–2 ⫻ 0.3–0.5 mm wide, scattered. ............................................................................... 2. S. sordida.

1. STIGMATOPTERIS LONGICAUDATA Stigmatopteris longicaudata (Liebm.) C. Chr., Bot. Tidsskr. 29: 300. 1909. Figs. 292A, B. Polypodium longicaudatum Liebm., Mexic. Bregn. 209 (reprint 57). 1849. Dryopteris longicaudata (Liebm.) Maxon, Contr. U.S. Natl. Herb. 13: 18. 1909. Lectotype (chosen by Smith, 1981: 213). Mexico. Veracruz: “Baranca de Huitamalco,” Liebmann s.n. [Pl. Mex. 2694, Fl. Mex. 737] (C!; isosyntype K!).

Rhizomes suberect; rhizome scales dark-castaneous, lustrous, lanceolate, 5–7(–10) ⫻ 1–3 mm, margins entire to denticulate; fronds 1–1.4 m long; stipes stramineous to tan, 25–70 cm long, 1⁄2–2⁄3 the frond length, with brown, ovate-lanceolate scales at stipe bases; blades pinnate, 40–60 ⫻ 20–26 cm; pinnae 12–18 pairs, 12–24 ⫻ 2–3 cm, serrately incised or lobed less than 1⁄4 their width, proximal pinnae stalked to 5 mm, all but proximal 2–4 pairs adnate and long-decurrent onto rachises, those of distal 1⁄3 of blade connected by a wing; rachises with sparse bifid or trifid scales, tips clavate; costae abaxially with sparse, tan, lanceolate, scales 0.3–0.8 mm long with clavate tips; adaxial costal grooves minutely pubescent, hairs especially dense near pinna bases; veins 3–6 pairs per segment, not united or rarely the basalmost veins casually united; sori in a double row between costules, 2–5 pairs of sori per segment.

Distribution. Terrestrial in wet montane forests; 450–1700 (–2250) m. Mexico; Guat, Hond, CR, Pan; Col, Ven, Fr Gui, Ec, Peru, Bol. Selected Specimens Examined. Chis (Breedlove 35128, DS, LL, 56494, CAS, ENCB, MICH, MO, NY; Rovirosa 822, K, PH). Oax (Mickel 5662, ENCB, NY, UC, US, 6543, NY, UC). Pue (Ventura A. 484, ENCB, LL, NY). Ver (Finck 169, UC; Nee et al. 26382, F, NY, XAL, Z).

The once-pinnate blades with shallowly lobed and longdecurrent pinnae distinguish S. longicaudata from S. sordida.

2. STIGMATOPTERIS SORDIDA Stigmatopteris sordida (Maxon) C. Chr., Index Filic., Suppl. 3: 175. 1934. Figs. 292C, D. Dryopteris sordida Maxon, Contr. U.S. Natl. Herb. 24: 60. 1922. Type. Guatemala. Alta Verapaz: Cubilquitz, Tu¨rckheim II 1432 (US!; isotype US!). Stigmatopteris chimalapensis Mickel & Beitel, Mem. New York Bot. Gard. 46: 356. 1988. Type. Mexico. Oaxaca: Mpio. Sta. Marı´a Chimalapa, Can˜ada E de la vereda a La Gloria, ca. 8 km SE de Sta. Marı´a, Herna´ndez G. 1282 (NY!; isotype MEXU!).

Rhizomes short-creeping to suberect; rhizome scales dark brown, mostly 3–7 ⫻ 1–3 mm, margins entire to denticulate; fronds 70–100⫹ cm; stipes stramineous to brownish, ca. 1⁄2 the frond length; blades pinnate-pinnatifid; pinnae 15–20 pairs, (8–)13–23 ⫻ 1.3–4 cm, lobed (1⁄2–)2⁄3–3⁄4 their width, somewhat inequilateral at base (acroscopic segments longer, basiscopic segments more oblique), pinnules serrate in distal 1⁄3, proximal pinnae short-stalked, middle ones sessile to partially adnate, distal ones fully adnate and short-decurrent, only the distalmost few joined by a wing; rachises and costae abaxially with scattered to numerous tan, ovate-lanceolate, fibrillose scales 1–2 ⫻ 0.3–0.5 mm; adaxial costal grooves pubescent, hairs especially dense near pinna bases; veins (4–)6–11 pairs per segment, not united; sori (2–)5–10 pairs per segment.

TECTARIA

609

on blades absent or very often of reddish, short, jointed (ctenitoid) hairs, especially on rachises and costae adaxially; veins netted (ours), often with free, included, sometimes forked veinlets; sori abaxial and round to oblong, rarely the sporangial in continuous marginal coenosori; indusia peltate or reniform, rarely continuous (with marginal coenosori); spores bilateral, perispores cristate; x⫽40.

Distribution. Terrestrial in wet lowland and lower montane forests; 150–600(–1000) m. Mexico; Guat, Bel (Holst 5897, UC), Hond, Nic, CR, Pan; Col, Ec. Specimens Examined. Chis (Ghiesbreght 288, YU; Purpus 6870, BM, MO, NY, UC, US). Oax (Mickel 6403, ENCB, NY, UC, US; Wendt et al. 5745, CHAPA, MEXU, NY). Ver (Lira 163, MEXU, UAMIZ, XAL; Ortı´z y Martiniano 159, IEB, NY).

Stigmatopteris chimalapensis was distinguished by its smaller blades, less deeply incised pinnae, and fewer veins per segment. These characteristics are unusually variable in Stigmatopteris, even on fronds from a single plant, and smaller variants have a tendency to become precociously fertile. A recent collection of typical S. sordida (Wendt et al. 5745, 13 km NE of Sta. Marı´a) from the vicinity of the type collection of S. chimalapensis further suggests that the two are conspecific. Stigmatopteris sordida differs from S. longicaudata by the more deeply incised pinnae and the distal pinnae being only weakly or not at all decurrent onto the rachises.

1 18. T E C T A R I A Tectaria Cav., Anales Hist. Nat. 1: 115. 1799. Type: Tectaria trifoliata (L.) Cav.[⬅ Polypodium trifoliatum L.]. Aspidium Sw., J. Bot. (Schrader) 1800(2): 4, 29. 1801. Lectotype (first chosen by J. Smith?, Hist. Fil. 200. 1875): Aspidium trifoliatum (L.) Sw. [⬅ Tectaria trifoliata (L.) Cav.] Dictyoxiphium Hook., Gen. Fil., t. 62. 1840. Type: Dictyoxiphium panamense Hook. [⬅ Tectaria panamensis (Hook.) R. M. Tryon & A. F. Tryon] For additional synonymy, see Kramer & Green (1991).

Terrestrial or epipetric; rhizomes mostly woody, stout, short, compact, short-creeping to erect, bearing brownish, concolorous, non-clathrate scales; fronds small to often large, monomorphic or somewhat dimorphic, rarely strongly dimorphic, clumped; blades membranaceous to chartaceous, simple to mostly pinnatifid or 1–2-pinnate, rarely more divided; indument

Tectaria is a large pantropical genus (ca. 200 species); it is most developed in Southeast Asia and adjacent Pacific islands. There are 25–30 species in the Neotropics, mostly in South America, with seven in Mexico. Three additional species are known from adjacent parts of Guatemala and Belize: T. nicotianifolia (Baker) C. Chr., T. plantaginea (Jacq.) Maxon, and T. rivalis (Mett. ex Kuhn) C. Chr. [misidentified as T. rheosora (Baker) C. Chr. by Stolze, 1981; see Grayum, Phytologia 64: 30– 35. 1987]. Mexican species generally occur on limestone at low or middle elevations, below 1500 m. Tectaria is distinct in its finely netted veins (the areoles sometimes with free included veinlets) and round, abaxial sori (except T. panamensis). It has often been considered most closely related to the free-veined genus Ctenitis (see, e.g., Holttum, 1986), agreeing with that in the non-grooved midribs adaxially and the presence of pluricellular hairs on the blades; however, molecular data (e.g., Hasebe et al., 1995; Cranfill, unpubl. data) suggest that this relationship may be more superficial than real. Some pteridologists have chosen to recognize Tectaria (as well as Ctenitis and a number of segregate tectarioid genera) in a family Tectariaceae (e.g., Moran in Davidse et al., 1995), separate from the Dryopteridaceae, but molecular evidence does not support a family with this circumscription. Dictyoxiphium has been segregated as a monotypic genus, distinct by its linear, dimorphic blades and marginal coenosori. It is very closely allied to Tectaria and crosses with that genus (T. incisa) to form the hybrid “genus” Pleuroderris Maxon (see Wagner et al., 1978 for more details and additional references). The misshapen-spored hybrid, Pleuroderris michleriana (D. C. Eaton) Maxon, has been collected from Guatemala to Colombia and can be expected wherever its two parents co-occur (as in Veracruz, Oaxaca, and Chiapas). Intermediate, but highly irregular in morphology, the fronds of Pleuroderris range from shallowly lobed to pinnatisect or fully pinnate at base, and the sori are round to elongate and scattered irregularly along and near the segment margins. References Holttum, R. E. 1986. Studies in the fern-genera allied to Tectaria Cav. VI. A conspectus of genera in the Old World regarded as related to Tectaria, with description of two genera. Gard. Bull. Singapore 39: 153–167; Morton, C.V. 1966. The Mexican species of Tectaria. Amer. Fern J. 56: 120–137; Wagner, W. H., Jr., F. S. Wagner & L. D. Go´mez P. 1978. The singular origin of a Central American fern, Pleuroderris michleriana. Biotropica 10: 254–264.

Key to the Mexican Species of Tectaria 1. Sporangia in a continuous submarginal band on each blade; blades linear-oblong, entire. ............................................................... 5. T. panamensis. 1. Sporangia in discrete round or oblong sori back from the blade margins; blades ovate to usually deltate, usually lobed, pinnatifid, or 1–2-pinnate.

610

TECTARIA

2. Blades 3–10 cm long, generally merely lobed or shallowly pinnatifid; rhizomes creeping, 1–3(–5) mm diam.; blades membranaceous, generally hairy on veins and tissue between veins. .................................................................................................. 1. T. fimbriata. 2. Blades usually greater than 15 cm long, usually pinnatifid to 1–2-pinnate; rhizomes creeping to suberect, more than 3 mm diam.; blades papyraceous, firm, glabrous to hairy. 3. Blades 2-pinnate-pinnatifid to subtripinnate; free included veinlets mostly absent in vein areoles; rhizome scales with dense hairs on surfaces and margins; pinna margins ciliate with hairs 0.3–0.8 mm long, especially in sinuses. .............................. 4. T. mexicana. 3. Blades pinnate-pinnatifid or entire with cordate bases; free included veinlets often present in vein areoles; rhizome scales glabrous or with papillate surfaces and ciliate to entire margins; pinna margins non-ciliate. 4. Indusia peltate, 1.5–2.5(–3.5) mm wide; 0–2(–4) lateral pinna pairs, with cordate bases; apical segment of each blade with a cordate base; proximal pinnae (if present) each with long stalk, a single acroscopic lobe, and 1 or 2 basiscopic lobes. 5. Blades glabrous on both sides, or occasionally with short hairs 0.1–0.2 mm on costae abaxially. ........................... 2. T. heracleifolia. 5. Blades densely hirsute on both sides, the hairs 0.5–1.5 mm long on costae, veins, and tissue between veins. ........... 7. T. trichodes. 4. Indusia round-reniform, attached at sinuses, 1–1.3(–1.5) mm wide; 3–10 lateral pinna pairs with decurrent bases; terminal segment of each blade with a cuneate base; proximal pinnae with short stalks and either only a single basiscopic lobe or a small acroscopic lobe and 2–3 basiscopic lobes. 6. Blade surfaces glabrous abaxially, except along costae; basal pinnae entire, repand, or very shallowly lobed, each with a single large basiscopic lobe, occasionally with an acroscopic lobe; sori in 2 distinct rows between main lateral veins on each blade; 0–800(–1200) m. ................................................................................................................................................. 3. T. incisa. 6. Blade surfaces hairy abaxially; basal pinnae coarsely serrately lobed and with 2–3 large basiscopic lobes and a single reduced acroscopic lobe; sori somewhat irregularly scattered or in 2–4 rows between main lateral veins; (850–)1050– 1750 m. ........................................................................................................................................................................................ 6. T. transiens.

1. TECTARIA FIMBRIATA Tectaria fimbriata (Willd.) Proctor & Lourteig, Bradea 5: 386. 1990. Figs. 293A–C. Aspidium fimbriatum Willd., Sp. Pl., ed. 4, 5(1): 213. 1810. Type. Plumier, Traite´ Foug. Ame´r., pl. 149. 1705, based on a plant from Hispaniola. Polypodium lobatum Rich. ex Willd., Sp. Pl., ed. 4, 5(1): 164. 1810, hom. illeg., non Hudson, 1762. Sagenia lobata C. Presl, Suppl. Tent. Pterid. 87. 1836. Tectaria lobata (C. Presl) C. V. Morton, Amer. Fern J. 56: 127. 1966. Type. Bahama Islands. s. coll. s.n. (not found in Herb. Willd., presumably in P). Tectaria minima Underw., Bull. Torrey Bot. Club 33: 199. 1906. Type. U.S.A. Florida: Between Cutler and Longview Camp, Small & Carter 855 (NY!; isotype US!).

Rhizomes short-creeping to suberect, 1–3(–5) mm diam.; rhizome scales uniformly tan to brown, glabrous, 1–3 ⫻ 0.2–0.5 mm; fronds mostly 3–15 cm long; stipes stramineous, slender, 0.3–1 mm diam., often longer (to 2.5 times) than blades, moderately to densely hairy, hairs 0.1–0.3 mm long; blades membranaceous, ovate to deltate, simple or broadly lobed to pinnatifid or with one free pair of proximal pinnae, 3–6(–10) ⫻ 1.5–3(–7) cm, cordate at bases; pinnae 0–1 lateral pair per blade, stalked to ca. 1 mm, to 4 ⫻ 2.5 cm, shallowly lobed on both sides, basiscopic lobes more pronounced; rachises (midribs) glabrescent to often moderately or densely hairy, buds absent; vein areoles mostly lacking included veinlets; indument of fine but dense hairs on blade margins, midribs, veins, and often laminae (between veins) abaxially, also on veins and laminae adaxially, hairs 0.1–0.3 mm long, septate; sori round, irregularly arranged between the principal lateral veins; indusia tan, thin, round-reniform, glabrous or sparsely hairy, each attached at a sinus; 2n⫽80 (Fla).

Distribution. Sinkholes and shaded ledges, on limestone; low elevations, ca. 100 m or less. USA (Fla); Mexico; Guat (Stevens et al. 25562, MEXU); Bah, Cuba, Hisp, PR. Specimens Examined. Yuc (Schott 780, MO, US–2 sheets, 2 localities, Cenote de Sta. Ana and Pozo de Guayma).

This species is easily distinguished from other Mexican tectarias by the relatively small, simple or merely lobed blades (rarely with one pair of free pinnae), narrow, short-creeping to suberect rhizomes 1–3 mm thick, and finely pubescent blades. It is primarily an Antillean species, rare on the continent.

2. TECTARIA HERACLEIFOLIA Tectaria heracleifolia (Willd.) Underw., Bull. Torrey Bot. Club 33: 200. 1906. Figs. 294B, D. Aspidium heracleifolium Willd., Sp. Pl., ed. 4, 5(1): 217. 1810. Type. based on Plumier, Traite´ Foug. Ame´r., pl. 147. 1705, illustrating a plant from Hispaniola. Polypodium cordifolium M. Martens & Galeotti, Me´m. Foug. Mexique 31,

TECTARIA INCISA pl. 4, f. 2., hom. illeg., non L., 1753. Type. Mexico. Veracruz: “Riviere de l’Antigua,” Galeotti 6313 (BR!–photo US!; isotype BR!). Tectaria heracleifolia (Willd.) Underw. var. maxima C. V. Morton, Amer. Fern J. 56: 126. 1966. Type. Mexico. Veracruz: Near Pueblo Nuevo, Reeder & Reeder 1975 (US!; isotype YU!). The placement of T. heracleifolia var. maxima in synonymy here is tentative. The type of this differs from all other Mexican specimens seen in being 2-pinnate at the base, with four pairs of pinnae. These characteristics suggest that it may be a hybrid, as Morton surmised, or perhaps more than just a variety of a widespread, common, and otherwise morphologically fairly uniform species.

Rhizomes suberect to erect, woody, 0.5–2 cm diam.; rhizome scales bicolorous, central portion dark brown, lustrous, margins thin, occasionally ciliate, slightly fimbriate, pale, lanceolate, 2–5 ⫻ 0.3–0.5 mm; fronds mostly (12–)30–70 cm long; stipes often longer than blades, stramineous to reddish brown at base, lustrous, with few scales at base and scattered hairs 0.1 mm long; blades firmly chartaceous, simple to 1-pinnate, deltate, (14–)20– 35(–50) ⫻ (2.5–)14–40 cm; rachises with scattered hairs 0.1 mm long, buds absent; pinnae 0–2(–4) lateral pairs (juvenile fronds merely cordate or trilobate), pentagonal, lateral and terminal ones with cordate base, basal pair much the largest, 12–25 cm long, long-stalked (3–30 mm), with 1–2 prominent basiscopic lobes and a shorter acroscopic lobe, lobes acute, 5–15 cm long, entire to sinuate; vein areoles pentagonal or hexagonal, commonly with a single, free, included veinlet; indument abaxially absent on costae and costules, or of sparse hairs 0.1–0.2 mm long, or of short glandular hairs, adaxially of dense to scattered hairs 0.1 mm long along costae, otherwise usually glabrous; sori round, in 2 rows, one on either side of main lateral veins; indusia persistent, thick, peltate, 1.5–2.5(–3.5) mm diam., with scattered short hairs on surface; 2n⫽80 (SLP, CR), 160 (Fla, Jam), ca. 160 (Trin).

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(Ferna´ndez N. 3464, ENCB, IEB, NY). QR (Te´llez 1714, CAS). SLP (Sundberg 2519, UC). Tab (Hammel & Merello 15530, UC). Tam (Palmer 184, NY, US). Ver (Delgadillo 12277, UC). Yuc (Butterwick 240, CAS, TEX, UC). Unverified, Doubtful, or Mistaken Reports. Tres Marı´as (reported by Lenz, 1995, but not verified).

This is one of the commonest ferns at low and middle elevations in southern and eastern Mexico. It often grows on calcareous rocks or in sink holes, and may occasionally hybridize with other Tectarias, particularly T. incisa. The peltate indusia, 0–2 (rarely 3 or 4) lateral pinna pairs with cordate bases, longstalked proximal pinnae with 1–2 prominent basiscopic lobes and a single acroscopic lobe, narrow, bicolorous scales, and cordate-based terminal segment distinguish T. heracleifolia from T. incisa and T. transiens (q.v.). Most specimens of T. heracleifolia have 0–2 pairs of lateral pinnae, but a few (e.g., Robles G. 207, XAL) have three or even four pinna pairs; occasional specimens are precociously fertile, with blades small and merely lobed at base (e.g., Martı´nez S. 15811, TEX, with blades 10 ⫻ 3 cm). See T. mexicana for comparison with that species. A few specimens of T. heracleifolia from Mexico are minutely pubescent with hairs 0.1–0.2 mm long on the costae (both sides) and also on the lamina between veins abaxially, e.g., Matuda 27185 (US) and 27539 (UC), both from Edo. Me´xico. Neither of these specimens approaches T. trichodes (which see) in pubescence.

3. TECTARIA INCISA Tectaria incisa Cav., Descr. Pl. 249. 1801. Type. Puerto Rico. Ventenat s.n. (MA, seen by Christensen, Dansk Bot. Ark. 9(3): 14. 1937). For additional synonymy, see Morton (1966).

Distribution. Wet stream banks and rock outcrops (usually limestone); 50–1000(–2100) m. USA (Fla); Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Bah, Gr & L Ant; Col, Ven, Trin, Ec. Selected Specimens Examined. Camp (Martı´nez S. 29490, UC). Chis (Croat & Hannon 65032, UC). Coah (Lyonnet 421000005, MEXU). Col (Sanders 10365, UC). Gro (Mexia 8894, ARIZ, CAS, LL, UC, US). Hgo (Clarke 6882, NY). Jal (Dı´az L. 16254, UC). Me´x (Matuda 27539, UC). Mich (Hinton 16290, DS, LL, UC). Mor (Lyonnet 755, NY). Nay (Nelson 4280, US). NL (Pringle 1983, ARIZ, CAS, DS, TEX, UC, US). Oax (Wendt et al. 5834, NY). Pue (Butterwick B-66, TEX, UC). Qro

Rhizomes suberect, woody, 1–2 cm diam.; rhizome scales concolorous, brown, 3–5 ⫻ 0.8–1 mm, margins strongly fimbriate; fronds mostly 70–170 cm long; stipes about equaling blade, stramineous to brownish, with scattered hairs 0.1 mm long; blades chartaceous, 1-pinnate, oblong to ovate-oblong, 40–75 ⫻ 20–60 cm; rachises glabrous or glabrescent, buds present or absent; pinnae 3–10 pairs, oblique, apical pinna with decurrent base, distal pinnae adnate basiscopically with decurrent base, basal pair short-stalked (to 5 mm), with an elongate, acute, basal, basiscopic lobe, otherwise subentire or undulate, other pinnae mostly 20–25 ⫻ 3–6(–7) cm; indument abaxially absent on costae and costules, or of scattered hairs 0.1 mm long on veins, adaxially of dense to scattered hairs 0.1 mm long on costae, elsewhere hairs absent; vein areoles pentagonal or hexagonal, commonly with included free veinlets; sori round, in 2 rows, one row on each side of main lateral veins; indusia persistent, reniform or round, with a basal sinus, lobes overlapping, indusia thus appearing peltate, 1–1.3 mm diam., glabrous; 2n⫽80 (CR, Jam, Trin, Tobago, Peru), 160 (Jam, PR, Trin).

612

TECTARIA

The non-peltate indusia, 3–10 lateral pairs of pinnae with decurrent bases, terminal segment with a cuneate base, and proximal pinnae with a single basiscopic lobe distinguish T. incisa from T. heracleifolia.

Differing from f. incisa in the presence of one or more proliferous buds at the bases of the pinnae on the adaxial side of the lamina.

Key to the Mexican Forms of Tectaria incisa 1. Blades lacking proliferous buds. ......................................... 3a. f. incisa. 1. Blades with one to several proliferous buds on the adaxial side, usually at pinna bases. ............................... 3b. f. vivipara.

3a. Tectaria incisa f. incisa

Figs. 293H, J.

Aspidium martinicense Spreng., Anleit. Kenntn. Gew. 3: 133. 1804. Aspidium macrophyllum Rudolphi, Bemerk. Geb. Naturges. 2: 103. 1805, hom. illeg. superfl. Tectaria martinicense (Spreng.) Copel., Philipp. J. Sci., Bot. 2: 410. 1907. Type. Santo Domingo [Hispaniola]. Poiteau s.n. [incorrectly said to be from Martinique by Sprengel].

Differing from f. vivipara by the lack of buds in the axils of distal pinnae.

Distribution. Terrestrial, in wet forests; 150–250 m. Mexico; Guat, Bel, Nic, CR, Pan; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, s Braz, Bol. Moran (in Davidse et al., 1995) also included Honduras and Jamaica in the range, but Proctor (1985) did not mention a viviparous form in Jamaica. Selected Specimens Examined. Chis (Martı´nez S. 24934, MEXU; Rovirosa 866, NY). Oax (Herna´ndez G. 861, CHAPA, 1814, CAS, CHAPA, 2587, CHAPA, NY). Tab (Cowan 3535, NY; Gilly & Herna´ndez X. 306, US). Ver (Dorantes 3731, IEB; Orcutt 2979, ASU, DS, US; Smith 2015, UC, US).

Distribution. Slopes and stream banks; 0–800 (–1200) m. Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol, Parag, n Arg. Selected Specimens Examined. Chis (Breedlove 22522, DS, NY, 34506, DS). Gro (Hinton 11021, LL, NY; Lorea 2447, FCME, IEB). Oax (Solano C. 519, CHAPA, NY). Tab (Cowan et al. 3535, UC; Magan˜a A. 215, CAS; Zamudio R. 1407, 1430, IEB). Ver (Wendt et al. 3981, UC). Unverified, Doubtful, or Mistaken Reports. Tam (reported by Sharp, 1954, but not verified, and probably based on a misidentification).

Both diploid and tetraploid cytotypes are known and both are reported from Jamaica where f. vivipara is not known to occur. Further morphological and cytological studies are needed on T. incisa from throughout its very wide range. 3b. Tectaria incisa f. vivipara (Jenman) C. V. Morton, Amer. Fern J. 56: 131. 1966. Figs. 294H, J–L. Nephrodium macrophyllum Baker var. viviparum Jenman, Bull. Bot. Dept. 3: 238. 1896. Tectaria martinicensis (Spreng.) Copel. var. vivipara (Jenman) Domin, Rozpr. Kra´l. Ceske´ Spolecˇn. Nauk., Trˇ. Mat.-Prˇ´ır. 2: 231. 1929. Lectotype (chosen by Mickel & Beitel, 1988: 360). Guyana. Isorooroo R., Pomeroon, Jenman s.n. (NY!).

Forma vivipara differs from the typical form primarily by its viviparous habit. We are unable to find additional characters by which it differs from f. incisa. It appears that old, senescent blades gradually sag and come in contact with the ground. At this stage, the buds become active and generate new plantlets. Viviparous forms of T. incisa occur nearly throughout the range of the species, except in the Antilles and in southern Brazil, Argentina, and Paraguay. Most often, buds occur on sterile fronds, the pinnae of which tend to be entire, rather than repand, as on fertile fronds. Jermy and Walker (1985, Bull. Brit. Mus. (Nat. Hist.), Bot. 13: 223) described a diploid from Trinidad, Tectaria vivipara Jermy & T. G. Walker, which they believed to be the same as the Tectaria incisa f. vivipara, as recognized here. Because their new type was of known genetic constitution, they treated T. vivipara as a new species, giving it a new type. They described their new species as differing from T. incisa by its viviparous nature and the presence of “hairs on the abaxial side of the costae”. Mexican, Central American, and South American specimens of T. incisa f. vivipara are essentially glabrous on the costae abaxially, and it may be that the Trinidad specimens are not conspecific with viviparous plants from elsewhere in the range. Hence, we exclude T. vivipara from synonymy. Moran (in Davidse et al., 1995) recognized T. vivipara Jermy & T. G. Walker as distinct in Flora Mesoamericana, and ascribed to it a wide range, including Mexico. In Trinidad, T. incisa was found to be a tetraploid and formed sterile triploid hybrids with T. vivipara (Jermy & Walker, 1985,

TECTARIA PANAMENSIS

613

Bull. Brit. Mus. (Nat. Hist.), Bot. 13: 269). This hybrid, T. ⫻bulbifera Jermy & T. G. Walker, has abortive bulblets in the pinna axils and malformed spores. Cowan 3535 (NY) from Tabasco, Mexico, was suggested by Mickel and Beitel (1988) as being this hybrid, but in view of the above discussion, we now think it should be referred to Tectaria incisa f. vivipara, a stage with undeveloped buds.

4. TECTARIA MEXICANA Tectaria mexicana (Fe´e) C. V. Morton, Amer. Fern J. 56: 133. 1966. Figs. 294E–G. Sagenia mexicana Fe´e, Me´m. Foug. 5: 313. 1852. Syntypes. Mexico. Oaxaca: Galeotti 6542 (P!; isosyntype RB); Etlapa, Galeotti 6484 (P!–2 sheets; isosyntypes BR!, RB). Aspidium latifolium C. Presl, Reliq. Haenk. 1(1): 30. 1825, non Tectaria latifolia (G. Forst.) Copel., 1907. Type. Mexico. Haenke s.n. (PRC ?). Tectaria mexicana (Fe´e) C. V. Morton var. pilosula C. V. Morton, Amer. Fern J. 56: 137. 1966. Type. Mexico. Sinaloa: Banks at Panuco, Pennell 20009 (US!).

Rhizomes short-creeping to erect, woody, mostly 1–2 cm diam.; rhizome scales concolorous, brown, lanceolate-subulate, (5–)6–9 mm long, surfaces and margins densely hairy; fronds mostly 40–100 cm long; stipes shorter than or about equaling blade length, stramineous to usually reddish brown, with scattered to dense hairs 0.1–0.3 mm long; blades chartaceous, bipinnate-pinnatifid to subtripinnate at the base, deltate, 20–50 cm long; rachises densely pubescent with hairs 0.1–0.3 mm long and scattered hairs 0.5–0.8 mm long, buds absent; pinnae 3–6 pairs, only the distal ones decurrent, proximal pair deltate, 12– 38 ⫻ 7–30 cm, long-stalked (to 4 cm), margins crenate with ciliate sinuses (hairs 0.3–0.8 mm long), basal basiscopic pinnule of proximal pinna pair the largest, to 22 cm long; vein areoles mostly elongate-pentagonal, free included veinlets usually absent; indument abaxially of glandular hairs 0.1–0.3 mm long on costae and costules, veins and margins with scattered hairs 0.3– 0.5(–0.8) mm long, or hairs absent, indument adaxially of dense hairs 0.1–0.3 mm long on costae and costules, veins with hairs 0.1–0.3 mm long, laminar surfaces in areoles glabrous or with one or more jointed hairs per areole; sori round, in 2 rows, one on each side of costule; indusia persistent, round, with a deep sinus and rounded, overlapping lobes, thus appearing peltate, remaining membranous and not thickened at maturity, margins ciliate; 2n⫽80 (Chis).

Distribution. Wet forests; 50–1600(–2000) m. Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Col. Moran (in Davidse et al., 1995) also included western Ecuador in the range. Selected Specimens Examined. Chis (Breedlove 57030, CAS). Gro (Hinton 11344, LL, NY; Martı´nez S. 5541, CAS, ENCB, MEXU). Jal (Mexia 1493, CAS, UC). Mich (Hinton 16039, UC, US). Nay (Palmer 1946, DS, NY, UC, US). Oax (Croat & Hannon 65542, UC). Sin (Ortega 6182, US). Ver (Purpus 4341, UC, US; Reeves R5909, ASU). Unverified, Doubtful, or Mistaken Reports. Tab (reported by Magan˜a, 1992, but not verified).

The names T. dilacerata (Kunze) Maxon and T. cicutaria (L.) Copel. have been misapplied to this species in Mexico and Central America (see Morton, 1966), but the former was regarded by Morton as a synonym of the latter, which is now thought to be confined to the Greater Antilles. The bipinnate-pinnatifid fronds, concolorous rhizome scales, ciliate pinna margins (especially in the sinuses), and lack of free included veinlets will distinguish this from other species of Tectaria in Mexico; the non-peltate indusia also separate it from T. heracleifolia. Plants with pilose blades, var. pilosula C. V. Morton, have been collected in Sinaloa, Guerrero, and Veracruz, but do not seem to differ in any other significant way. Tectaria rivalis (Mett. ex Kuhn) C. Chr., from Guatemala to Ecuador, is similar to T. mexicana in blade dissection (2-pinnate-pinnatifid), but has included free veinlets in the areoles. Tectaria rivalis (Mett. ex Kuhn) C. Chr., known from Guatemala and Belize to Panama, Colombia, and Ecuador, is very similar to T. mexicana in blade dissection (2- pinnate-pinnatifid), but the latter differs primarily in having free, included veinlets in many areoles.

5. TECTARIA PANAMENSIS Tectaria panamensis (Hook.) R. M. Tryon & A. F. Tryon, Rhodora 83: 136. 1981. Fig. 294A. Dictyoxiphium panamense Hook., Gen. Fil., t. 62. 1840. Lindsaea panamensis (Hook.) Mett., Fil. Hort. Bot. Lips. 105. 1856. Type. Panama. Isthmus, Pacific coast, Cuming 1124 (K!–2 sheets).

Rhizomes short, erect, 0.5–1 cm diam.; rhizome scales dark brown, lanceolate to ovate, 3–8 ⫻ 0.8–1.5 mm, remotely dentic-

614

TECTARIA

ulate; fronds clumped; stipes brown, 2–7 cm long, 1⁄10 or less the frond length, with scattered scales like those of rhizomes; blades firmly chartaceous, dimorphic, fertile longer than the sterile, linear-oblong, margins entire, (40–)70–110 ⫻ 1–3.5 cm, sterile blades narrowly elliptic, (26–)45–60 ⫻ 4–6.5 cm, attenuate at bases and apices, margins entire, usually revolute; rachises (midribs) with sparse scales at the bases, otherwise glabrous, buds absent; pinnae none; vein areoles obscure, with free included veinlets extending in all directions; indument absent on both sides of blades; sori continuous (coenosoral) and submarginal on the distal half of the fertile blades; indusia linear, continuous, 0.3–0.5 mm wide, entire or very sparingly and minutely ciliate; 2n⫽80 (CR).

Ⳳ equaling blade length, reddish brown, glabrescent or minutely hairy, hairs 0.1–0.2 mm long; blades chartaceous, 1pinnate, to 60 cm long; rachises densely hairy with hairs 0.1–0.3 mm long, buds absent; pinnae ca. 4–6 lateral pairs, basal pair largest, to 40 cm long, short-stalked (to ca. 5 mm), with 2–3 prominent, basal, basiscopic lobes, with or without exaggerated acute acroscopic lobes, rest of pinna margins serrately lobed on both sides of the costae; vein areoles pentagonal to hexagonal, commonly with free included veinlets; indument abaxially of moderate to dense glandular hairs 0.1–0.2 mm long on costae and veins, similar hairs also on leaf tissue, adaxially with similar hairs confined to the costae and costules, veins sparingly hairy to glabrous, laminae glabrous; sori round, in 2–4 rows between the main lateral veins, often more irregular than in T. incisa or T. heracleifolia; indusia persistent, 1–1.5 mm diam., round to reniform, with a basal sinus, lobes overlapping, indusia thus appearing peltate, margins and sometimes surfaces glandular-hairy; 2n⫽160 (Trin).

Distribution. Terrestrial in wet woods; 100–300 m. Mexico; Guat, Bel, Hond, Nic, CR, Pan; Col, Ec (Holm-Nielsen et al. 25765, UC). Selected Specimens Examined. Chis (Breedlove 34167, DS, ENCB, MEXU, NY; Martı´nez S. 8252, ARIZ, NY, 10835, ENCB, LL, MEXU, 18250, 25029, MEXU; Matuda 3606, MEXU, US). Oax (Cruden 1112, UC; Va´zquez T. et al. 1555, CHAPA). Ver (Brigada Dorantes 2451, MEXU, XAL; Riba 812, XAL).

For a discussion of relationships, see comments on Dictyoxiphium following the generic description of Tectaria. Tectaria panamensis differs from all other congeners in Mexico by the simple, undivided, strongly dimorphic blades and linear, submarginal sori.

6. TECTARIA TRANSIENS Tectaria transiens (C. V. Morton) A. R. Sm., Amer. Fern J. 70: 27. 1980. Figs. 293D, E. Tectaria incisa Cav. subsp. transiens C. V. Morton, Amer. Fern J. 56: 133. 1966. Type. Mexico. Veracruz: Cordoba, Finck 57 (US). Tectaria ramkissoonii Jermy & T. G. Walker, Bull. Brit. Mus. (Nat. Hist.), Bot. 13: 272. 1985. Type. Trinidad. Arima Ward. Aripo Heights, Walker T7088 (BM; isotypes TRIN, herb. Walker).

Rhizomes erect, stout, woody, mostly 1–2 cm diam.; rhizome scales dark brown, concolorous, lanceolate to ovate-lanceolate, 5–7 ⫻ 1–2.5 mm, surfaces glabrous or nearly so, margins entire to minutely and sparingly ciliate; fronds to ca. 130 cm long; stipes

Distribution. Terrestrial in wet forests; 850–1750 m. Mexico; Guat, Hond (Lagos-Witte et al. 77, MO), Nic, CR, Pan; Hisp; Trin, Ec, Peru. Moran (in Davidse et al., 1995) also included El Salvador and Colombia in the range. Selected Specimens Examined. Chis (Breedlove 26054, 32953, 38941, DS). Oax (Martı´nez S. 8748, MO, Wendt et al. 4681, MO, NY). Qro (Carranza 1264, IEB; Rzedowski 42807, IEB). Ver (Calzada 8324, XAL; Sa´nchez 63, UC). Unverified, Doubtful, or Mistaken Reports. Tab (reported by Magan˜a, 1992, but not verified).

This species is somewhat intermediate in morphology between T. heracleifolia and T. incisa (see Smith, 1981, for further discussion) but differs from both species (as well as T. vivipara) in its hairy laminar tissue abaxially, broad (1–2.5 mm wide), papillate-ciliate rhizome scales, and occurrence at higher elevations (1050–1750 m in Oaxaca, 1250–1700 m in Chiapas). It can be distinguished from T. heracleifolia by having 4–6 pairs of lateral pinnae, cuneate-based terminal segments, decurrent distal pinnae, relatively smaller acroscopic lobes on the basal pinnae, and reniform indusia. Unlike T. incisa, T. transiens has serrately lobed pinnae and 2–3 large basiscopic lobes on the proximal pinnae.

TERPSICHORE

7. TECTARIA TRICHODES Tectaria trichodes (C. V. Morton) A. R. Sm., comb. nov. Figs. 293F, G. Tectaria heracleifolia (Willd.) Underw. var. trichodes C. V. Morton, Amer. Fern J. 56: 126. 1966. Type. Guatemala. Alta Verapaz: Cerro de Agua Tortuga (Sahococ), near Cubilgu¨itz, Steyermark 44586 (US!).

Differs from T. heracleifolia by the dense, silky, spreading, septate hairs mostly 0.5–1.5 mm long on the rachises, costae, veins, and leaf surfaces between veins on both sides of the blades. Blades simple, lobed at the base, or 3-foliolate.

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four ranks of sori. The indusia are large, appearing round but actually with a narrow sinus and overlapping lobes, and the blade is glabrous or with minute glandular hairs less than 0.1 mm along the costae abaxially; costae adaxially are glabrous. Morton has written the name Pleuroderris michleriana [⫽ Tectaria incisa ⫻ panamensis] on the US sheet in 1963, and the Mu¨nch collection is somewhat similar to that in blade dissection but not in soriation. Because no other material of T. buchtienii has been seen from Mexico, or, for that matter, from Mesoamerica, the Antilles, or northern South America, it may be that this specimen is mislabeled, and we prefer to exclude the species from the flora. The type is from Bolivia, and the species also occurs in southern Brazil.

1 1 9 . T E R P SI C H O R E Terpsichore A. R. Sm., Novon 3: 479. 1993. Type: Terpsichore asplenifolia (L.) A. R. Sm. [⬅ Polypodium asplenifolium L.].

Distribution. Apparently terrestrial (ours) or on limestone rocks or cliffs, ca. 100 m (300–900 in Guat). Mexico; Guat. Specimen Examined. Tab (Zamudio 193.5, ENCB).

This species differs from all others in Mexico and the Neotropics by its dense pubescence on both sides of the blades. Such hairs are entirely absent in T. heracleifolia, or hairs, if present, are very short (0.1–0.2 mm) and sparse to only moderately dense. The hairs in the single specimen seen from Mexico are so dense that they give the blade a rusty appearance (because of the reddish septae in the hairs). We see no evidence of hybridization (morphological intermediacy) between T. heracleifolia and T. trichodes in either southern Mexico or Guatemala. Aside from the relationship to T. heracleifolia, T. trichodes seems close to T. pilosa (Fe´e) R. C. Moran from Costa Rica, Panama, and southern Brazil, and also T. pubens R. C. Moran from Ecuador and Peru. Tectaria trichodes differs from both of these in the apical segment truncate or slightly cordate at the base and leaf tissue more densely pubescent, especially on the adaxial surfaces. Excluded Species Tectaria buchtienii (Rosenst.) Maxon [syn. ⫽ Aspidium plumieri C. Presl var. brasiliense Rosenst.]. A single specimen of what appears to be this species is labelled as from Chiapas, Mu¨nch 100 (DS, US). This species has long-decurrent pinnae, so much so that the rachis is sometimes completely winged (as at DS). Sori are in two ranks between the main lateral veins, but not as regularly as in T. incisa; sometimes, there are three or even

Epiphytic, rarely epipetric or terrestrial; rhizomes shortcreeping to ascending, generally weakly dorsiventral to radially symmetric; rhizome scales brown to blackish, infrequently orangish, dull to usually lustrous, non-clathrate, glabrous or with variously colored setulae; fronds monomorphic, determinate or indeterminate; stipes articulate or not, nearly absent to equaling the blades in length; blades pinnatisect to 1-pinnate, rarely 1pinnate-pinnatifid, forking in a few species, gradually tapering distally; indument on rachises, costae, veins, and sometimes laminar tissue (both sides) of reddish brown to tawny or whitish setae 0.5–3 mm, also sometimes of branched or unbranched hairs 0.1–0.2 mm, these sometimes glandular; hydathodes present adaxially, sometimes producing calcareous secretions (lime-dots); veins simple, pinnate from the costae, free, hidden or easily visible; sori round, not sunken into the laminae, exindusiate; soral paraphyses absent; sporangia glabrous or setose on the capsules; spores tetrahedral-globose, oblate, or reniform, with a trilete or monolete scar; x⫽37. Terpsichore comprises about 50 neotropical species and one, T. elastica (Bory ex Willd.) A. R. Sm., in Africa, Madagascar, and the Mascarenes. Characteristics defining Terpsichore include the presence of rather conspicuous hydathodes on the adaxial surface of the blades (thus, differing from Ceradenia and Enterosora); reddish, atropurpureous, tawny, or hyaline setae often greater than 1 mm long on the stipes, rachises, costae, veins, and sometimes laminar tissue (thus, differing from Lellingeria); absence of glandular paraphyses in the sori; non-clathrate, usually castaneous or atropurpureous, concolorous rhizome scales that are usually setulose along the margins, or at least at tips (thus, differing from Melpomene); rhizomes generally weakly dorsiventral to occasionally radially symmetric; more than one (often 8–40) sorus per segment or pinna (thus, differing from Micropolypodium); and free, unbranched veins from the costae. Based on molecular data using rbcL and atpB genes, Terpsichore may be polyphyletic, as currently construed, although the species groups defined by Smith (1993) are quite likely mono-

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phyletic (Ranker et al., unpubl. data). Group 1, in Mexico, contains only T. asplenifolia, and this group forms one of the basalmost elements in Grammitidaceae, a family probably of Neotropical origin. Group 2, the T. taxifolia group, comprises Terpsichore semihirsuta and T. anfractuosa, among many others, and is very likely monophyletic; T. subtilis (Group 4) is closely allied to, and probably inseparable from, Group 2, on the basis of both morphological and molecular evidence. Nearly all mem-

bers of this clade have black clavate fungi on the blades, the ascomycete genus Acrospermum. Group 3 comprises the remaining five Mexican species T. cultrata, T. delicatula, T. mollissima, T. senilis, and T. spathulata, and this group forms a separate clade, in preliminary molecular analyses. Reference Smith, A. R. 1993. Terpsichore, a new genus of Grammitidaceae (Pteridophyta). Novon 3: 478–489.

Key to the Mexican Species of Terpsichore 1. Stipes well developed, mostly 6–30 cm long; blades determinate, arching or suberect; setae at segment margins solitary (unpaired). 2. Sporangia setulose; stipes atropurpureous, lustrous; rachises and costae abaxially lacking black clavate fungi; lime-dots absent adaxially. ..................................................................................................................................................................................................... 2. T. asplenifolia. 2. Sporangia glabrous; stipes reddish brown, dull; rachises and costae abaxially with black, clavate fungi; lime-dots present adaxially. .................................................................................................................................................................................................... 6. T. semihirsuta. 1. Stipes very reduced, less than 3 cm and often less than 1 cm long (except some specimens of T. spathulata); blades indeterminate, usually laxly pendent; setae at segment margins solitary, paired, or in clusters. 3. Blades 1-pinnate proximally; pinnae short-stalked to 1 mm, at least the proximal ones; sporangia glabrous. ............................. 8. T. spathulata. 3. Blades pinnatisect, if 1-pinnate then pinnae partially or entirely adnate, not short-stalked; sporangia glabrous or setose. 4. Stipes and blades densely glandular-pubescent, the glands oblong, generally not branched; sporangia lacking hairs. ......... 4. T. delicatula. 4. Stipes and blades lacking glands or, if glands present, these generally sparse, often at the tips of branched hairs; sporangia bearing hairs (except T. subtilis). 5. Hairs of laminae and rachises abaxially all dark red-brown; rachises often with black clavate fungi abaxially; sporangia glabrous; new plants formed from filamentous stolon-like roots. ......................................................................................... 1. T. anfractuosa. 5. Hairs of laminae and rachises abaxially, at least some, hyaline or hyaline at tips; rachises lacking black clavate fungi abaxially (except T. subtilis); sporangia glabrous or setose; new plants not arising from stolon-like roots. 6. Setae at segment margins solitary, not in pairs or threes; hydathodes producing lime-dots adaxially; sporangia glabrous; blades abaxially bearing black clavate fungi. ........................................................................................................... 9. T. subtilis. 6. Setae at segment margins paired or in threes; hydathodes not producing lime-dots adaxially; sporangia setose; blades lacking black clavate fungi. 7. Rhizome scales absent; 350–1350 m. ............................................................................................................................... 5. T. mollissima. 7. Rhizome scales present; 1400–2800 m. 8. Pinnae generally (0.8–)1.2–2 ⫻ 0.4–0.8 cm, length 2–4 times the width, short- to long-decurrent, adnate to rachises 2⁄3 or more of their width; blade surfaces and rachises generally with a mixture of solitary, paired, and sessile-stellate setae; sporangial setae 1–2 times length of capsules. .................................................................. 3. T. cultrata. 8. Pinnae usually 0.5–1 ⫻ 0.2–0.4 cm, length to twice the width, not or shortly decurrent, adnate to rachises 1⁄3– 1 ⁄2 their width; blade surfaces and rachises generally with solitary setae; sporangial setae 3–5 times length of capsules. .............................................................................................................................................................................. 7. T. senilis.

1. TERPSICHORE ANFRACTUOSA Terpsichore anfractuosa (Kunze ex Klotzsch) B. Leo´n & A. R. Sm., Amer. Fern J. 93: 86. 2003. Figs. 296F, G. Polypodium anfractuosum Kunze ex Klotzsch, Linnaea 20: 375. 1847. Ctenopteris anfractuosa (Kunze ex Klotzsch) Copel., Philipp. J. Sci. 84: 431. 1955 [1956]. Grammitis anfractuosa (Kunze ex Klotzsch) Proctor, Rhodora 63: 35. 1961. Melpomene anfractuosa (Kunze ex Klotzsch) A. R. Sm. & R. C. Moran, Novon 2: 429. 1992. Type. Venezuela. Me´rida, Moritz 330 (B!; isotype US!). Polypodium saxicola Baker, J. Bot. 15: 264. 1877 [“saxicolum”], hom. illeg., non Swartz, 1817. Polypodium induens Maxon, Bull. Torrey Bot. Club 32: 75. 1905. Type. Jamaica. Mar 1877, [Jenman 84] (K).

Rhizomes ascending, proliferating by root buds, plants connected by slender (0.1–0.3 mm wide) stolon-like roots; rhizome scales clathrate, 0.5–0.8 mm long, entire; fronds clumped; stipes 1 ⁄10–1⁄8 the frond length, dark gray to black, very narrowly winged, with stiff reddish brown setae 0.5–0.8 mm long; blades pinnatisect, linear-elliptic, apices subcaudate, 7–12 ⫻ 0.8–1.4

cm, rachises abaxially with stiff, reddish setae 0.5–0.8 mm long, also with dense hairs 0.1 mm long; segments oblong to deltate, alternate (thus giving the blades a zig-zag appearance), ca. 15–25 pairs, largest 7–9 ⫻ 2–2.5 mm at the dilated bases, rounded to acute at tips; costae and veins hidden and inconspicuous; hydathodes not producing lime-dots; indument abaxially of rare, twisted, branched hairs 0.3 mm long on abaxial laminar surfaces, glabrous abaxially except for short hairs on rachises and scattered, straight, red-brown hairs 0.3–0.5 mm long on margins in distal 1⁄3 of pinnae; costae black, veins obscure, pinnate; sori with many, straight, reddish brown hairs 1–1.5 mm long; sporangia glabrous; spores tetrahedral-globose.

TERPSICHORE CULTRATA

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abaxially, setae solitary (unpaired) on segment margins, indument also of minute hairs 0.1 mm; sori to ca. 12 pairs per segment, receptacles glabrous; sporangia with 1 or 2 setae 0.2–0.3 mm long on capsules; spores tetrahedral-globose; 2n⫽74 (Jam).

Distribution. Epiphytic in wet woods; 1200–2300 m. Mexico; Guat, Hond, Salv, Nic, CR, Pan; Gr Ant, Guadeloupe; Col, Ven, Guy, Ec, Peru, Bol. Selected Specimens Examined. Chis (Breedlove 25423, MEXU, 30159, 31872, DS, 32644, DS, NY, 33585, 34598, DS). Oax (Mickel 977, ISC, 4292, US, 5562, ENCB, NY, UC, 6771, NY, UC).

Black, club-shaped fruiting bodies of the ascomycete, Acrospermum maxonii Farlow in Riddle are visible on most specimens. Terpsichore anfractuosa was recently transferred to Terpsichore on the basis of unpublished molecular evidence (Ranker et al., ined.) allying it with T. semihirsuta and other fungusbearing species, rather than with Melpomene. The combination of small frond size, proliferous stolon-like roots giving rise to new plants, small rhizome scales, and the usual presence of fungal fruiting bodies along the rachises and sometimes in the sori characterize this species.

2. TERPSICHORE ASPLENIFOLIA Terpsichore asplenifolia (L.) A. R. Sm., Novon 3: 485. 1993. Figs. 297J, K. Polypodium asplenifolium L., Sp. Pl. 2: 1084. 1753. Ctenopteris asplenifolia (L.) Copel., Gen. Fil. 219. 1947. Grammitis asplenifolia (L.) Proctor, Brit. Fern Gaz. 9: 76. 1962. Type. Petiver, Pter. Ame´r., t. 7, f. 16, 1712, which is a redrawing of Plumier, Traite´ Foug. Ame´r., pl. 102A, 1705, representing a plant from Martinique. Polypodium laxifrons Liebm., Mexic. Bregn. 204 (reprint 52). 1849. Lectotype (chosen by Smith, 1981: 120). Mexico. [Veracruz:] Mirador, Liebmann s.n. [Fl. Mex. 232] (C!; isotypes K, US!, frag. NY!).

Rhizomes short-creeping; rhizome scales ferrugineous, lustrous, 0.5–2 mm long, densely setose on margin and surfaces; fronds determinate, mostly 25–75 cm long; stipes dark-castaneous to atropurpureous, lustrous, 10–30 cm long, 0.3–0.8 mm diam., 1⁄3– 1⁄2 the frond length, with reddish setae 1.5–3 mm long, sometimes glabrescent; blades pinnatisect, linear-lanceolate, 32–45 ⫻ 2.5– 4(–5.5) cm, abruptly narrowed at base, lacking black clavate fungi; segments 24–44 pairs, 14–30 ⫻ 7–10 mm at the dilate base, acute at tips; costae and veins hidden or inconspicuous; hydathodes not producing lime-dots; indument of setae 0.8–2 mm long on rachises and sometimes sparsely on costae and veins

Distribution. Epiphytic, pendent (often with the stipe apex abruptly bent downward) on trunks in wet montane forests; 900–2350 m. Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Ec. Selected Specimens Examined. Chis (Breedlove 32415, DS, 56491, CAS, ENCB; Dressler 1602, US). Gro (Langlasse´ 898, MEXU, US; Lorea 2002, IEB, 3030, IEB, UAMIZ). Oax (Mickel 1012, US, 5724, ENCB, UC, US). Ver (Pringle 8158, LL, MO, US; Ventura A. 5119, ARIZ, ENCB, 11610, IEB, MO).

Determinate fronds, solitary (unpaired) setae on segment margins, and long stipes distinguish T. asplenifolia from T. cultrata. Smith (1981) and Mickel and Beitel (1988) cited Michoaca´n in the range, but we are now unable to confirm this. Specimens cited or determined as this species from Peru (Tryon & Stolze, 1993), Brazil, and Bolivia appear to be mostly, if not entirely, T. chrysleri (Copel.) A. R. Sm. Terpsichore lehmanniana (Hieron.) A. R. Sm., from Guatemala and Belize to Ecuador, is closely related to T. asplenifolia and differs in the dull (vs. lustrous) rhizome scales that are densely setulose on the surfaces and margins, and the setose blades abaxially.

3. TERPSICHORE CULTRATA Terpsichore cultrata (Willd.) A. R. Sm., Novon 3: 486. 1993. Figs. 297A, B. Polypodium cultratum Willd., Sp. Pl., ed. 4, 5(1): 187. 1810. Ctenopteris cultrata (Willd.) Copel., Gen. Fil. 219. 1947. Grammitis cultrata (Willd.) Proctor, Rhodora 63: 35. 1961. Type. Presumably Jamaica. Swartz s.n. (S; isotype BWilld. 19674!, photo US!).

Rhizomes very short-creeping to suberect; rhizome scales castaneous, lustrous, 1–1.5 mm long, setose; fronds indeterminate, mostly 12–40 cm long; stipes gray-brown to blackish, dull, 1–2 cm long, 0.5 mm diam., with dense hyaline to pale yellow setae 2–2.5 mm long; blades pinnatisect to fully pinnate, linear, 6–38 ⫻ 1.8–4 cm, gradually tapering to both ends, lacking black cla-

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vate fungi; segments 13–52 pairs, (8–)12–20 ⫻ 4–8 mm at base, broadly adnate but less so on acroscopic edge, rounded to subacute at apices; costae flexuous, brownish nearly to apices; veins not or faintly visible; hydathodes not producing lime-dots; indument of pale yellowish to slightly reddish setae 0.8–1.5 mm long on rachises, costae, veins, and laminar tissue, setae abaxially along rachis and on segment margins solitary, paired, or in fascicles of 3, also of scattered branched hairs to 0.3 mm; sori 3–7 pairs per pinna, receptacles glabrous; sporangia with several setae 0.3–0.6 mm long on capsules; spores ellipsoid or tetrahedraloblate, monolete or trilete.

Polypodium delicatulum M. Martens & Galeotti, Me´m. Foug. Mexique 35, pl. 7, f. 1. 1842. Ctenopteris delicatula (M. Martens & Galeotti) J. Sm., Bot. Voy. Herald 227. 1854. Grammitis delicatula (M. Martens & Galeotti) Proctor, Brit. Fern Gaz. 9: 219. 1965. Type. Mexico. Oaxaca: Llano Verde, Galeotti 6378 (BR!, numbered “6328,” photo US!; isotypes BR–2 sheets, numbered 6378, K, frag. US!).

Rhizomes ascending; rhizome scales brown, lustrous, 0.3–0.5 mm long, densely covered with very short glandular hairs; fronds indeterminate, mostly 4–15(–20) cm long; stipes gray-brown to black, dull, to 1.5 cm long, with reddish setae 0.8–1 mm long and clear, blunt hairs 0.3 mm long; blades pinnatisect, linearlanceolate, 6–15(–20) ⫻ 0.6–2 cm, with to ca. 6 pairs of gradually reduced proximal pinnae; segments (20–)24–46(–55) pairs, 4–12 ⫻ 1.5–2.5 mm, bases broadly adnate, slightly decurrent, apices acute; costae black, flexuous; veins hidden; hydathodes not producing lime-dots; indument of two types, reddish setae ca. 0.8–1 mm long and more copious, pale, ca. 3-celled glandular hairs ca. 0.1–0.3 mm long; sori mostly 5–7 pairs per pinna; sporangia glabrous; spores oblate, trilete.

Distribution. Pendent epiphyte on trunks (often on tree ferns) or occasionally on wet rocks in wet montane forests, cloud forests; 1400–2800 m. Mexico; Guat, Hond, Salv, CR, Pan; Cuba, Jam, Hisp; Col, Ven, Ec, Peru, s Braz, Bol. Selected Specimens Examined. Chis (Breedlove 19946, DS, MEXU, 29971, 33641, DS, MEXU, 55544, CAS). Gro (Hinton 14280, ARIZ, ENCB, LL, MO, NY, TEX, US; Rzedowski 18612, ENCB). Oax (Mickel 4271, 5295, 6743, NY, UC; Smith 453, UC).

The shorter stipes, longer sporangial setae, clustered setae on segment margins, and often pinnate blades pinnae distinguish T. cultrata from T. asplenifolia. The presence of rhizome scales and larger pinnae distinguish it from T. mollissima. Terpsichore senilis differs in having longer, more numerous sporangial setae and more distant, smaller pinnae that are contracted to half their width at their bases. A chromosome report, n⫽37, for Ctenopteris cultrata in Costa Rica was published by Evans (Caryologia 16: 671–677. 1963), but we identify the voucher (UC 58-234-1) as Terpsichore lanigera (Desv.) A. R. Sm., a close relative not known from Mexico. A second voucher (UC 58-233-1) cited for another count (also n⫽37) for Ctenopteris cultrata by Evans, supposedly in UC, has not been found.

4. TERPSICHORE DELICATULA Terpsichore delicatula (M. Martens & Galeotti) A. R. Sm., Novon 3: 486. 1993. Figs. 295F, G.

Distribution. Pendent epiphyte or, less often, on rocks in wet pine-oak forests, cloud forests; 1500–3200 m. Mexico; Guat, Hond (Moran 5610, MO); Jam. Selected Specimens Examined. Chis (Breedlove 22927, DS, MEXU, 40450, DS, MEXU, MO). Gro (Lorea 4208, IEB; Martı´nez S. 890, MO). Hgo (Gimate 808, MEXU, NY; Pringle 13494, US). Oax (Mickel 4893, NY, UC; Smith 471, UC). Ver (Bohs et al. 1777, XAL; Mu¨ller 137, NY).

The dense glandular hairs of the blades, stipes, and rhizome scales easily distinguish T. delicatula. The closest relative is T. subscabra (Klotzsch) B. Leo´n & A. R. Sm. [syn.: T. jamesonioides (Fe´e) A. R. Sm.], from Costa Rica to Peru, which differs in lacking reddish setae on the blades.

5. TERPSICHORE MOLLISSIMA Terpsichore mollissima (Fe´e) A. R. Sm., Novon 3: 487. 1993. Fig. 295E. Polypodium mollissimum Fe´e, Me´m. Foug. 11: 47, t. 12, f. 2. 1866. Grammitis mollissima (Fe´e) Proctor, Rhodora 63: 35. 1951. Ctenopteris mollissima (Fe´e) Copel., Philipp. J. Sci. 84: 423. 1955 [1956]. Type. Guadeloupe. 1861, L’Herminier s.n. (presumably P; isotypes K!, MO, frags. NY!, US!).

TERPSICHORE SEMIHIRSUTA

Rhizomes suberect; rhizome scales absent, but with dense, tan to ferruginous setae 1–1.5 mm long; fronds indeterminate, less than 20(–30) cm long; stipes blackish, less than 1(–2) cm long and 0.3 mm diam., with dense, whitish setae 1–1.5 mm long; blades pinnatisect or fully pinnate proximally, linear, 9–15 ⫻ 0.8– 1.5 cm, gradually tapering proximally, lacking black clavate fungi; segments mostly 25–50 pairs, 4–7 ⫻ 1.5–3 mm, bases adnate, decurrent basiscopically, apices rounded to subacute; costae and veins obscure; hydathodes not producing lime-dots; indument of whitish to pale tan setae 0.8 mm long on rachises and laminar tissue, setae often reddish and 1–1.5 mm long at segment margins, often in clusters of 2–3; sori 1–4 pairs per segment, receptacles glabrous or setose; sporangia with setae 0.3–0.5 mm long; spores ellipsoid, monolete.

Distribution. Pendent epiphyte on trunks or epipetric on wet limestone rocks in wet montane forests, pine-oak-Liquidambar forests; 350–1350 m. Mexico; Guat, Bel, Hond, CR, Pan; Gr & L Ant; Col, Ven, Trin, Sur, Ec. Citation of this species for Nicaragua by Smith (1981) was based on the report by Go´mez (1976); we are unable to verify the occurrence of this species there.

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6. TERPSICHORE SEMIHIRSUTA Terpsichore semihirsuta (Klotzsch) A. R. Sm., Novon 3: 488. 1993. Figs. 295A–D. Polypodium semihirsutum Klotzsch, Linnaea 20: 379. 1847. Ctenopteris semihirsuta (Klotzsch) Copel., Philipp. J. Sci. 84: 450. 1955 [1956]. Grammitis semihirsuta (Klotzsch) C. V. Morton, Contr. U.S. Natl. Herb. 38: 113. 1967. Type. Peru. Ruiz & Pavo´n s.n. [Herb. Ruiz n. 9] (B!, frag. NY!; isotype US!). Polypodium semihirsutum Klotzsch var. fuscosetosum Hieron., Bot. Jahrb. Syst. 34: 575. 1904. Grammitis semihirsuta (Klotzsch) C. V. Morton var. fuscosetosa (Hieron.) Lellinger, Proc. Biol. Soc. Wash. 89: 715. 1977. Syntype. Guatemala. Quiche´: San Miguel Uspanta´n, Heyde & Lux s.n. [Donn. Sm. 3256] (isosyntype US!). Other syntypes are from Colombia and Peru (see Lellinger, op. cit.).

Rhizomes short-creeping to suberect; rhizome scales castaneous, lustrous, 5–7 mm long, margins with setulae 0.3 mm long; fronds determinate, mostly 25–55 cm long; stipes reddish brown, dull, 6–15 cm long, 1 mm diam., ca. 1⁄5–1⁄3 the frond length, with reddish brown setae 2–4 mm long; blades pinnatisect to fully pinnate proximally, linear-lanceolate, 27–60 ⫻ 3.5–7 cm, proximal 3–6 pairs gradually to subabruptly reduced, bearing black clavate fungi along rachis and costae abaxially; segments (pinnae) 45–70 pairs, to 30 ⫻ 6 mm, base adnate, apices rounded; costae and veins black, evident, veins 6–20 pairs per segment; hydathodes producing conspicuous lime-dots adaxially; indument of reddish brown, sparse setae mostly along rachis and at segment margins near apices, setae 1.5–2 mm long, laminar tissue glabrous on both sides; sori 5–15 pairs per pinna, often not close to base (first vein pairs often sterile), receptacles glabrous; sporangia glabrous; spores tetrahedral-globose; 2n⫽ca. 148 (Jam).

Selected Specimens Examined. Chis (Breedlove 29871, 32521, DS, 33033, DS, MEXU, 48308, CAS, ENCB, 57683, CAS, ENCB, NY; Martı´nez S. 18808, MEXU, TEX, UAMIZ). Oax (Hallberg 1592, NY; Vera C. 92, CHAPA, MEXU). Ver (Purpus 206, US; Va´zquez et al. 4103, XAL).

None of the Mexican and Central America specimens identified as T. mollissima in herbaria matches type material seen, and it may be that continental specimens are specifically different from those of the Lesser Antilles. For the present, we maintain usage of the name in its traditional application. Generally, T. mollissima occurs at lower elevations than allied species (T. cultrata, T. senilis). The lack of rhizome scales distinguishes T. mollissima from small forms of T. cultrata (q.v.). Terpsichore alfarii (Donn. Sm.) A. R. Sm. of southern Central America also lacks rhizome scales, but has a conspicuous hump along the acroscopic margin of each segment. Mickel 5561a (NY), cited by Mickel and Beitel (1988) as this species, is T. cultrata.

Distribution. Erect or arching epiphytes in wet montane forests and cloud forests; 1900–2700(–4100) m. Mexico; Guat, Hond, Salv, Nic, CR, Pan; Jam, Hisp; Col, Ven, Ec, Peru, s Braz, Bol. Selected Specimens Examined. Chis (Breedlove 24377, MEXU, NY, 31873, DS, MEXU, 55963, CAS; Matuda 2879, MEXU, US). Oax (Breedlove 59992, CAS; Mickel 4285, ENCB, MEXU, NY, 4902, US, 6762, ENCB, NY). Ver (Nee 23139, F, XAL). Unverified, Doubtful, or Mistaken Reports. Gro (Campos 1716, FCME, reported by Lorea-Herna´ndez & Vela´zquez M., 1998, but not verified).

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Terpsichore semihirsuta is distinguished by the pectinate fronds, lime-dots over the hydathodes adaxially, and setulose rhizome scales. Nearly all specimens in this group of Terpsichore have black, club-shaped fruiting bodies of the ascomycete Acrospermum maxonii (found also on T. anfractuosa, T. taxifolia (L.) A. R. Sm., and several related South American species of Terpsichore) along the rachises, costae, and in the sori abaxially.

7. TERPSICHORE SENILIS Terpsichore senilis (Fe´e) A. R. Sm., Novon 3: 488. 1993. Figs. 296D, E. Polypodium senile Fe´e, Me´m. Foug. 7: 60. 1857. Ctenopteris senilis (Fe´e) Copel., Philipp. J. Sci. 84: 398. 1955 [1956]. Grammitis senilis (Fe´e) C. V. Morton, Contr. U.S. Natl. Herb. 38: 103. 1967. Type. Colombia. Norte de Santander: Schlim 364 (RB). Polypodium subflabelliforme Rosenst., Repert. Spec. Nov. Regni Veg. 7: 306. 1909. Grammitis subflabelliformis (Rosenst.) C. V. Morton, Contr. U.S. Natl. Herb. 38: 104. 1967. Type. Ecuador. Mt. Abitagua, Spruce 5271 (K; isotypes NY!, US).

Rhizomes short-creeping to suberect; rhizome scales castaneous, 0.3–1.5 mm, with numerous, pale marginal setae; fronds indeterminate, 10–25+ cm long; stipes brownish, dull, 0.5–2 cm long, 0.3–0.4 mm diam., with numerous hyaline to pale reddish setae 1–2 mm long; blades 8–45 ⫻ 1–1.5 cm, 1-pinnate, gradually reduced proximally, lacking black clavate fungi; pinnae 0.5–1 ⫻ 0.2–0.4 cm, adnate 1⁄3–2⁄3 their width, the bases abruptly rounded acroscopically, short-decurrent or not decurrent basiscopically, apices obtuse; costae and veins inconspicuous or hidden; hydathodes not producing lime-dots; indument of hyaline to pale reddish setae 1–2 mm long on rachises, costae, and laminar tissue, also of paired (or in clusters of 3) hairs ca. 1–2 mm, pinna margins with solitary and paired (occasionally in clusters of 3) setae; sori glabrous or setose from the receptacles; sporangia with numerous setae ca. 0.5⫹ mm long on capsules; spores ellipsoid, monolete.

Distribution. Laxly pendent epiphyte in evergreen cloud forests; 2750 m. Mexico; Guat, Hond, CR, Pan; Col, Ven, Ec, Peru, Bol.

Specimens Examined. Chis (Breedlove 68357, CAS, UC; Mu¨nch 119, US).

Stolze (1981) and Lellinger (1989) treated this species as a synonym of T. cultrata, but T. senilis differs in a suite of characters that seem at least as significant as those that distinguish T. lanigera (Desv.) A. R. Sm. (recognized by both Tryon & Stolze, 1993, and Lellinger, 1989) from T. cultrata. Tryon and Stolze (1993) did recognize Grammitis subflabelliformis (Rosenst.) C. V. Morton as distinct; we treat this as a synonym of T. senilis. The differences between T. cultrata and T. senilis are especially evident in occasional mixed collections, e.g., Stolze 1513 (UC), Scamman & Holdridge 8097 (US), and Stork 1620 (UC, US), all from Costa Rica. In general, specimens of T. senilis have pinnae less than 1 ⫻ 0.4 cm and 1–2 times longer than wide (vs. pinnae 1–2 ⫻ 0.3–0.7 cm and 2–4 times longer than wide in T. cultrata), pinnae adnate 1⁄3–2⁄3 their width (vs. adnate more than 2⁄3 their width in T. cultrata), and surfaces of the blades and rachises abaxially and along pinna margins with setae generally solitary, almost never paired (vs. a mixture of solitary, paired, and sessilestellate hairs in T. cultrata).

8. TERPSICHORE SPATHULATA Terpsichore spathulata A. R. Sm., Novon 3: 483. 1993. Type. Mexico. Me´xico: Sierra de las Cruces, Pringle 4145b (UC!; isotypes MO!, NY!, US!). Figs. 297C–H. Rhizomes very short-creeping to suberect; rhizome scales orangish to brownish, dull, 0.3–1 mm long, somewhat obscured by dense, hyaline setae on margins and surfaces; fronds indeterminate, crowded, often more than 50 per plant, (12–)25–60 (–90) cm long; stipes brown, 2–15 cm long, 0.2–0.4 mm diam., with numerous, mostly unbranched, tawny setae 1–2 mm long; blades 1-pinnate (occasionally the rachises 1–2-bifurcate with forks ca. 30⬚ apart), linear, 10–85 ⫻ 0.4–1.2(–1.6) cm, very gradually narrowed proximally and with pinnae more widely separated, distally the pinnae often contiguous or imbricate, blades lacking black clavate fungi; pinnae often more than 50 pairs per frond, 2–5(–7) ⫻ 2–4 mm, sessile or proximal ones stalked to 2 mm, irregularly ovate, obovate, rhombic, or slightly lobed, larger ones often inequilateral; costae darkened in basal half, otherwise obscure, veins obscure; hydathodes not producing lime-dots; indument of pale setae 1–2 mm long on rachises, costae, veins, and sometimes laminar tissue, rachises abaxially and pinna margins often with paired or fasciculate pale reddish setae in addition to solitary ones, blades also with 1–3-branched hairs 0.1–0.2 mm long; sori 1–3 pairs per pinna, receptacles glabrous; sporangia glabrous; spores tetrahedral-oblate.

THELYPTERIS

621

or short-decurrent basiscopically, apices obtuse; costae and veins hidden or inconspicuous; hydathodes producing lime-dots; indument abaxially or pale reddish setae 1–2 mm on rachises, costae, veins, and laminar surfaces, adaxially with fewer but similar setae, setae on segment margins 1–2 mm long, solitary, reddish; sori mostly 2–3 per pinna, receptacles setose; sporangia glabrous; spores tetrahedral-globose.

Distribution. Pendent in moist, shaded rock crevices and on cliffs, under rock overhangs, and in shallow caves and grottoes; 2600–4100 m. Mexico; Guat. Selected Specimens Examined. Chis (Breedlove 24303, LL, MEXU, NY; Matuda 2890, ARIZ, LL, MEXU, MO, TEX). DF (Lyonnet 232, CAS, MO, NY; Rzedowski 35578, IEB). Jal (McVaugh 10043, NY, TEX). Me´x (Rzedowski 29309, ASU; Vega A. 591, CHAPA, ENCB, MEXU, NY). Mich (Leavenworth & Hoogstraal 1187, MO, NY). Pue (Reeves R5803, ASU). Ver (Nee & Diggs 24843, MO, UC).

Specimens of T. spathulata from Mexico and Guatemala have previously been determined as Grammitis heteromorpha (Hook. & Grev.) C. V. Morton. The latter species is restricted to the Andes from Colombia to Peru and differs by the larger (1–1.6 mm), black rhizome scales, thicker blades and stouter rachises, and presence of sporangial setae. In addition, specimens of Terpsichore heteromorpha (Hook. & Grev.) A. R. Sm. have blades markedly and regularly bifurcate, with the blade forks diverging 60–90⬚. Blades of T. spathulata are usually unbranched or sometimes once-forked with the branches divergent less than 60⬚ (e.g., Matuda 2890, MO). More collections have been seen of this species from Edo. Me´xico (20) than from all the other states combined (18). Curiously, the species has not yet been collected in Oaxaca.

9. TERPSICHORE SUBTILIS Terpsichore subtilis (Kunze ex Klotzsch) A. R. Sm., Novon 3: 488. 1993. Figs. 296A–C. Polypodium subtile Kunze ex Klotzsch, Linnaea 20: 375. 1847. Ctenopteris subtilis (Kunze ex Klotzsch) J. Sm., Hist. Fil. 184. 1875. Grammitis subtilis (Kunze ex Klotzsch) C. V. Morton, Contr. U.S. Natl. Herb. 38: 104. 1967. Type. Venezuela. Me´rida: Moritz 325 (B?; isotypes K! US!).

Rhizomes short-creeping to suberect; rhizome scales black or dark brown, 0.3–0.7 mm long, with sparse, pale marginal setulae; fronds determinate or indeterminate, to ca. 20 cm long; stipes black, 0.1–1.5 cm long, 0.2–0.4 mm diam., with sparse to dense, yellowish to reddish setae; blades pinnatisect or 1-pinnate proximally, 4–18 ⫻ 0.6–1.2 cm, tapering proximally with 1–5 pairs of shortened pinnae, with black clavate fungi along rachises and costae abaxially; segments 3–6 ⫻ 1.5–2.5 mm, base adnate, dilate

Distribution. Pendent epiphyte in montane rain forests; 1300– 2100 m. Mexico; Guat, Hond, CR, Pan; Col, Ven, Ec, Peru, Bol. Specimens Examined. Chis (Breedlove 32207, DS, 67840, CAS; Mu¨nch 51, DS).

Terpsichore subtilis is characterized by having small black or dark brown rhizome scales 0.3–0.7 mm long, short stipes usually less than 1 cm long, narrow, pendent blades, black clavate fungi on the blades abaxially, solitary (unpaired) setae, hydathodes producing lime-dots, and glabrous sporangia. Its closest relative is T. cretata (Maxon) A. R. Sm., from the Greater Antilles.

120. THELYPTERIS Thelypteris Schmidel, Icon. Pl. (ed. Keller) 45, t. 11, 13. Oct 1763. Type: Thelypteris palustris Schott [⬅Acrostichum thelypteris L.]. Some authors adopt Thelypteris thelypteroides (Michx.) Holub as the name for this taxon, but Tryon and Tryon (1973) argued against this. Terrestrial to rarely epipetric or epiphytic; rhizomes longcreeping to short-creeping to erect, dictyostelic; rhizome scales entire and usually ciliate on the margins and/or surfaces; fronds simple to pinnate to usually pinnate-pinnatifid, to decompound (Old World groups); stipes in cross section with two crescentshaped vascular bundles at bases, these fusing distally; blade axes (stipes, rachises, and costae) adaxially sulcate, grooves not running continuously into axis of the next order; indument of stipes, blades, and/or rachises nearly always hairy, sometimes sparingly, at least in the adaxial grooves, often on axes and leaf tissue abaxially, hairs acicular (unicellular or multicellular), hamate, or stellate; sori dorsal on the veins, occasionally submarginal, with or without reniform indusia; spores bilateral (tetrahedral in a few

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THELYPTERIS

Old World species), with a usually prominent perispore; x⫽36, 35, 34, 33, 32, 31, 30, 29, 27. In its broadest sense, Thelypteris comprises nearly 1000 species, distributed pantropically but with a few species in temperate areas. Using a broad circumscription, there are perhaps 300 species of Thelypteris in the Neotropics. In its narrowest sense, Thelypteris includes only two species, one (T. palustris) in Mexico. Distinguishing characters of Thelypteris include the stipe vasculature of two bundles (many bundles in dryopteroid ferns); acicular hairs on many parts of the frond; usually bilateral spores with a prominent perispore; and chromosome base numbers from 27 to 36 (x⫽40, 41 in dryopteroids and athyrioids). Thelypteris s.l. has been subdivided by pteridologists into natural groups variously treated as genera, subgenera, or sections. Most of the New World groups can be well circumscribed using a combination of characters, and thus could stand as genera, but we segregate only one of these, Macrothelypteris, naturalized in the New World, in this flora. For now, the interests of taxonomists and other biologists seem best served by adopting a large genus Thelypteris with several subgenera. Those subgenera occurring in Mexico can be distinguished by the key following the references. Species are individually keyed under each subgenus. Although precise relationships are still unresolved, Thelypteris s.l., in the sense applied here, and Thelypteridaceae, are hypothesized to have sister group relationships with a suite of athyrioid, blechnoid, onocleoid, and woodsioid ferns (Smith & Cranfill, 2002). The family comprises two main lineages, one phegopteroid (including Macrothelypteris, which see) the other thelypteroid (including Thelypteris s.l. and Cyclosorus s.l., the latter including the segregate genera Goniopteris, Meniscium, Steiropteris, and Stegnogramma in Mexico). All segregates possessing x⫽36 (Cyclosorus sensu Smith in Kubitzki, 1991), form a strongly supported clade in a recent analysis (Smith & Cranfill, 2002); these

species also usually have veins that either unite below the sinuses or run to the sinuses. The free-veined species groups (Thelypteris sensu Smith in Kubitzki, 1991), with lower base chromosome numbers, include both Amauropelta and Thelypteris s.str. in the Neotropics; these groups insert below Cyclosorus in the hypothetical phylogenetic tree. References Christensen, C. 1907. Revision of the American species of Dryopteris of the group of D. opposita. Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 7, 4: 247–336; Christensen, C. 1913. A monograph of the genus Dryopteris. Part. I. The tropical American pinnatifid-bipinnatifid species. Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 7, 10: 55–282; Holttum, R. E. 1971. Studies in the family Thelypteridaceae III. A new system of genera in the Old World. Blumea 19: 17–52. 1971; Holttum, R. E. 1974. Thelypteridaceae of Africa and adjacent islands. J. S. African Bot. 40: 123–168; Maxon, W. R. & C. V. Morton. 1938. The American species of Dryopteris, subgenus Meniscium. Bull. Torrey Bot. Club 65: 347–376; Riba, R. 1993. First continental record of Thelypteris guadalupensis (Wikstro¨m) Proctor in the state of Quintana Roo, Mexico. Amer. Fern J. 83: 39; Smith, A. R. 1971. Systematics of the neotropical species of Thelypteris section Cyclosorus. Univ. Calif. Publ. Bot. 59: 1–143; Smith, A. R. 1973. The Mexican species of Thelypteris subgenera Amauropelta and Goniopteris. Amer. Fern J. 63: 116–127; Smith, A. R. 1974. A revised classification of Thelypteris subgenus Amauropelta. Amer. Fern J. 64: 83–95; Smith, A. R. 1980. Taxonomy of Thelypteris subgenus Steiropteris, including Glaphyropteris (Pteridophyta). Univ. Calif. Publ. Bot. 76: 1–38; Smith, A. R. 1983. Polypodiaceae– Thelypteridoideae [Fam. 14(4)]. Flora of Ecuador 18: 1–148; Smith, A. R. & R. B. Cranfill. 2002. Intrafamilial relationships of the thelypteroid ferns (Thelypteridaceae). Amer. Fern J. 92: 131–149; Tryon, A. F. & R. F. Tryon. 1973. Thelypteris in northeastern North America. Amer. Fern J. 63: 65–76; Tryon, A. F., R. F. Tryon & F. Badre´. 1980. Classification, spores, and nomenclature of the Marsh fern. Rhodora 82: 461–474; Watkins Jr., J. E. & D. R. Farrar. 2002. A new name for an old fern from north Alabama. Amer. Fern J. 92: 171–178; Yatabe, Y., M. Takamiya & N. Murakami. 1998. Variation in the rbcL sequence of Stegnogramma pozoi subsp. mollissima (Thelypteridaceae) in Japan. J. Pl. Res. 111: 557–564; Yatabe, Y., J. E. Watkins, D. R. Farrar & N. Murakami. 2002. Genetic variation in populations of the morphologically and ecologically variable fern Stegnogramma pozoi subsp. mollissima (Thelypteridaceae). J. Pl. Res. 115: 29– 38.

Key to the Mexican Subgenera of Thelypteris 1. Venation predominantly areolate, with uniformly sized areoles (formed by cross-veins between primary veins) in rows of three or more between costae and pinna margins; pinnae entire, crenate, serrate, or shallowly lobed at the margins. 2. Stellate or furcate trichomes present, especially in adaxial grooves of rachises and on rhizome apex scales (also often on costae, veins, and occasionally tissue between veins abaxially). ................................................................................................. subg. Goniopteris (spp. 38–61). 2. Stellate or furcate trichomes absent on all parts of plant. 3. Sori round, in two rows between the primary veins (costules). .......................... subg. Goniopteris (T. ghiesbreghtii, T. hondurensis) (spp. 40, 43). 3. Sori oblong, often arcuate, in a single row between the primary veins (costules). .............................................. subg. Meniscium (spp. 62–66). 1. Venation not predominantly areolate, the veins free or running to the sinuses, if anastomosing below the sinuses then only single areoles formed below the sinuses; pinnae usually pinnatifid. 4. Stellate or furcate trichomes present, especially in adaxial grooves of rachises and on rhizome apex scales (also often on costae, veins, and occasionally tissue between veins abaxially). ................................................................................................. subg. Goniopteris (spp. 38–61). 4. Stellate or furcate trichomes absent on all parts of plant. 5. Sporangia setose and the sori elongate; distal pinnae strongly adnate to and decurrent on the rachises. .......... subg. Stegnogramma (sp. 67). 5. Sporangia not setose, the sori nearly round, oblong, or rarely elongate (T. linkiana); distal pinnae not or weakly adnate to rachises, not strongly decurrent. 6. Blades with all veins extending to the margins above the sinuses. 7. Blades narrowed proximally, with one or usually several pairs of greatly reduced pinnae, the proximal pinnae often auriculiform or glanduliform; veins simple. .................................................................................................. subg. Amauropelta (spp. 1–24). 7. Blades not narrowed proximally, lacking greatly reduced pinnae; veins of sterile blades forked. ................. subg. Thelypteris (sp. 69). 6. Blades with at least some of the proximal veins extending to, connivent at, or united below the sinuses. 8. Sori indusiate; aerophores absent. ..................................................................................................................... subg. Cyclosorus (spp. 25–37). 8. Sori exindusiate; scale-like aerophores usually at least 2 mm long at pinna bases abaxially. ......................... subg. Steiropteris (sp. 68).

THELYPTERIS SUBG. AMAUROPELTA

623

THELYPTERIS SUBG. AMAUROPELTA

large to small or absent; sporoderm evenly granular (light microscope), finely reticulate (scanning electron microscope); x⫽29.

Thelypteris Schmidel subg. Amauropelta (Kunze) A. R. Sm., Amer. Fern J. 63: 121. 1973.

Thelypteris subg. Amauropelta has about 200 species, mostly at middle to higher elevations on margins of rain forests, along trails, wet roadside embankments, and along streams, in tropical and subtropical America (Mexico, Florida, Antilles, Central America, and South America to northern Argentina and Chile), with three species on Pacific islands and a few in Africa, Madagascar, and the Mascarene Islands (Holttum, 1974; Smith, 1974). Subgenus Amauropelta is distinguished from other New World subgenera by the reduced pinnae at the bases of blades; veins reaching the margins above the sinuses; finely reticulate sporoderm; and x⫽29.

Amauropelta Kunze, Farnkra¨uter 1: 86, 109. 1843. Type: Amauropelta breutelii Kunze [⫽ Thelypteris limbata (Sw.) Proctor].

Rhizomes suberect to erect, occasionally decumbent; blades pinnate-pinnatifid (rarely pinnate or decompound), proximal pinnae reduced, sometimes nearly to rhizomes; aerophores present at bases of pinnae, or absent; veins usually simple, occasionally furcate, proximal pair from adjacent segments meeting margins of segments above sinuses; sori round to occasionally somewhat elongate, medial to submarginal on the veins, indusia

Key to the Mexican Species of Thelypteris subg. Amauropelta 1. Indusia absent, or minute and completely obscured by sessile glands. 2. Sori oblong or elongate along veins. 3. Proximal pair of veins from adjacent segments meeting the margins above the sinuses; tissue between veins with hamate hairs abaxially; pinnae incised to within 1 mm of costae. ........................................................................................................... 1. T. atrovirens. 3. Proximal pair of veins from adjacent segments meeting the margins at or near the sinuses; tissue between veins glabrous abaxially or very sparsely hairy; pinnae incised to ca. 2 mm from costae. .................................................................................. 8. T. linkiana. 2. Sori round or nearly so. 4. Hairs fasciculate or subfasciculate on costae and rachises abaxially, appearing stellate; red or orange hemispherical glands present on tissue between veins abaxially; aerophores at bases of costae to 6 mm long. 5. Hairs subfasciculate; aerophores absent at bases of costules. .............................................................................................. 12. T. nubigena. 5. Hairs fasciculate; aerophores present at bases of costules. ................................................................................................. 23. T. thomsonii. 4. Hairs not fasciculate and not appearing stellate; glands absent (present in T. loreae); costal aerophores absent or less than 1 mm long. 6. Sporangia setulose; costae and rachises abaxially minutely puberulous, the hairs less than 0.1 mm long. .................... 6. T. concinna. 6. Sporangia glabrous; costae and rachises abaxially not minutely puberulous. 7. Costae abaxially with at least a few scales, especially near bases, the scales Ⳳ clathrate. 8. Fronds ca. 15 cm long; pinnae ca. 2 cm long; reduced proximal pinnae 1–3 pairs. ......................................... 3. T. caucaensis. 8. Fronds mostly greater than 25 cm long; pinnae mostly greater than 5 cm long; reduced pinnae more than 5 pairs. 9. Hairs on costae abaxially mostly erect, to 1.5 mm long; receptacular tissue with stiff hairs like those of costae; segments Ⳳ perpendicular to costae, rounded at tips. ............................................................. 15. T. pilosohispida. 9. Hairs on costae abaxially more or less antrorse, mostly less than 0.6 mm long; receptacular tissue glabrous or with a few short hairs; segments somewhat oblique, rounded or acute at tips. ........................ 18. T. rudis. 7. Costae abaxially without scales. 10. Blades abaxially with sessile orangish to reddish glands. 11. Pinnae inequilateral; reduced proximal pinnae 3–7 pairs; fronds mostly 15–25(–40) cm long. ................. 19. T. sancta. 11. Pinnae equilateral; reduced proximal pinnae 3–20 pairs; fronds mostly 60-95 cm long 12. Reduced proximal pinnae 3–5 pairs, subabruptly reduced; costae nearly or quite glabrous abaxially; rhizomes erect. ................................................................................................................................................. 9. T. loreae. 12. Reduced proximal pinnae 10–20 pairs, gradually reduced; costae with scattered hairs abaxially; rhizomes short-creeping. ........................................................................................................................... 11. T. mortonii. 10. Blades abaxially lacking sessile glands. 13. Rachises glabrous or nearly so; costules and veins glabrous adaxially. ........................................................... 7. T. deflexa. 13. Rachises rather densely short-hairy; stiff hairs on costules and veins adaxially. ..................................... 14. T. oligocarpa. 1. Indusia present, usually not obscured by glands. 14. Hairs fasciculate or subfasciculate on costae and rachises abaxially; red or orange hemispherical glands on tissue between veins abaxially; aerophores present. ................................................................................................................................................................... see couplet (5). 14. Hairs hamate or acicular, not fasciculate; hemispherical glands present or absent; aerophores present or absent. 15. Hamate hairs on tissue between veins abaxially and/or on indusia; glands absent on tissue between veins. 16. Reduced pinnae usually 3–6 pairs; rachises often darkened, densely hairy, the hairs 0.1–0.2 mm long with a few longer ones intermixed; indusia tan. ..................................................................................................................................... 14. T. oligocarpa.

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THELYPTERIS 16. Reduced pinnae 7 or more pairs, nearly to the bases of frond; rachises stramineous, not densely hairy; indusia tan to blackish. 17. Indusia dark-castaneous to blackish. ...................................................................................................................... 10. T. melanochlaena. 17. Indusia stramineous to tan. ................................................................................................................................................ 20. T. scalaris. 15. Hamate hairs absent on tissue between veins and indusia; glands present or absent on tissue between veins. 18. Stipe and rachis hairs stiff, numerous, mostly 0.3–1.5 mm long. 19. Sori submarginal, often partially hidden by the revolute leaf margins; costal and rachial hairs septate; veins immersed abaxially, often darkened; basal basiscopic segment of each well developed pinna longer than the others, parallel to rachises. .......................................................................................................................................... 4. T. cheilanthoides. 19. Sori medial to supramedial, not hidden by the margins; hairs mostly unicellular (not septate); veins not immersed abaxially, usually slightly raised, stramineous; proximal segments of pinnae not enlarged or elongate. ............................................................................................................................................................................... 16. T. pilosula. 18. Stipes and rachises glabrous or hairs mostly less than 0.3 mm long, or, if longer, hairs flexuous. 20. Fronds mostly less than 45 cm long, with stipes 1–1.5 mm diam.; reduced proximal pinnae 1–4 pairs; rachises glabrous or minutely puberulous with hairs less than 0.1 mm long. 21. Laminae between veins abaxially lacking glands. 22. Stipes and rachises stramineous to tan; hairs absent on costae, costules, and tissue between veins abaxially. ............................................................................................................................................................... 22. T. tablana. 22. Stipes and rachises castaneous to brownish; hairs of costae, costules, and tissue between veins abaxially uniformly short, moderately dense, 0.05–0.1 mm long. .............................................................................. 24. T. venturae. 21. Laminae between veins abaxially with sessile orangish to reddish glands. 23. Hairs of the costae and rachises abaxially uniformly short, less than 0.1 mm long; stipes and rachises usually darkened; indusia reddish brown. .......................................................................................................... 5. T. cinerea. 23. Hairs of the costae and rachises abaxially absent or sparse and 0.1–0.3 mm long; stipes and rachises stramineous; indusia tan. ...................................................................................................................................... 19. T. sancta. 20. Fronds usually more than 45 cm long, with stipes more than 2 mm diam.; reduced proximal pinnae 5⫹ pairs; rachises glabrous or hairy with hairs greater than 0.1 mm (not puberulous). 24. Rachises, costae, costules, and tissue between veins abaxially with uniform short hairs 0.05–0.1 mm long; stipes and rachises castaneous to brownish. ........................................................................................................... 24. T. venturae. 24. Rachises, costae, costules, and tissue between veins abaxially glabrous or with hairs mostly greater than 0.1 mm long; stipes and rachises stramineous to tan. 25. Rhizomes short-creeping; midribs glabrous abaxially; proximal pinnae very gradually reduced, 10–20 pairs. .................................................................................................................................................................... 11. T. mortonii. 25. Rhizomes suberect to erect; midribs glabrous to hairy abaxially; proximal pinnae gradually to abruptly reduced, usually fewer than 10 pairs. 26. Rachises, costae, veins, and tissue between veins abaxially without hairs. 27. Segments oblong, not falcate, rounded at tips; indusia rather large, persistent; pinnae incised to ca. 1 mm from costae; glands absent on blades abaxially. ..................................................... 13. T. oaxacana. 27. Segments linear to linear-lanceolate, acutish at tips, subfalcate; indusia relatively small, somewhat obscured in mature sori; pinnae incised to within 0.5 mm of costae; glands present or absent on blades abaxially. ................................................................................................ 21. T. struthiopteroides. 26. Rachises, costae, veins, and tissue between veins abaxially sparsely to densely hairy. 28. Sori submarginal, often partially hidden by the revolute leaf margins; usually a few long, septate hairs on costae and rachises abaxially; veins immersed abaxially, often darkened. ...................................................................................................................................... 4. T. cheilanthoides. 28. Sori medial to submarginal, usually not hidden by segment margins; septate hairs present or absent; veins slightly raised abaxially, not darkened. 29. Pinna segments Ⳳ perpendicular to costae, occasionally oblique; pinnae usually abruptly reduced abaxially, with widely spaced smaller pinnae; segments mostly 4–6 times longer than wide. ...................................................................................................................................... 2. T. balbisii. 29. Pinna segments oblique, usually falcate; pinnae gradually reduced nearly to the bases of frond; pinna segments mostly 2–3 times as long as broad. ................................................ 17. T. resinifera.

1. THELYPTERIS ATROVIRENS Thelypteris atrovirens (C. Chr.) C. F. Reed, Phytologia 17: 261. 1968. Figs. 302D–F.

Dryopteris atrovirens C. Chr. in Christ, Bull. Herb. Boissier, se´r. 2, 7: 263. 1907. Syntypes. Costa Rica. Foreˆts de l’Achiote, Tonduz 10727 (P); Navarro, Werckle´ s.n. (P); La Luna, Werckle´ s.n. (P).

Rhizomes erect; fronds generally (35–)50–100 cm long; stipes 5–15 cm ⫻ 2–4 mm, scaly only at bases; blades mostly 40–80 cm, with 4–10 pairs of gradually to subabruptly reduced proximal pinnae; pinnae 15–30 pairs, horizontal, alternate, the medial

THELYPTERIS CAUCAENSIS

the largest, 5–10 ⫻ 1.3–2 cm, 2–3 cm distant, incised 0.8 the way to the costae, the proximal ones reduced to less than 1 cm long (often to 1 mm), 2–6 cm distant; aerophores absent at pinna bases; segments spreading to slightly oblique, 3–4 mm wide, oblong, apices rounded or obtuse, proximal pair somewhat shortened (especially the anterior one); veins 7–9 pairs per segment; indument abaxially of moderate to dense erect hairs 0.1 mm long on costae, veins, and tissue between veins, also blades with scattered hamate hairs 0.2 mm long between veins, adaxially the tissue between veins with numerous appressed hairs 0.1 mm long; sori oblong, supramedial, exindusiate; sporangia glabrous; 2n⫽58 (Chis).

625

maica, Sci. Ser. 5: 65. 1953. Neotype (chosen by Proctor, 1977: 281). Dominica. Hodge & Hodge 1203 (GH, not found, 2002). Complete synonymy given in Smith (1983).

Rhizomes erect; fronds 50 to often 100(–150) cm or more long; stipes stramineous to tan, sometimes mucilaginous when young, mostly 5–20 cm ⫻ 3–7 mm diam.; blades to 130 ⫻ 40 cm, with to 10 pairs of gradually to subabruptly reduced, widely spaced pinnae proximally; pinnae to (4–)10–16(–22) ⫻ 1.5–3 cm, deeply incised to within 1 mm of costae, usually opposite, horizontal, most pinnae with proximal segments somewhat elongate; aerophores peg-like at pinna bases; segments perpendicular or nearly so to costae, linear, 2–3 mm wide; veins to 18 pairs per segment; indument abaxially of sparse to dense hairs on rachises, costae, and costules, hairs unicellular or often septate, to 1.5 mm long, occasionally hairs absent, tissue between veins often with numerous orangish to reddish, sessile, hemispherical glands; sori medial to submarginal, with indusia persistent, thin, tan, often glandular; sporangia glabrous.

Distribution. In montane rain forests, cloud forests; 1200– 1700 m. Mexico; Guat, Bel (Holst 5756, MO), Nic, CR, Pan; Col. Specimens Examined. Chis (Breedlove 32185, 32664, 38796, DS; Mu¨nch 50?, DS). Hgo (Arreguı´n S. 1101, NY). Oax (Arreguı´n 702, ENCB, 703, CHAPA, MEXU; Mickel 986, NY). Ver (Ramı´rez 1041, XAL).

Thelypteris atrovirens differs from most Mexican species by the combination of oblong, exindusiate sori and hamate hairs on the tissue between veins abaxially. From T. linkiana it differs by the greater number of reduced proximal pinnae, lack of buds in axils of distal pinnae, and veins meeting the margins above the sinuses. The type of this species usually has been regarded as from Guatemala (Alta Verapaz: Between Sepacuite´ and Secanquin, Maxon & Hay 3281, NY!, US!), but the name was first published by Christensen in Christ, and syntypes were cited from Costa Rica; the Guatemalan specimen was not cited until a publication by Christensen (1907) later in the same year.

2. THELYPTERIS BALBISII Thelypteris balbisii (Spreng.) Ching, Bull. Fan Mem. Inst. Biol. 10: 250. 1941. Fig. 301C. Polypodium balbisii Spreng., Nov. Actorum Acad. Caes. Leop.-Carol. German. Nat. Cur. 10: 228. 1821. Aspidium sprengelii Kaulf., Flora 6: 365. 1823, nom. superfl. (see Morton, Amer. Fern J. 53: 62. 1963). Dryopteris sprengelii (Kaulf.) Kuntze, Revis. Gen. Pl. 2: 813. 1891. Dryopteris balbisii (Spreng.) Urb., Symb. Antill. 4: 14. 1903. Thelypteris sprengelii (Kaulf.) Proctor, Bull. Inst. Ja-

Distribution. In wet woods, especially along stream banks and roadsides; 100–1400 m. Mexico; Guat, Bel, Hond, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Guy, Ec, Peru, Bol. Selected Specimens Examined. Chis (Breedlove 22439, DS; Calzada et al. 4088, XAL; Matuda 2766, LL). Oax (Mickel 5183, ENCB, NY, UC, 5891, NY, US; Torres C. 11554, MEXU). Tab (Linden 1494, K, W; Rovirosa 581, NY). Ver (Nee & Taylor 26280, F, NY; Orcutt 2977, ASU, DS, NY, TEX, US).

Thelypteris balbisii is closely related to T. resinifera, and it would not be surprising to find both species growing together and hybridizing. Two collections vary in the direction of T. resinifera: Mickel 1301 (NY, UC) and Hallberg 1319 (NY, UC). Usually T. balbisii can be distinguished by the presence of a least a few long, septate hairs on the costae abaxially, by the more spreading and longer pinna segments, by its larger size, by the lack of hairs on the indusia, and by the more remote and fewer reduced proximal pinnae. See T. struthiopteroides for additional comments.

3. THELYPTERIS CAUCAENSIS Thelypteris caucaensis (Hieron.) Alston, J. Wash. Acad. Sci. 48: 233. 1958. Figs. 302H, J–L.

626

THELYPTERIS

Nephrodium caucaense Hieron., Bot. Jahrb. Syst. 34: 444. 1904. Dryopteris caucaensis (Hieron.) C. Chr., Index Filic. 257. 1905. Lectotype (chosen by Christensen, 1907). Colombia. Cauca: Lehmann 3102 (B!, isolectotype BM!).

Rhizomes suberect; fronds ca. 15(–50) cm long; stipes to ca. 5 cm ⫻ 1 mm, tan to brownish; blades subcoriaceous, 10–45 ⫻ 4– 8(–12) cm, with 1–3 pairs of abruptly reduced proximal pinnae; pinnae to ca. 10 pairs, to 2(–6) ⫻ 0.8(–1.5) cm, incised to ca. 1 mm from costae; aerophores absent; segments slightly oblique, apices obtuse to subacute, ca. 2(–4) mm wide; veins mostly 4–7 pairs per segment; indument abaxially of antrorse acicular hairs mostly 0.3–0.5 mm long on costae and rachises, the costae also with a few narrow subclathrate scales ca. 2–3 cells wide, tissue between veins glabrous and eglandulose on both sides; sori supramedial, exindusiate, sometimes with a few receptacular setae; sporangia glabrous.

3(–4) cm, incised to within 1(–3) mm of costae; aerophores prominent, tuberculiform or peg-like at pinna bases; segments with revolute margins, ca. 3–5 mm wide, proximal segments often elongate, the basiscopic one the longest and often dentate; veins (7–)10–18(–23) pairs per segment, usually immersed and darkened abaxially; indument abaxially usually of a few stiff septate hairs to 2(–2.5) mm long on rachises and costae, or sometimes glabrous or glabrescent, tissue between veins glabrous on both sides, or with sessile yellowish to orange glands; sori submarginal, often partially hidden by revolute margins, with indusia relatively large, persistent, often glandular on the margins; sporangia glabrous; 2n⫽58, 116 (Oax). Key to the Mexican Varieties of Thelypteris cheilanthoides 1. Segments falcate, with strongly revolute margins; blades usually with septate hairs abaxially. ..................... 4a. var. cheilanthoides. 1. Segments straight or subfalcate, with margins plane or slightly revolute; blades glabrous. ................................. 4b. var. mucosa.

4a. Thelypteris cheilanthoides var. cheilanthoides Figs. 298A–C. Phegopteris amplificata Fe´e, Me´m. Foug. 8: 89. 1857. Dryopteris amplificata (Fe´e) C. Chr., Index Filic. 251. 1905. Type. Mexico. [Veracruz]: Cordoba, Schaffner 219 (P? possible authentic specimen, Schaffner s.n., P!). Aspidium kunzei Fe´e, Me´m. Foug. 10: 37, t. 41, f. A, B. 1865. Type. Mexico. Veracruz: Orizaba, Schaffner s.n. (P?). Complete synonymy given in Smith (1983).

Distribution. Montane rain forests; 2750 m. Mexico; Guat, CR; Col, Ven, Ec, Peru, Bol.

Differs from var. mucosa by the characters given in the key; 2n⫽58 (Oax).

Specimen Examined. Chis (Martı´nez S. 19577, MEXU).

This species occurs at high elevations throughout its wide range. Because of the subclathrate scales on the costae abaxially, the exindusiate sori, and the abruptly reduced proximal pinnae, it is believed to be related to T. rudis, from which it differs in its small size, narrower costal scales, sparser hairs on the costae abaxially, and more coriaceous blades.

4. THELYPTERIS CHEILANTHOIDES Thelypteris cheilanthoides (Kunze) Proctor, Bull. Inst. Jamaica, Sci. Ser. 5: 58. 1953. Aspidium cheilanthoides Kunze, Linnaea 22: 578. 1849. Dryopteris cheilanthoides (Kunze) C. Chr., Index Filic. 257. 1905. Amauropelta cheilanthoides (Kunze) A˚. Lo¨ve & D. Lo¨ve, Taxon 26: 325. 1977. Type. Brazil. Regnell II 326 (LZ destroyed; isotype S).

Rhizomes erect, caudices (1–)2–4(–5) cm diam.; fronds mostly 60–150(–200) cm long, usually mucilaginous when coiled; stipes stramineous to tan, to 25 ⫻ 0.4–1 cm; blades subsucculent when living, chartaceous to subcoriaceous when dried, mostly 75–175 ⫻ 6–50 cm, with 2–5 pairs of abruptly reduced proximal pinnae, the lowermost glanduliform; pinnae (3–)10–25(–30) ⫻ (1–)1.5–

Distribution. Wet pine-oak woods and seepage areas, especially along streams and roadside ditches; 950–2700 m. Mexico; Guat, Hond, CR, Pan; Jam, Hisp; Col, Ven, Ec, Peru, s Braz, Bol. Selected Specimens Examined. Chis (Breedlove 32713, DS). Gro (Mexia 9001, GH, K, MICH, MO, NY, UC, US). Gto (Rzedowski 51229, IEB, UC). Hgo (Equipo 6, FCME). Jal (Mexia 1768, CAS, UC). Me´x (Hinton 1392, GH, MEXU, NY). Mich (Hinton 15376, GH, LL, NY, US). Oax (Mickel 4223, MEXU, NY). Pue (Ventura A. 22170, CHAPA, UAMIZ, XAL). Ver (Lira 212, MEXU, UAMIZ).

THELYPTERIS CONCINNA Unverified, Doubtful, or Mistaken Reports. Tab (reported by Magan˜a, 1992, probably based on a misidentification).

This species is distinguished from others in Mexico by the persistent indusia, somewhat succulent blades with immersed veins, and a basiscopic segment of larger pinnae somewhat elongate. Variety cheilanthoides is further distinguished from var. mucosa by the revolute segment margins and septate hairs on rachises and costae abaxially. Te´llez 7867 (MEXU) is unusual in having pinnatifid segments.

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2.5–10 cm, 1–3 pairs of proximal pinnae reduced and auriculiform; pinnae ca. 15(–30) pairs, 2–4(–6) ⫻ 1.3(–2) cm, inequilateral with basiscopic segments shorter and more oblique, falcate; aerophores absent; segments oblique, 2–3 mm wide, apices obtuse to acute; veins 3–5(–8) pairs per segment; indument abaxially absent or of minute hairs 0.1 mm long along rachises, costae, adaxially the costae setose, tissue between veins on both sides glabrous or with sessile yellowish to reddish glands abaxially; sori medial to supramedial, with indusia persistent, reddish brown, glossy, to 1 mm diam., often sessile-glandular; sporangia glabrous; 2n⫽58 (Oax, Chis).

4b. Thelypteris cheilanthoides var. mucosa A. R. Sm., Amer. Fern J. 63: 123. 1973. Figs. 298D, E. Amauropelta mucosa (A. R. Sm.) A˚. Lo¨ve & D. Lo¨ve, Taxon 26: 325. 1977. Type. Mexico. Oaxaca: Dto. Ixtla´n, along road from Ixtla´n to Tuxtepec, Km 126–129, Smith 451 (UC!).

Similar to var. cheilanthoides except: segments less falcate, with margins less revolute; indument completely absent on blades abaxially; 2n⫽116 (Oax).

Distribution. Along wet roadbanks, montane rain forests; 1600–2200 m. Mexico; Guat, CR, Pan; Col, Ven, Ec, Bol. Specimens Examined. Chis (Breedlove 23162, 24404, 31678, 32682, DS, MEXU, 23127, 32389, 42566, DS; Matuda 5118 (DS, F, MEXU, UC, US). Oax (Smith 462, UC).

Distribution. In wet woods; 1350–1900 m. Mexico. Specimens Examined. Oax (Breedlove 59843, CAS; Mickel 4317, ENCB, MEXU, NY; Smith 463, UC). Pue (Arse`ne 1935, MEXU, US). Ver (Copeland herb. 31, BM, GH, MEXU, MICH, UC, US; Ventura A. 229, DS, ENCB, NY).

5. THELYPTERIS CINEREA Thelypteris cinerea (Sodiro) A. R. Sm., Phytologia 34: 233. 1976. Figs. 199M, N. Nephrodium cinereum Sodiro, Anales Univ. Centr. Ecuador 22: 103. 1908. Type. Ecuador. Corazo´n, Aug 1903, Sodiro s.n. (not found at K or P, but authentic and topotypic specimen at P!–collected Dec 1907). Dryopteris sanctiformis C. Chr., Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 7, 10: 130, f. 12d. 1913. Thelypteris sanctiformis (C. Chr.) C. F. Reed, Phytologia 17: 312. 1968. Amauropelta sanctiformis (C. Chr.) A˚. Lo¨ve & D. Lo¨ve, Taxon 26: 325. 1977. Type. Panama. [Chiriquı´:] Volca´n Chiriquı´, Maxon 5294 (US!; isotypes NY!, P!). Complete synonymy given in Smith (1983).

Rhizomes erect; fronds fasciculate to 45 cm long; stipes brownish, lustrous, 6–12 cm ⫻ ca. 1–2 mm, glabrous; blades 12–35 ⫻

Thelypteris cinerea is easily distinguished by the combination of short, uniform pubescence less than 0.1 mm long on the rachises and costae abaxially, the relatively few pairs of reduced proximal pinnae, inequilateral pinnae, and indusiate sori.

6. THELYPTERIS CONCINNA Thelypteris concinna (Willd.) Ching, Bull. Fan Mem. Inst. Biol. 10: 251. 1941. Figs. 300D–F. Polypodium concinnum Willd., Sp. Pl., ed. 4, 5(1): 201. 1810. Dryopteris concinna (Willd.) Kuntze, Revis. Gen. Pl. 2: 812. 1891. Amauropelta concinna (Willd.) Pic. Serm., Webbia 31: 251. 1977. Type. Venezuela. Caracas, Bredemeyer s.n. (B-Willd. 19698). Phegopteris elongata E. Fourn., Mexic. Pl. 1: 89. 1872. Thelypteris concinna var. elongata (E. Fourn.) C. F. Reed, Phytologia 17: 268. 1968. Syntypes. Mexico. All apparently Veracruz, Botteri 1442 (P?, not found; possible isosyntype US), Hahn s.n. (P!), Linden s.n. (P, photo US!). Complete synonymy given in Smith (1983).

Rhizomes suberect to erect; fronds to ca. 80 cm long; stipes brownish or dark purplish, to ca. 15(–20) cm ⫻ 2–4 mm; blades chartaceous, to 60 ⫻ 28 cm, with to 12 pairs of gradually reduced proximal pinnae; pinnae to ca. 30 pairs, (3–)8–14 ⫻ (0.7–)1–2 cm, incised to ca. 1 mm from costae; aerophores absent; segments slightly oblique, apices obtuse to subacute, ca. 2–3 mm wide;

628

THELYPTERIS

veins mostly 6–8(–10) pairs per segment; indument abaxially of dense uniform-sized non-hamate hairs less than 0.1 mm long on rachises and costae, tissue between veins glabrous, eglandulose, adaxially with appressed hairs 0.1 mm long on tissue between veins, or glabrescent; sori supramedial, exindusiate; sporangia minutely setulose, hairs ca. 0.05 mm long; 2n⫽58 (Ver, Oax, Chis, Jam).

herbaceous, generally 40–80 ⫻ 5–16 cm, with 2–4(–6) pairs of reduced proximal pinnae; pinnae opposite or subopposite, mostly 2–7(–9) ⫻ 0.7–1.4(–1.8) cm, widest at bases, incised to within 1(–1.5) mm of costae; aerophores absent; segments slightly oblique, rounded at apices, ca. 2–4 mm wide; veins 4–6(–8) pairs per segment; indument abaxially absent on rachises, costae, veins, and tissue between veins or costae sparsely hairy, hairs 0.2–0.4 mm long, adaxially the blades glabrous except along rachial and costal grooves; sori medial, exindusiate; sporangia glabrous; 2n⫽116 (Oax).

Distribution. In woods, especially along trails and near streams; 700–2000 m. Mexico; Guat, Hond, Salv, Nic, CR, Pan; Gr & L Ant; Col, Ven, Ec, Peru, Bol, nw Arg. Selected Specimens Examined. Chis (Purpus 6711, GH, NY, UC, US). Coah (Lyonnet 421000004, CAS). Gro (Lorea 2964, FCME, IEB). Hgo (Gonza´lez Q. 1549, DS, MICH, NY). Jal (Mora s.n., IBUG). Mor (Lyonnet 3141, MEXU, US). NL (Pringle 1984, CAS, GH, LL, MICH, NY, UC, US). Oax (Rzedowski 32802, MEXU). Pue (McGregor 16454, MICH). Qro (Rzedowski 46625, IEB, XAL). SLP (Pringle 3395, MEXU, UC). Ver (Copeland herb. 24, BM, MICH, UC).

Thelypteris concinna is distinguished from all other members of subg. Amauropelta in Mexico by having minutely setulose sporangia. In laminar pubescence T. concinna is similar to T. cinerea, but differs from that by the exindusiate sori and a greater number of reduced proximal pinnae. It is by far most common in Chiapas, Oaxaca, and Veracruz, with more than 60 collections seen; collections from all other states in Mexico are fewer than half this number.

Distribution. In wet oak and pine-oak woods and along trails, roadside embankments; 1300–2700 m. Mexico; Guat, Hond, Salv, CR, Pan; Col, Ven, Ec, Peru, Bol. Go´mez P. (1976) also reported the species from Nicaragua, but no specimens have been seen. Selected Specimens Examined. Chis (Breedlove 21999, DS, 23153, DS, LL, 32327, DS; Gittins 4180, NY). Gro (Breedlove 36081, DS). Oax (Rzedowski 33362, UAMIZ; Smith 402, UC).

Thelypteris deflexa differs from other members of subg. Amauropelta in Mexico by the combination of exindusiate sori, nearly or quite glabrous rachises and costae abaxially, and relatively few pairs of reduced proximal pinnae. Breedlove 31739 (DS), from Chiapas, is atypical in bearing sessile reddish glands on the blades and receptacles, fibrillose scales on the costae and costules abaxially, and small aerophores at the pinna bases.

7. THELYPTERIS DEFLEXA Thelypteris deflexa (C. Presl) R. M. Tryon, Rhodora 69: 5. 1967. Figs. 300J, K. Nephrodium deflexum C. Presl, Reliq. Haenk. 1(1): 36, t. 5, f. 2. 1825. Dryopteris lindigii C. Chr., Index Filic. 275. 1905, non D. deflexa (Kaulf.) C. Chr. Thelypteris lindigii (C. Chr.) Alston, J. Wash. Acad. Sci. 48: 233. 1958. Amauropelta deflexa (C. Presl) A˚. Lo¨ve & D. Lo¨ve, Taxon 26: 325. 1977. Type. Peru. Haenke s.n. (PR?). Complete synonymy given in Smith (1983).

Rhizomes erect or suberect, caudices ca. 1–1.5 cm diam.; fronds (15–)30–60(–120) cm long; stipes stramineous with darkened bases, 5–12 cm ⫻ 1–2 mm, glabrous except for sparse scales at bases, scales tan, ovate, ca. 3 mm long, glabrous; blades thin,

8. THELYPTERIS LINKIANA Thelypteris linkiana (C. Presl) R. M. Tryon, Rhodora 69: 6. 1967. Figs. 300A–C. Gymnogramma polypodioides Link, Hort. Berol. 2: 50. 1833, hom. illeg., non Spreng., 1827. Grammitis linkiana C. Presl, Suppl. Tent. Pterid. 209. 1836. Dryopteris linkiana (C. Presl) Maxon, J. Wash. Acad. Sci. 14: 199. 1924. Type. Described from cultivated specimens (B). Gymnogramma diplazioides Desv., Me´m. Soc. Linn. Paris 6: 214. 1827 [“diplazoides”]. Dryopteris diplazioides (Desv.) Urb., Symb. Antill. 4: 21. 1903. Thelypteris diplazioides (Desv.) Proctor, Bull. Inst. Jamaica, Sci. Ser. 5: 59. 1953, hom. illeg., non (Moritz ex Mett.) Ching, 1936. Type. Hispaniola. Collector not stated (P!).

THELYPTERIS LOREAE Gymnogramma cheilosora Fe´e, Me´m. Foug. 8: 79. 1857 [“cheilosorus”]. Type. Mexico. Veracruz: near Huatusco, Schaffner 215 (P?). Complete synonymy given in Smith (1983).

Rhizomes erect; fronds (25–)60–140 cm long; stipes 15–25(–45) cm long; blades chartaceous to thick herbaceous, mostly 25–95 ⫻ 15–30 cm, with 2–6 proximal pinna pairs gradually reduced, lowermost 1–2 cm long or more, often short-stalked (ca. 1 mm), nearly always bearing proliferous buds in axils of distal pinnae; pinnae (4–)8–16 ⫻ 1–2.5(–3) cm, broadest at bases, incised 0.5– 0.8 to the costae; aerophores tuberculiform or peg-like at bases of larger pinnae; segments oblong, 3–6 mm wide, rounded to truncate at apices, spreading or slightly oblique; veins 4–8(–12) pairs, lowermost meeting margins at or slightly above sinuses; indument abaxially of scattered hamate hairs ca. 0.2 mm long on rachises, costae, and veins, tissue between veins on both sides glabrous or glabrescent, or with appressed hairs 0.1–0.2 mm long adaxially; sori oblong to linear, to ca. 2 mm long, exindusiate; sporangia glabrous; 2n⫽58 (Oax, Ver), 116 (Chis).

629

(Tlalpitzahuac), 2100 m, Lorea 4540 (holotype XAL!; isotypes FCME!, MEXU!, NY, UC!). Figs. 303A–E. Species habitu et glandibus aurantiacis sessilibus in laminis adaxialiter cum T. balbisii et T. resinifera optime congruens, sed indusiis minutis inconspicuis differt; a T. mortonii paribus paucioribus pinnarum proximalium subabrupte reductis, rhizomatibus erectis differt. (Dedicated to Dr. Francisco (Paco) LoreaHerna´ndez (1956–; Instituto de Ecologı´a, Xalapa, Mexico), keen observer and premier collector of ferns from Guerrero, as well as specialist in Lauraceae.)

Rhizomes erect, caudices to 12 cm long; fronds 65–85 cm long; stipes stramineous, mostly ca. 15 cm ⫻ 2–4 mm, bearing scattered, thin, persistent, tan, ovate, glabrous, somewhat appressed to slightly spreading scales to ca. 3 ⫻ 2 mm; blades 50–70 ⫻ 14–25 cm, with 3–5 pairs of subabruptly reduced pinnae proximally, the shortest ca. 1(–3) cm long, lacking proliferous buds; pinnae to 8–13 ⫻ 1.5–2.5 cm, deeply incised to within 1 mm of costae, usually opposite or subopposite proximally, reduced proximal pinnaedeflexed, others patent, proximal segments of individual pinnae about the same size as adjacent more distal segments; aerophores absent or tuberculate at pinna bases; segments perpendicular or slightly ascending with respect to costae, linear, 2–4 mm wide; veins ca. 10 pairs per segment; indument abaxially of sparse hairs 0.1–0.2 mm long on rachises, costae, and costules, or axes nearly glabrous, costae also with very narrow, amorphous, tan scales to ca. 0.5 mm long, 2–3 cells wide, tissue between veins with numerous orangish sessile hemispherical glands, adaxially the blades glabrous between veins; sori supramedial to submarginal, with minute indusia 0.1–0.2 mm diam., these nearly or quite obscured by orangish glands and/or mature sporangia; sporangia glabrous.

Distribution. In wet montane forests, especially along streams, wet banks; 500–2000 m. Mexico; Guat, Bel (Holland 29, UC), Hond, Nic, CR; Gr & L Ant; Col, Ven, Guy, Ec, Peru, Bol. Selected Specimens Examined. Chis (Breedlove 32219, DS, 56485, CAS, ENCB). Gro (Lorea 1522, FCME; Velazquez 285, FCME, IEB). Oax (Conrad & Conrad 3239, MO, UC). Pue (Riba 463, MEXU; Sharp 45396, TENN, US). Qro (Dı´az B. & Carranza 6518, IEB, UC). Ver (Purpus 5690, UC, US; Ventura A. 10451, ASU, ENCB, IEB).

Thelypteris linkiana is easily distinguished by the combination of exindusiate, elongate sori, glabrous sporangia, and presence of hamate hairs on the axes abaxially (see T. atrovirens for additional discussion). Nearly all collections seen from Mexico have one or more minute buds in the axil(s) of distal pinnae. The only other species of subg. Amauropelta in Mexico that has buds on the blades is T. scalaris.

9. THELYPTERIS LOREAE Thelypteris loreae A. R. Sm., sp. nov. Type. Mexico. Guerrero: Mpio. Malinaltepec, ca. 3 km S de Paraje Montero, sobre el camino a San Miguel Progreso, ladera NO del Cerro Tepilzahuac

Distribution. In montane forests, soils with abundant organic material; 2100–2150 m. Mexico. Specimens Examined. Gro (Lorea 1913, NY, UC, XAL).

This species seems to combine characters of several species in different subgroups of subg. Amauropelta. Like species of sect. Amauropelta, T. loreae bears orangish sessile glands on the tissue between veins abaxially; however, unlike most species in that section, the indusia are very small, so small that it is difficult to see them except in very young specimens. Other species belonging to sect. Amauropelta in Mexico include T. balbisii, T. resinifera, and T. struthiopteroides, all of which have relatively large,

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conspicuous indusia. Another species in this group is T. sancta, which does have very small indusia. However, it differs strongly from T. loreae in having inequilateral pinnae, much smaller blades, and in being rheophytic; it seems unlikely to be closely related. The only other species in sect. Amauropelta in Mexico is T. mortonii, an uncommon and small-indusiate species restricted to the states of Morelos and Me´xico; but T. mortonii has 10–20 pairs of gradually reduced proximal pinnae and shortcreeping rhizomes, and also seems more distantly related. In T. loreae, there are a few amorphous and fibrillose scales on the costae abaxially, but these are unlike the clathrate or subclathrate scales in species of sect. Lepidoneuron. I find no evidence of hamate or fasciculate hairs on the blade axes, eliminating sects. Uncinella and Phacelothrix as possible homes. In the sum of its characters, T. loreae could be transitional to taxa such as T. deflexa, in sect. Apelta. Species in sect. Apelta lack indusia and have relatively few reduced proximal pinnae, but they also lack laminar glands. Thus, T. loreae is placed in sect. Amauropelta with slight uncertainty, and with no obvious sister species relationships.

10. THELYPTERIS MELANOCHLAENA Thelypteris melanochlaena (C. Chr.) C. F. Reed, Phytologia 17: 292. 1968. Figs. 301K–M. Dryopteris melanochlaena C. Chr., Smithsonian Misc. Collect. 52: 384. 1909. Type. Guatemala. Alta Verapaz: Coba´n, Donnell Smith 168 (US! p.p.).

Rhizomes suberect; fronds to 110 cm long; stipes stramineous, 5–15 cm ⫻ 2.5–3.5 mm, deciduously pubescent; blades 60–100 ⫻ 8–22 cm, with 7–14 pairs of gradually reduced pinnae proximally, lacking proliferous buds; pinnae to 45 pairs, 4–12 ⫻ 1.2–2.2 cm, deeply pinnatifid to within 1 mm of costae; aerophores absent or essentially so; segments patent or slightly oblique, ca. 2–3.5 mm wide, rounded or subacute at apices; veins 7–10 pairs per segment; indument abaxially of patent hairs 0.2–0.3 mm long on rachises, costae, scales absent, tissue between veins with numerous patent hamate hairs ca. 0.2 mm long, adaxially the blades with adpressed hairs ca. 0.1 mm long between veins; sori supramedial to submarginal, with indusia thick, dark brown to blackish, lustrous, margins ciliate or hamate-hairy; sporangia glabrous; 2n⫽116 (Chis).

Distribution. Roadbanks and wet pine-oak forests; 800–2500 m. Mexico; Guat, Hond, Salv, Nic. Selected Specimens Examined. Chis (Breedlove 21842, 28113, 31310, 34654, DS). Oax (Hallberg 1590, NY, UC; Mickel 4867, NY, 5110, MEXU, NY). Ver (Purpus 4340, UC; Ventura A. 8141, MEXU, 19431, CHAPA, ENCB).

Thelypteris melanochlaena is closely related to T. scalaris and differs from that species primarily by the dark brown to black, thick, opaque indusia. We have not observed buds in the pinna axils of T. melanochlaena, whereas such buds are commonly found in Mexican specimens of T. scalaris.

11. THELYPTERIS MORTONII Thelypteris mortonii A. R. Sm., Amer. Fern J. 63: 125. 1973. Type. Mexico. Me´xico: Bosencheve, Sa´nchez 38 (US!; isotypes UC!, US!–3 sheets). Figs. 299A–C. Rhizomes short-creeping, to 8 mm diam.; fronds monomorphic, 60–95 cm long; stipes stramineous, darkened at bases, 3– 10 cm ⫻ 2–4 mm, with a few pale brown, adpressed, glabrous scales at bases; blades chartaceous, to 85 ⫻ 9–13 cm, with to 50 pinna pairs per blade, proximally with 10–20 pairs of gradually reduced pinnae, lacking proliferous buds; pinnae to 7 ⫻ 1.6 cm, incised 0.8–0.9 the way to the costae, opposite to subopposite; aerophores absent; segments to 25 pairs per pinna, to 8 ⫻ 2–3 mm, oblong, not falcate, slightly oblique, margins not revolute, tips rounded to subacute, separated by very narrow sinuses or contiguous, basal segments slightly longer and wider than more distal segments on a given pinna and often overlapping the rachises; veins mostly 6–9 pairs per segment; indument abaxially of scattered hairs ca. 0.2 mm long on costae, costules, and veins, scales absent, veins and tissue between veins bearing orangish sessile glands; sori supramedial to submarginal, with small epilose indusia that are nearly obscured by sessile, orange-red glands; sporangia glabrous.

THELYPTERIS OAXACANA

Distribution. Terrestrial in wet woods; 1500–2800 m. Mexico. Specimens Examined. Me´x (Sa´nchez 65, US). Mor (Lyonnet 521100005, MEXU; Lyonnet & Cha´vez 3390, MEXU, US).

This is most similar and closely related to T. resinifera, from which it differs in the short-creeping rhizomes and the small indusia that are nearly obscured by sessile, orange-red glands; it also occurs at somewhat higher elevations.

12. THELYPTERIS NUBIGENA Thelypteris nubigena A. R. Sm., Proc. Calif. Acad. Sci., ser. 4, 40: 229, f. 7C. 1975. Type. Mexico. Chiapas: Mpio. San Cristo´bal Las Casas, Zontehuitz, Breedlove 22057 (DS!; isotypes CAS!, ENCB!). Figs. 301A, B. Rhizomes suberect; fronds to ca. 65 cm long; stipes to 15 cm ⫻ 2–3 mm, darkened and sparsely scaly proximally, scales ovate, tan, sparsely hairy on margins, less than 2 mm long; blades to ca. 50 ⫻ 24 cm, 4–5 proximal pairs of pinnae gradually reduced, the lowermost to ca. 1 mm long, lacking proliferous buds; pinnae ca. 20–25 pairs, the largest 7–12 ⫻ 1.5–3 cm, broadest at bases, lobed ca. 0.9 the way towards costae; aerophores to 1 mm long at bases of proximal pinnae; segments slightly oblique, subfalcate, to 15 ⫻ 4 mm, rounded to subacute at apices; veins to 12 pairs per segment; indument abaxially of scattered hairs ca. 0.1–0.2 mm long on rachises, costae, and costules, the hairs often in pairs or threes (subfasciculate), also with numerous orangish to reddish, sessile, hemispherical glands on tissue between veins; sori medial, with minute indusia obscured by orangish glands; sporangia glabrous; 2n⫽ca. 116 (Chis).

631

Distribution. Along stream banks in pine-oak woods; 2600– 2800 m. Mexico; Guat. Specimens Examined. Chis (Breedlove 31739, DS, 53593, CAS, ENCB; Little & Sharp 9843, US). Oax (Mickel 5533, ENCB, MEXU, NY).

Thelypteris nubigena differs from other Mexican species in its tendency for the hairs on the costae and rachises abaxially to be fasciculate, but not to the same degree as in T. thomsonii. From that species, T. nubigena also differs in the absence of costular aerophores and generally smaller fronds.

13. THELYPTERIS OAXACANA Thelypteris oaxacana A. R. Sm., Amer. Fern J. 63: 125, f. 7, 8. 1973. Type. Mexico. Oaxaca: Ixtla´n–Tuxtepec road, Km 126–129, Smith 459 (UC!). Fig. 301H. Rhizomes erect; fronds ca. 60–95 cm long, mucilaginous when young; stipes stramineous to tan, 8–20 cm ⫻ 3–5 mm; blades 50– 100 ⫻ 14–25 cm, with 6–10 pairs of gradually reduced proximal pinnae, lacking proliferous buds; rachises adaxially with scattered hairs along the sulcus, glabrous abaxially; pinnae 7–13 ⫻ 1.3–2.5(– 3) cm, broadest at their bases, incised ca. 0.8 to the costae; aerophores prominent, tuberculate or peg-like at pinna bases; segments 3–5 mm wide, slightly oblique, not falcate, apices rounded or subacute; veins 5–12 pairs per segment, glabrous; indument abaxially generally absent, but with a few stramineous linearlanceolate appressed scales towards bases of costae, adaxially with scattered acicular hairs 0.5 mm long along costae, veins and tissue between veins glabrous on both sides of blades; sori medial, round or slightly oblong, with prominent, persistent, glabrous indusia ca. 0.7 mm diam.; sporangia glabrous.

632

THELYPTERIS

veins, adaxially with stout, stiff hairs 0.5–1 mm long on costae, costules, and veins, both sides eglandulose; sori supramedial to submarginal, with very small tan indusia concealed by mature sporangia, margins short-ciliate; sporangia glabrous; 2n⫽58 (Ver, Chis, Trin).

Distribution. Along streams in wet montane forests; 1500– 2250 m. Mexico; Salv, CR, Pan. Go´mez (1976) also reported this species from Nicaragua, but no specimens have been seen. Selected Specimens Examined. Chis (Breedlove 32326, 22845, DS, 34402, DS, ENCB, 42099, DS). Gro (Lorea 2058, FCME; Millan Espin 124, FCME). Oax (Germa´n R. & Sousa 47, MEXU; Smith 403, UC).

Thelypteris oaxacana differs from Mexican congeners in the costae and tissue between veins lacking hairs and glands abaxially and in having large, persistent indusia. It is very closely related to T. pachyrhachis (Kunze ex Mett.) Ching, a variable species of southern Central America, the Antilles, and South America.

14. THELYPTERIS OLIGOCARPA Thelypteris oligocarpa (Humb. & Bonpl. ex Willd.) Ching, Bull. Fan Mem. Inst. Biol. 10: 253. 1941. Figs. 300G, H. Polypodium oligocarpum Humb. & Bonpl. ex Willd., Sp. Pl., ed. 4, 5(1): 201. 1810. Dryopteris oligocarpa (Humb. & Bonpl. ex Willd.) Kuntze, Revis. Gen. Pl. 3: 378. 1898. Amauropelta oligocarpa (Humb. & Bonpl. ex Willd.) Pic. Serm., Webbia 31: 251. 1977. Type. Venezuela. Caripe, Humboldt & Bonpland s.n. (B-Willd. 19699, part!; isotype P-Bonpl.!). Dryopteris columbiana C. Chr., Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 7, 4: 279, f. 8. 1907. Thelypteris columbiana (C. Chr.) C. V. Morton, Leafl. W. Bot. 8: 194. 1957. Type. Colombia. [Magdalena:] Santa Marta, Smith 998 (P, not found; isotype UC!). Phegopteris pilosula Fe´e, Me´m. Foug. 8: 90. 1857. Type. Mexico. Linden 126 (RB?; see Windisch, 1982. Placed here on the authority of Christensen, 1920: 125.). Complete synonymy given in Smith (1983).

Rhizomes erect; fronds mostly (30–)40–75 cm long; stipes often brownish, 4–13 cm ⫻ (1–)2–3 mm, puberulous; blades chartaceous, mostly 40–65 ⫻ 10–20 cm, with 3–6 pairs of reduced proximal pinnae, lowest of these a few mm long, 2–6 cm apart, lacking proliferous buds; pinnae ca. 15–25 pairs, largest 5–8(–10) ⫻ (1–)1.2–2(–2.6) cm, deeply incised to within 1 mm of costae; aerophores absent; segments oblique, subfalcate, 2–3 mm wide, obtuse to acutish; veins 8–10 pairs per segment; indument abaxially of numerous acicular hairs, dense short hairs ca. 0.2 mm long on rachises, sometimes with a few longer hairs intermixed, hairs mostly 0.2–0.4 mm long on costae, veins, and tissue between veins, also with a few hamate hairs on tissue between

Distribution. In wet woods, especially along streams and trails, sometimes in coffee understory; 600–2200 m. Mexico; Guat, Hond, Nic, CR, Pan; Gr Ant; Col, Ven, Ec, Peru, Braz, Bol, nw Arg. Selected Specimens Examined. Chis (Breedlove 32633, DS). Col (Va´zquez 625, MEXU, UC). Gro (Tenorio L. 451, ENCB, MEXU). Gto (Dı´az B. & Martı´nez 3689, ENCB, IEB). Hgo (Yatskievych 83-357, ARIZ, UC). Jal (Mexia 1518, CAS, UC). Mich (Rees 461, MEXU). Nay (McVaugh 18956, IEB, MICH). Oax (Mickel 7009, NY, UC). Pue (Rzedowski 29995, ENCB). Qro (Rzedowski 42840, IEB, XAL). Rev (Flores P. 718, UC). SLP (Rzedowski 10057, ISC, MICH). Tam (Harrell 408, US). Ver (Copeland herb. 22, UC). Unverified, Doubtful, or Mistaken Reports. Me´x (Aranda 13 and Mojica P. 10, both IZTA, cited by Tejero-Dı´ez & Arreguı´n-Sa´nchez, 2004, but these specimens are T. rudis).

Thelypteris oligocarpa differs from T. pilosula by the combination of hamate hairs on the tissue between veins abaxially, supramedial to submarginal sori with small indusia (smaller than in T. pilosula), fewer pairs of reduced proximal pinnae, and densely and uniformly short-hairy rachises abaxially. Most of the ca. 75 collections seen are from Veracruz, Oaxaca, and Chiapas.

15. THELYPTERIS PILOSOHISPIDA Thelypteris pilosohispida (Hook.) Alston, J. Wash. Acad. Sci. 48: 233. 1958. Fig. 300L. Nephrodium pilosohispidum Hook., Sp. Fil. 4: 105. 1862. Dryopteris pilosohispida (Hook.) C. Chr., Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 7, 10: 148. 1913. Type. Ecuador. Spruce s.n. (K!). Alsophila pilosa M. Martens & Galeotti, Me´m. Foug. Mexique 78. 1842, non Thelypteris pilosa (M. Martens & Galeotti) Crawford. Type. Mexico. [Veracruz:] Totutla, Galeotti 6405 (BR!; isotypes BR!, K!).

Similar to T. rudis except: fronds generally larger; segments more or less perpendicular to costae, not falcate, the margins only slightly or not at all revolute; indument abaxially of longer hairs to ca. 1.5 mm long on the costae, hairs spreading rather than antrorse, costal scales longer, castaneous, not so clathrate;

THELYPTERIS RESINIFERA

sori with receptacles having with a few stiff acicular hairs; 2n⫽58 (Oax).

633

epaleate abaxially, tissue between veins glabrous to pubescent on both sides, often with short-stipitate glands abaxially; sori medial to supramedial, with persistent, prominent, marginally setose indusia ca. 0.4 mm diam., these a few short-stipitate glands ca. 0.1 mm long; sporangia glabrous; 2n⫽58, 116 (Oax, Chis).

Distribution. Along trails and embankments in wet forests; 1200–2200 m. Mexico; Guat, Hond, Nic; Hisp; Col, Ven, Ec, Peru, Bol. Selected Specimens Examined. Chis (Breedlove 32712, DS). Oax (Geith 95-M101, NY; Pe´rez C. C26, MEXU; Rzedowski 30600, CAS, 32815, CHAPA, ENCB; Smith 455 (UC). Pue (Rzedowski 31720, ENCB). Ver (Dorantes et al. 5117, NY; Fink 55, US; Linden 75, K; Lira 18, MEXU, UAMIZ, XAL).

Thelypteris pilosohispida is most closely related to T. rudis and is distinguished by the characters mentioned above. From other Mexican species it differs in the presence of costal scales, exindusiate sori, and lack of glands or hamate hairs on the tissue between veins abaxially. The species is probably conspecific with the Andean T. retrorsa (Sodiro) A. R. Sm., as treated by Smith (in R. Tryon & Stolze, 1992), but the inter-relationships need more study; in any case, T. pilosohispida is the older name.

16. THELYPTERIS PILOSULA Thelypteris pilosula (Klotzsch & H. Karst. ex Mett.) R. M. Tryon, Rhodora 69: 7. 1967. Fig. 301J. Aspidium pilosulum Mett., Fil. Hort. Bot. Lips. 130. 1856. Dryopteris pilosula (Klotzsch & H. Karst. ex Mett.) Hieron., Hedwigia 46: 332. 1907. Amauropelta pilosula (Mett.) A˚. Lo¨ve & D. Lo¨ve, Taxon 26: 325. 1977. Type. Cultivated plant, said by Mettenius to have originated from Caracas; a probably authentic specimen is at B!, probable isotype at BR. Complete synonymy given in Smith (1983).

Rhizomes suberect or erect; fronds 50–100 cm long; stipes tan to brownish, to 25 cm ⫻ 5 mm, with stiff, straight hairs mostly 1 mm long or more, also with persistent spreading linearlanceolate scales at bases; blades chartaceous (infrequently subcoriaceous), to 90 ⫻ 10–35 cm, with ca. 5–7(–9) pairs of gradually reduced proximal pinnae; pinnae mostly 5–12(–18) ⫻ 1.2– 2.2 cm, incised ca. 0.8–0.9 to the costae; aerophores absent; segments 3–4 mm wide, slightly oblique, not (or slightly) falcate, apices rounded to subacute; veins 6–10 pairs per segment; indument on both surfaces of stout patent acicular hairs mostly 0.3–1 mm long on rachises, costae, costules, and veins, costae

Distribution. In wet oak and pine-oak woods, especially near streams and along trails; (1200–)1600–2800 m. Mexico; Guat, Hond, Salv, CR, Pan; Jam, Hisp; Col, Ven, Ec, Peru, Bol. Go´mez (1976) reported this species from Nicaragua, but no specimens have been seen. Selected Specimens Examined. Chis (Croat 40727, UC). Gro (Martı´nez S. 3286, ENCB, MEXU). Hgo (Lyonnet 2008, US). Jal (McVaugh & Koelz 1229, IEB, MEXU, MICH). Me´x (Tejero Dı´ez 2950, IZTA). Mich (Hinton 13444, ARIZ, GH, K, LL, NY, US). Mor (Espejo 4993, UAMIZ). Oax (Yatskievych 83-149, ARIZ, UC). Pue (Ventura A. 22133, ENCB). Qro (Dı´az B. & Zamudio 3888, ENCB, IEB). Tlax (Vela´zquez s.n., MEXU). Ver (Ventura A. 256, ENCB, NY). Unverified, Doubtful, or Mistaken Reports. Gto (Dı´az B. 3689, IEB, cited as this species by Dı´az-Barriga and Palacios-Rios, 1992, but the specimen is T. oligocarpa).

Thelypteris pilosula differs from other Mexican species by the combination of indusiate sori, axes on both blade surfaces with stout acicular hairs to 1 mm long, and epaleate costae. It lacks hamate hairs and sessile glands, but often has some shortstipitate glands on the blades abaxially and also on the indusia. Possible hybrids involving this species are Mickel 4728 (ENCB, NY), from Oaxaca, and Breedlove 32703 (DS), from Chiapas. The latter differs from typical T. pilosula primarily in having only fragmentary indusia.

17. THELYPTERIS RESINIFERA Thelypteris resinifera (Desv.) Proctor, Bull. Inst. Jamaica, Sci. Ser. 5: 63. 1953. Polypodium resiniferum Desv., Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck. Gesammten Naturk. 5: 317. 1811. Dryopteris resinifera (Desv.) Weath., Contr. Gray Herb. 114: 32. 1936. Amauropelta resinifera (Desv.) Pic. Serm., Webbia 31: 251. 1977. Type. “Habitat in America calidiore.” collector not stated (P!).

634

THELYPTERIS

Rhizomes erect; fronds (15–)25–100(–130) cm long; stipes stramineous to tan, usually mucilaginous when young, 2–10(–25) cm ⫻ 1–3(–6) mm diam., mostly less than 10(–25) cm long; blades to 110 ⫻ 4–25(–40) cm, with to 12 pairs of gradually reduced proximal pinnae, lowermost pinnae hastate to auriculate; pinnae to 12(–20) ⫻ (0.4–)0.7–1.5(–2) cm (excluding basal auricles), ascending in the distal third of blades, incised to within 1 mm of costae; aerophores short, peg-like at pinna bases, or seemingly absent; segments usually very oblique to the costae, often falcate, deltate to lanceolate, to ca. 2.5 mm wide; veins (3–) 5–10 pairs per segment; indument abaxially of usually short hairs mostly less than 0.3(–0.5) mm (occasionally glabrous) on rachises, costae, and costules, tissue between veins often shorthairy and usually with numerous golden to reddish, sessile, hemispherical, resinous glands; sori medial to supramedial, with persistent tan usually glandular indusia; sporangia glabrous; 2n⫽58 (Ver, var. resinifera), 116 (Jam). Key to the Mexican Varieties of Thelypteris resinifera 1. Segments slightly oblique, Ⳳ straight; blades narrowly elliptic; western Mexico (Chihuahua to Jalisco). .... 17b. var. proxima. 1. Segments strongly oblique and falcate; blades broadly elliptic; e. and s. Mexico, w. Mexico n. to Jalisco. ....................................................................................... 17a. var. resinifera.

17a. Thelypteris resinifera var. resinifera

Figs. 301D–G.

Nephrodium panamense C. Presl, Reliq. Haenk. 1(1): 35. 1825. Dryopteris panamensis (C. Presl) C. Chr., Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 7, 4: 292. 1907. Thelypteris panamensis (C. Presl) E. P. St. John, Amer. Fern J. 26: 44. 1936. Type. Panama. Haenke s.n. (PR?, probable isotype K!). Polypodium litigiosum Liebm., Mexic. Bregn. 205 (reprint 53). 1849. Type. Mexico. Veracruz: “ombring Mirador, Liebmann s.n. (C!; isotype K!). ?Aspidium producens Fe´e, Me´m Foug. 10: 38. 1865. Type. Mexico. Veracruz: Totutla, Schaffner 220 p.p. (P?). Placed here on the authority of Christensen (1920: 124). Aspidium exsudans E. Fourn., Mexic. Pl. 1: 93. 1872. Syntypes. Mexico. Veracruz: Bottieri 66 (P, photo US!), Mu¨ller 3, 726, 1642 (P?), Liebmann s.n. (P?); Venezuela. Fendler 176 (P?); Ecuador, collector not given (P?). Aspidium exsudans E. Fourn. var. majus E. Fourn., Mexic. Pl. 1: 93. 1872. Syntypes. Mexico. Veracruz: Zacuapan, Galeotti 6306 (P?; isosyntypes BR! K!); Oaxaca: Franco s.n. (P?). Aspidium exsudans E. Fourn. var. feei E. Fourn., Mexic. Pl. 1: 93. 1872. Syntypes. Mexico. Veracruz: Orizaba, Schaffner 498, 500 (P?); Orizaba, Bourgeau 2361 (P?; isosyntypes GH! K! NY! UC! US!), 2362 (P?); “in valle Cordobensi, Bourgeau 2226 (P?; isosyntype UC!). State unknown: Linden 126 [1526?] (P?). Aspidium ghiesbreghtii E. Fourn., Mexic. Pl. 1: 94. 1872. Syntypes. Mexico. Veracruz: Orizaba, Bourgeau 3615 (P?). State unknown: Ghiesbreght 407 (P, photo US!); Schiede s.n. (P). Dryopteris panamensis (C. Presl) C. Chr. var. gonzalezii Christ ex C. Chr., Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 7, 4: 294. 1907. Type. Mexico. Oaxaca: Cerro de San Felipe, Oct 1900, Gonza´lez s.n. (P). Complete synonymy given in Smith (1981, 1983).

Differing from var. proxima in the more broadly elliptic, generally larger blades and the more strongly oblique and falcate segments.

Distribution. In woods and clearings, especially along trails and near streams, also in seepage areas and roadside ditches; 100–1600(–2300) m. USA (Fla); Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Gr Ant; Col, Ec. Selected Specimens Examined. Chis (Breedlove 31263, DS). Col (Lott 2988 et al., UC). Gro (Tenorio L. 460, CAS, ENCB, MEXU). Jal (McVaugh 20461, IEB, MEXU). Me´x (Hinton 7323, GH, K, LL, MEXU, NY, US). Mich (Torres C. 363, ARIZ, ENCB, MEXU). Mor (Salas M. 33, IEB). Oax (Mexia 9203, ARIZ, BH, CAS, GH, LL, MICH, NY, U, UC, US). Pue (Sa´nchez 641, MEXU). Qro (Rzedowski 42839, IEB, XAL). Tab (Linden 1526, K). Ver (Mun˜oz 16130, UC). Unverified, Doubtful, or Mistaken Reports. Ags (De la Cerda & Garcı´a 762, HUAA, cited by Siqueiros-Delgado & Gonza´lez-Adame, 2004, but not verified).

This species is closely related to T. struthiopteroides and T. balbisii (which see for further discussion), and a few specimens are difficult to place. Thelypteris resinifera can be distinguished by the combination of numerous, rather closely placed, gradually reduced proximal pinnae, sessile glands on the tissue between the veins abaxially, inequilateral pinnae with oblique, often falcate segments, and persistent indusia. There is considerable variation in pubescence: some specimens are nearly glabrous abaxially (e.g., Medina A. & Va´zquez B. 371, XAL, from Veracruz), but most specimens have rather dense hairs on costae, veins, and tissue between veins. Indusia vary from glabrous to hairy, but most have at least a few hairs. 17b. Thelypteris resinifera var. proxima (C. Chr.) A. R. Sm., comb. nov. Fig. 302G. Dryopteris panamensis (C. Presl) C. Chr. var. proxima C. Chr., Smithsonian Misc. Coll. 52: 377. 1909. Type. Mexico. Morelos: Cuernavaca, Ross 326 (M; isotype BM, photo UC!).

Differing from var. resinifera by the characters mentioned in the key.

THELYPTERIS SANCTA

635

brous, the hairs mostly 0.2–0.3 mm long, those abaxially sometimes hamate; sori supramedial, exindusiate; sporangia glabrous; 2n⫽58 (Oax, Jam).

Distribution. Terrestrial in moist pine-oak forests, especially along streams and seepage areas, or in streambeds; 1000–1900 m. Mexico. Selected Specimens Examined. Chih (Knobloch 579, US). Jal (Rose & Painter 7320, US; Sada 16163, UC). Mor (Ross 326, BM, M). Nay (Alava & Cook 1623, UC; Tenorio L. 16793, MEXU). Sin (Breedlove 35694, CAS; Gonza´lez O. 5070, MEXU, US).

Distribution. Along roadsides, trails in ravines, and near streams, especially in or adjacent to wet oak and pine-oak woods; (900–)1400–3000 m. Mexico; Guat, Hond, Salv, Nic, CR, Pan; Gr Ant; Col, Ven, Guy, Ec, Peru, Bol.

Mexia 608 (CAS, F, GH, UC), from Nayarit, and Salazar 287 (US), from Sinaloa, are atypical in the dense septate hairs on the rachises and costae abaxially. They do not differ from other specimens of this variety in frond shape, size, and dissection. Additional study is needed to determine whether the type of var. proxima, from Morelos, is really convarietal with the other specimens of this distinctive variety.

Selected Specimens Examined. BCS (Carter 2323 et al., CAS, K, UC, US). Chih (LeSueur 1137, ARIZ, GH, TEX, US). Chis (Breedlove 68735, CAS). Dgo (Yatskievych 81-442, UC). Gro (Martı´nez S. 964, IEB, TEX). Hgo (Knobloch 684, GH, MEXU, MSC, US). Jal (Mexia 1762, UC). Me´x (Matuda 30552 et al., US). Mich (Hinton 15673, ARIZ, GH, LL, NY, TEX, US). Mor (Pringle 13773, ARIZ, CAS, CU, GH, MICH, MSC, US). Nay (Rose 2213, GH, NY, US). Oax (Yatskievych 83-401, ARIZ, UC). Pue (Pringle 8920, CAS, ENCB, MEXU, UC, US). Sin (Vega A. 3130, ENCB, MEXU). Ver (Ventura A. 190, ENCB, MO, NY).

18. THELYPTERIS RUDIS Thelypteris rudis (Kunze) Proctor, Bull. Inst. Jamaica, Sci. Ser. 5: 64. 1953. Fig. 300M. Polypodium rude Kunze, Linnaea 13: 133. 1839. Dryopteris rudis (Kunze) C. Chr., Index Filic. 289. 1905. Amauropelta rudis (Kunze) Pic. Serm., Webbia 31: 251. 1977. Type. Mexico. [Veracruz:] Jalapa, Schiede s.n. (LZ, destroyed). Neotype (chosen by Proctor, 1985: 317). Venezuela. Otto 612 (B). Phegopteris impressa Fe´e, Me´m. Foug. 8: 90. 1857. Type. Mexico. [Veracruz:] Huatusco, Schaffner 218 (RB; see Windisch, 1982). Complete synonymy given in Smith (1983).

Rhizomes short-creeping to suberect; fronds (35–)50–150 (–200) cm long; stipes to ca. 25(–45) cm ⫻ 2–5(–8) mm, darkened, often bearing tuberculiform wart-like pinnae at intervals of 2–3(–7) cm, pubescent (infrequently glabrescent); blades chartaceous to subcoriaceous, mostly 40–150 cm long, proximally with (3–)5–12 pairs of wart-like, easily overlooked pinnae; pinnae to (5–)10–25 ⫻ (1–)1.5–3(–4.5) cm, incised to ca. 1 mm from costae; aerophores rather small, tuberculiform, at pinna bases; segments slightly oblique and subfalcate, ca. 3–5 mm wide, apices rounded to acutish, margins slightly revolute, basal pair of segments considerably reduced in proximal pinnae; indument abaxially of antrorse acicular hairs and brownish to blackish subclathrate scales along rachises and costae, especially toward the bases, tissue between veins pubescent on both sides of blades, or gla-

Unverified, Doubtful, or Mistaken Reports. NL (reported by Aguirre-Clavera´n & Arreguı´n-Sa´nchez, 1988, but not verified and doubtful).

This is one of the most common and “weedy” species of Thelypteris in Mexico and throughout the Neotropics. Specimens show great variability in size, texture, and pubescence. The presence of subclathrate costal scales will distinguish this species from all others in Mexico except for the much less common T. pilosohispida (which see for differences). Other distinguishing characters include the exindusiate sori and antrorse costal hairs.

19. THELYPTERIS SANCTA Thelypteris sancta (L.) Ching, Bull. Fan Mem. Inst. Biol. 10: 254. 1941. Figs. 299F, G. Acrostichum cruciatum L., Sp. Pl. 2: 1072. 1753, non Thelypteris cruciata (Willd.) Tardieu [see Proctor & Lourteig, Bradea 5: 384. 1990]. Type. Plumier, Traite´ Foug. Ame´r., pl. 48B, 1705, based on a plant from Hispaniola. Acrostichum sanctum L., Syst. Nat., ed. 10, 2: 1320. 1759. Dryopteris sancta (L.) Kuntze, Revis. Gen. Pl. 2: 813. 1891. Amauropelta sancta (L.) Pic. Serm., Webbia 31: 251. 1977. Lectotype (chosen by Proctor, 1977: 277). Sloane, Voy. Jamaica 1: 91, t. 49, f. 2, 1707, based on a plant from Jamaica.

Rhizomes erect, to 1.5 cm diam.; fronds numerous, tightly clustered, mostly 15–25(–40) cm long; stipes stramineous, mostly 3–5 cm ⫻ 1 mm; blades thin-chartaceous, mostly 10–20 ⫻ 2–9 cm,

636

THELYPTERIS

oblanceolate, attenuate at apices, the proximal 3–7 pinna pairs gradually reduced, auriculate or unequally 3-foliate; pinnae 1–5 ⫻ 0.4–1.2 cm, usually inequilateral at the bases, incised to within 1 mm of costae; aerophores absent; segments strongly oblique, 1– 2 mm wide, rounded to acutish at apices, entire or crenateserrate, the proximal pair often free; veins 2–6(–10) pairs per segment; indument abaxially of relatively sparse thin short hairs ca. 0.1–0.3 mm on rachises, costae, costules, veins, and tissue between veins, or blades nearly glabrous, sometimes also with resinous-glands on tissue between veins; sori medial to supramedial, with very small, tan, ciliate, sometimes resinousglandular indusia; sporangia glabrous; 2n⫽58 (Jam).

of gradually reduced proximal pinnae, lowermost auriculiform, ca. 5 mm long, buds often present in axil(s) of distal pinnae; pinnae to 45 pairs, the largest 6–16 ⫻ 1.1–2.6 cm, incised to ca. 1 mm from costae, subopposite, horizontal; aerophores absent; segments 2–4 mm wide, mostly rounded at tips, perpendicular or slightly oblique to costae; veins 9–12 pairs per segment; indument abaxially of scattered to dense hairs 0.1–0.3 mm long on rachises, costae, and veins, tissue between veins eglandulose, with erect hamate hairs to 0.3 mm long, adaxially with scattered to dense appressed hairs 0.1 mm long between veins; sori supramedial, with small stramineous to tan indusia, margins hamatehairy; sporangia glabrous; 2n⫽58 (Oax, CR).

Distribution. Terrestrial in lowland rain forests, along streams; 200–300 m. Mexico; Guat, Bel (Holst 5968, UC), Hond; Gr & L Ant.

Distribution. In wet forests, especially along trails and streams; 600–2200 m. Mexico; Guat, Hond, Salv, Nic, CR, Pan; Col, Ven, Ec, Peru.

Specimens Examined. Chis (Breedlove 47427, CAS; Martı´nez S. 8398, ARIZ, CHAPA, MEXU, MO, UAMIZ, 11302, 13715, 14825, MEXU).

Thelypteris sancta is distinguished from congeners in Mexico by its small fronds (it is generally the smallest species of subg. Amauropelta in Mexico), inaequilateral pinnae, resinous glands between the veins abaxially, and small indusia. Of Mexican species, it is most closely related to T. resinifera, but the nearest relatives are several species in the Antilles.

20. THELYPTERIS SCALARIS Thelypteris scalaris (Christ) Alston, J. Wash. Acad. Sci. 48: 234. 1958. Figs. 301N–P. Aspidium scalare Christ, Bull. Herb. Boissier, se´r. 2, 6: 159. 1906. Dryopteris scalaris (Christ) C. Chr., Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 7, 4: 323. 1907. Amauropelta scalaris (Christ) A˚. Lo¨ve & D. Lo¨ve, Taxon 26: 325. 1977. Lectotype (chosen by Christensen, 1907: 324). Guatemala. Alta Verapaz: Cubilquitz, Tu¨rckheim s.n. (ed. Donn. Sm. 8357) (P, not found; isolectotypes GH, US!). Complete synonymy given in Smith (1983).

Rhizomes suberect to erect; fronds to mostly 50–100(–200) cm long; stipes to ca. 8 cm ⫻ 2–3 mm, with densely short-hairy scales at bases; blades chartaceous, 45–90 cm long, with 7–11 pairs

Selected Specimens Examined. Chis (Chater 160, MEXU, MO; Martı´nez S. 17614, UAMIZ). Hgo (Bartholomew 3413, CAS; Luna et al. 765, FCME). Oax (Croat 47991, UC; Rzedowski 32816, ENCB). Pue (Zelaya 19, UAMIZ). Ver (Ventura A. 15138, ENCB, MO).

Thelypteris scalaris is distinguished by the combination of hamate hairs on the indusia and tissue between veins abaxially, numerous (7–11) pairs of gradually reduced proximal pinnae, and blades adaxially with short, adpressed, acicular hairs. It is similar and closely related to T. melanochlaena, which see for differences. Eleven of sixteen collections seen from Oaxaca bear a small bud in the axil of a distal pinna, as does the sole collection from Puebla. The gemmiferous collections also have shorter and denser hairs on the costae abaxially. A few specimens seem to be exindusiate, e.g., Ra´mirez & Riba 667 (XAL), from Hidalgo. Chater 160, from Chiapas, has unusually long fronds to 200 cm and long pinnae. Three collections from the vicinity of Cerro Tres Picos in Chiapas (Breedlove 25404, 34386, 34434, DS) differ from typical forms in having more oblique pinna segments, denser hairs on the costae abaxially, longer, more spreading, and less hairy rhizome scales, broader and more abruptly reduced blades proximally with fewer pairs of reduced pinnae, and larger, more prominently green indusia. These specimens probably represent a distinct variety or perhaps a different species.

THELYPTERIS THOMSONII

21. THELYPTERIS STRUTHIOPTEROIDES Thelypteris struthiopteroides (C. Chr.) C. F. Reed, Phytologia 17: 316. 1968. Figs. 299H, J. Dryopteris struthiopteroides C. Chr., Smithsonian Misc. Collect. 52: 388. 1909. Type. Guatemala. Escuintla: Concepcio´n, Donnell Smith 2459 (US! frag. BM; isotype US!).

Differs from T. balbisii by: blades with proximal pinnae gradually (not subabruptly) reduced; segments rather oblique; indument abaxially completely absent on the rachises, costae, and tissue between veins. Also similar to T. resinifera except: pinnae broader; segments longer; indument absent on the blades abaxially.

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Rhizomes erect, 1 cm diam.; fronds to ca. 45 cm long; stipes stramineous, 20 cm ⫻ 1 mm; blades herbaceous, 25 ⫻ 12 cm, lacking proliferous buds; pinnae 6–6.5 ⫻ 1.3 cm, incised to ca. 1 mm from costae, opposite, proximal several pairs deflexed, lowermost auriculiform; aerophores not seen; segments ca. 3 mm wide, obtuse at apices, margins plane; veins 4–5 pairs per segment; indument abaxially on costae of strigose hairs, otherwise glabrous, tissue between veins glabrous and eglandular on both sides; sori supramedial, with persistent, glabrous indusia; sporangia glabrous.

Distribution. Terrestrial, habitat and elevation uncertain. Mexico. Known only from the type.

Distribution. Wet forests and along trails; 400–500(–1000) m. Mexico; Guat. Specimens Examined. Chis (Breedlove 22513, 31584, 31617, DS; Matuda 18117, DS, F, MEXU, US). Oax (Hallberg 1759, NY).

This species is closely related to and more or less intermediate between T. resinifera and T. balbisii (which see for differences). It may have had a hybrid origin or have been involved in the ancestry of these two species. Further collections and study are needed to clarify relationships among the three species. All three (and also T. mortonii and T. sancta) have persistent indusia and sessile glands on the tissue between veins abaxially, but lack costal scales and hamate hairs. By this combination of characters they can be distinguished from other Mexican species except T. cinerea, which differs by the uniformly short pubescence and small fronds, and T. cheilanthoides, which has immersed veins, thicker laminae, and submarginal sori. Thelypteris cheilanthoides usually grows at higher elevations than does T. struthiopteroides.

22. THELYPTERIS TABLANA Thelypteris tablana (Christ) A. R. Sm., Amer. Fern J. 63: 127. 1973. Figs. 299D, E. Aspidium tablanum Christ, Bull. Herb. Boissier, se´r. 2, 5: 727. 1905. Dryopteris tablana (Christ) C. Chr., Index Filic. 297. 1905. Type. Mexico. Chiapas: San Pablo, Mu¨nch 146 (P!; isotype DS!, photo and frag. US!).

The specific epithet “tablanum” was almost certainly an error for or misreading of “pablonum”. The species is closely related to T. oaxacana and may be only an extremely depauperate specimen of that species.

23. THELYPTERIS THOMSONII Thelypteris thomsonii (Jenman) Proctor, Bull. Inst. Jamaica, Sci. Ser. 5: 65. 1953. Figs. 299K, L. Polypodium thomsonii Jenman, J. Bot. 24: 272. 1886. Dryopteris thomsonii (Jenman) C. Chr., Index Filic. 298. 1905. Amauropelta thomsonii (Jenman) Pic. Serm., Webbia 31: 251. 1977. Type. Jamaica. Jenman J. P. 254 (IJ; isotypes NY, US). Dryopteris stuebelii Hieron., Hedwigia 46: 340, t. 6, f. 13. 1907. Syntypes. Colombia. Stu¨bel 439, 146 (B).

Rhizomes erect, massive; fronds large, to 250 cm long; stipes stramineous to tan, mucilaginous when young, to 50 cm ⫻ 6– 10 mm, with numerous, partly appressed, tan scales to ca. 8 ⫻ 5 mm; blades to 200 ⫻ 60 cm, proximal pinnae abruptly reduced toward bases, with numerous wart-like pinnae less than 5 mm long, blades lacking proliferous buds; pinnae to 32 ⫻ 2–4.4 cm, deeply incised to within 1 mm of costae, proximal pinnae somewhat narrowed at bases; aerophores prominent, scale-like, to 6 mm long at bases of costae abaxially, smaller aerophores to 1 mm long often at bases of costules; segments 3–5 mm wide, acute to rounded at tips; veins to 25 pairs per segment, meeting the mar-

638

THELYPTERIS

gins above the sinuses; indument abaxially of dense fasciculate hairs ca. 0.1 mm long on rachises, costae, and costules, tissue with numerous reddish, sessile, hemispherical, resinous glands; sori medial, with small indusia nearly obscured by dense reddish glands like those of blades; sporangia glabrous; 2n⫽116 (Jam).

supramedial, with small, tan indusia, these sometimes with a few marginal hairs and minute sessile glands; sporangia glabrous.

Distribution. Terrestrial and on rocks, lower montane forests; 400–600 m. Mexico. Distribution. In montane rain forests, along trails in woods; 1700–2300 m. Mexico; Guat, Salv, CR, Pan; Jam, Hisp; Col, Ec, Peru. Specimens Examined. Chis (Breedlove 23121, 31981, 31986, 32345, 32683, 33695, DS). Oax (Smith 499, UC).

Thelypteris thomsonii is readily distinguished by the combination of dense, fasciculate hairs on rachises and costae abaxially, and scale-like aerophores at the bases of costae and costules.

24. THELYPTERIS VENTURAE Thelypteris venturae A. R. Sm., sp. nov. Type. Mexico. Veracruz: Mpio. Atzalan, Pilares, Ventura A. 8141 (holotype ENCB!; isotype NY!). Figs. 302A–C. A T. melanochlaena indusiis pallide brunneolis (vs. denigratis), pilis hamatis in laminis carentibus, tantum 3–4 paribus pinnarum proximalium abrupte reductis differt; a T. scalari pilis hamatis in laminis carentibus, tantum 3–4 paribus pinnarum proximalium abrupte reductis differt; a T. concinna, qua pubescentia similiter brevi in rhachidibus et costis habet, sporangiis setulosis carentibus, paribus paucioribus pinnarum proximalium reductis, pilis erectis numerosis in laminis inter venas abaxialiter, pilis numerosis 0.1 mm longis in paginis adaxialibus laminarum differt. (Honoring Francisco Ventura A., avid plant collector in southern Mexico and especially in the state of Veracruz.)

Rhizomes not known, probably suberect; fronds 45–90 cm long; stipes castaneous, 10–15 cm ⫻ 1.5–3 mm, with short-hairy scales at bases; blades chartaceous, 35–75 cm long, with 3–5 pairs of abruptly reduced proximal pinnae, lowermost auriculiform, ca. 5–10 mm long, lacking proliferous buds; pinnae 20–26 pairs, the largest 5–12 ⫻ 1.6–2.2 cm, subopposite to alternate, incised to ca. 1 mm from costae; aerophores absent; segments 3–4 mm wide, mostly acute at tips, oblique to costae; veins 8–12 pairs per segment; indument abaxially of moderately dense, uniformly short hairs 0.05–0.1 mm long on rachises, costae, and veins, tissue between veins eglandular but with erect hairs 0.05–0.1 mm long, adaxially with dense appressed hairs to 0.1 mm long; sori

Specimen Examined. Ver (Ventura A. 8030, ENCB).

Specimens of this species were initially determined by Smith, in 1974, as T. aff. melanochlaena. Thelypteris venturae differs from that in having only 3–4 pairs of abruptly reduced proximal pinnae, indument abaxially of straight (non-hamate) hairs, and light tan (rather than blackish) indusia. From T. scalaris, it differs in having uniformly dense short, straight hairs 0.05–0.1 mm long on the axes and blades abaxially (hairs longer, many hamate, in T. scalaris). From T. concinna, which has similar uniformly short pubescence on the rachises and costae abaxially, T. venturae differs in having fewer reduced proximal pinnae, numerous erect hairs on the tissue between veins abaxially, numerous hairs 0.1 mm long on the adaxial surfaces of the blades, and absence of sporangial setulae. Unplaced Name Aspidium cheiloplotium Fe´e, Me´m. Foug. 8: 104. 1857. Type. Mexico. [Veracruz:] Orizaba, 1856, Schaffner s.n. (type not located).

THELYPTERIS SUBG. CYCLOSORUS Thelypteris subg. Cyclosorus (Link) C. V. Morton, Amer. Fern J. 53: 153. 1963. Cyclosorus Link, Hort. Berol. 2: 128. 1833. Dryopteris subg. Cyclosorus (Link) C. Chr., Index Filic. XXI. 1906. Type: Cyclosorus gongylodes (Schkuhr) Link [⬅ Aspidium gongylodes Schkuhr] ⫽ Thelypteris interrupta (Willd.) K. Iwats.

Rhizomes creeping to erect; fronds pinnate-pinnatifid, the proximal pinnae reduced or not (most American spp.); blades lacking aerophores or with only a small darkened protuberance where aerophores would be located, proliferous buds absent; lowermost veins of segments meeting margins at the sinuses or connivent just below the sinuses or meeting below the sinuses with an excurrent vein to the sinuses; indument abaxially of unbranched, unicellular, acicular hairs and sometimes capitate

THELYPTERIS SUBG. CYCLOSORUS

glands on the costae and veins; sori round, with usually large, persistent, hairy indusia; sporangia with or without hairs or glands from sporangial stalks; spores bilateral, dark, with an often broadly winged perispore; x⫽36. Subgenus Cyclosorus differs from other Mexican subgenera by the combination of the lowermost veins from adjacent segments running to the sinuses or uniting below the sinuses, absence of stellate hairs, pinnate-pinnatifid blades, round sori with relatively large, persistent indusia, and glabrous sporangia. Several species are among the most common and “weediest” Thelypteris in Mexico, and may occupy somewhat exposed sites on roadbanks, in ditches, and along paths. Hybridization appears to be common in subgenus Cyclosorus when species co-occur (Smith, 1971, 1981; Smith in Mickel & Beitel, 1988). The most common hybrids in Mexico involve T. puberula, which see for discussion. Other species, e.g., T. albicaulis and T. ovata are probably also involved. Without a thor-

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ough field study of suspected hybridizing populations, it is not usually possible to determine the exact parentage of a given hybrid, but, because of the production of malformed spores, it is often possible to detect hybrids among herbarium specimens. Chromosomal studies are also needed to provide more conclusive evidence of parentage. Holttum (1971) restricted the application of Cyclosorus to the type species and one other from Africa. Later, Holttum (1974) concluded that the American species previously ascribed to Cyclosorus by Christensen and others had closest affinities to Christella, a primarily Old World genus of about 60 species. For reasons discussed elsewhere (Smith, 1981, 1983), we prefer to recognize a more broadly circumscribed Cyclosorus, at the rank of subgenus, in the New World comprising 18 species distributed from southeastern and southwestern United States, through Mexico, to northern Argentina and the Antilles. The greatest diversity is found in southern Mexico and it is here where species delimitations are most difficult.

Key to the Mexican Species of Thelypteris subg. Cyclosorus 1. Basal veins of adjacent segments united below sinuses with an excurrent vein to the sinuses. 2. Rhizomes long-creeping, nearly glabrous; costae abaxially bearing tan scales; tissue between veins usually glabrous adaxially. ..................................................................................................................................................................................................................... 31. T. interrupta.

2. Rhizomes short-creeping to erect, scaly; costae abaxially without scales; veins and tissue between veins hairy adaxially. 3. Costae abaxially with predominantly uniformly short hairs 0.1(–0.2) mm long; stipes and rachises usually darkened; rhizomes short-creeping. ....................................................................................................................................................................... 29. T. dentata. 3. Costae abaxially with most hairs greater than 0.3 mm long with many exceeding 0.5 mm; stipes and rachises usually stramineous; rhizomes suberect. ........................................................................................................................................................ 30. T. hispidula. 1. Basal veins of adjacent segments free or connivent at sinuses. 4. Rhizomes erect, the caudices (including old stipe bases) 2–5 cm in diam., the stipes straight at the bases; basal segments of proximal and middle pinnae usually elongate parallel to rachises, prominent auricles often present at bases of proximal pinnae; costae abaxially without scales. ..................................................................................................................................................................... 35. T. patens. 4. Rhizomes long- to short-creeping, less than 1 cm in diam., the stipes curved at the bases; basal segments of proximal and middle pinnae not greatly enlarged to form auricles, in some species reduced; costae abaxially with or without scales. 5. Stipes and rachises conspicuously and persistently scaly. .......................................................................................................... 37. T. tuerckheimii. 5. Stipes and rachises not conspicuously and persistently scaly. 6. Indusia densely hairy, frequently nearly obscured by hairs 0.5–1 mm long; costae abaxially densely long-hairy with hairs mostly 0.5–1 mm long; proximal pinnae often with large auricles to 7 ⫻ 2 cm; Ver. ............................................................. 33. T. lanosa. 6. Indusia sparsely to densely hairy, with hairs mostly 0.1–0.5 mm long; proximal pinnae non-auriculate or with small auricles mostly less than 2 ⫻ 0.6 cm; throughout Mexico. 7. Costules and veins adaxially with at least a few rather stout hairs mostly greater than 0.3 mm long; blades adaxially often capitate-glandular; costae abaxially lacking scales. ...................................................................................................... 32. T. kunthii. 7. Costules and veins adaxially glabrous or with a few very slender, short hairs mostly less than 0.2 mm long; tissue between veins adaxially eglandular; costae abaxially generally with at least a few scales. 8. Terminal “pinna” mostly 5 times longer than wide; proximal pinnae often less than 1.2 cm wide; QR. ........... 26. T. augescens. 8. Terminal “pinna” absent or less than 5 times longer than wide; proximal pinnae often more than 1.2 cm wide. 9. Stipe base scales very dark brown, bristle-like; indusia densely long-hairy; costae abaxially glabrous or sparsely hairy. ............................................................................................................................................................... 27. T. blepharis. 9. Stipe base scales tan to light brown, linear-lanceolate to ovate-lanceolate; indusia hairy or glabrous; costae abaxially hairy or glabrous. 10. A few narrow scales 1–3 mm long persistent on the rachises; proximal segments of pinnae usually somewhat elongate parallel to rachises; pinnae incised more than 0.8 the way to the costae; sori submarginal; hairs on costae abaxially and indusia sparse or moderately dense, acicular. .... 34. T. ovata var. lindheimeri. 10. Scales mostly absent from rachises, if present less than 1 mm long; proximal segments of pinnae not elongate parallel to rachises; pinnae incised 0.7–0.8 the way to the costae; sori medial to submarginal; indusia and costae abaxially glabrous or with dense subflexuous hairs. 11. Indusia glabrous; costae abaxially glabrous or sparsely hairy. ................................................................ 25. T. albicaulis. 11. Indusia pubescent; costae abaxially sparsely to densely hairy.

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THELYPTERIS 12. Stipe base scales ovate-lanceolate, glabrous; rhizomes short-creeping. ........................................... 28. T. cretacea. 12. Stipe base scales linear-lanceolate, sparingly hairy; rhizomes often long-creeping. 13. Largest pinnae mostly 15–35 cm long; costae abaxially sparsely hairy. .................................. 25. T. albicaulis. 13. Largest pinnae mostly less than 20 cm long; costae abaxially usually moderately to densely hairy. ............................................................................................................................................... 36. T. puberula.

25. THELYPTERIS ALBICAULIS Thelypteris albicaulis (Fe´e) A. R. Sm., Univ. Calif. Publ. Bot. 59: 94. 1971. Fig. 304J. Aspidium albicaule Fe´e, Me´m Foug. 8: 102. 1857. Type. Mexico. [Me´xico:] Valley of Mexico, Schaffner 245 (P!). Aspidium geropogon Fe´e ex E. Fourn., Mexic. Pl. 1: 95. 1872. Dryopteris geropogon (Fe´e ex E. Fourn.) C. Chr., Index Filic. 267. 1905. Dryopteris augescens (Link) C. Chr. var. geropogon (Fe´e ex E. Fourn.) C. Chr., Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 7, 10: 185. 1913. Syntypes. Six collections from southern Mexico and one from Cuba; Bourgeau 1319 (P!–3 sheets; isosyntypes C!, GH!) and Galeotti 6311 (P!) are this species.

Rhizomes long-creeping, to 8 mm diam.; fronds mostly 150– 200 cm long; stipes glabrous or glabrescent, epaleate or nearly so, 25–100 cm ⫻ 4–7 mm; blades to 100 cm long, often narrowed rather abruptly toward the somewhat elongate tips; rachises usually glabrous or sparsely hairy; pinnae (10–)15–35 ⫻ (0.9–)1.4–3.5 cm, lowermost the longest, never auriculate, narrowed at bases especially along basiscopic side, with 2–4 basiscopic segments often reduced, incised 0.5–0.8 the way to the costae; segments oblique, falcate, acute at tips, (2–)3–4 mm wide at bases; veins 10–20 pairs per segment, proximal 1–2 pairs of veins from adjacent segments connivent at the raised sinus membrane; indument abaxially of a few dark-castaneous short-hairy scales on costae, hairs absent or relatively sparse, 0.1–0.2 mm long on costae and costules, veins and tissue between veins glabrous, adaxially the blades glabrous except for sparse hairs 0.1 mm long along costae; sori medial to supramedial, with indusia glabrous or infrequently sparsely hairy with hairs 0.1–0.15 mm long, usually drying pinkish or reddish brown.

(Schaffner 84, P). Mich (Hinton 15584, ARIZ, LL, TEX, US). Mor (Lyonnet 2865, MEXU, US). Oax (Wendt et al. 4700, CHAPA, MEXU; Yatskievych 83-402, ARIZ, UC). Ver (Langman 3428, MEXU, US).

Thelypteris albicaulis is most closely related to T. puberula, from which it differs in the glabrous indusia and glabrous or sparsely hairy costae abaxially. From T. hispidula, T. dentata, and T. kunthii, it differs in lacking hairs and glands on the blades adaxially (costae excepted). Another close relative is T. grandis A. R. Sm., from southern Central America, South America, and the Greater Antilles. Previously, Smith (in Mickel & Beitel, 1988) applied the name T. grandis var. pallescens to very large-fronded specimens with short-hairy costae and indusia from western Mexico. We now believe such plants are better referred to T. albicaulis, and that the true T. grandis var. pallescens is confined to southern Central America, South American Andes, and the Greater Antilles. The following Mexican specimens are representative of this hairy variant of T. albicaulis: Jal (McVaugh 13775, MICH, US; Mexia 1764, CAS, UC; Rzedowski 27500, ENCB). Oax (Somers et al. 319, NY). Thelypteris albicaulis is suspected of hybridization with T. puberula (Smith, 1971). Several putative hybrids have been seen from Oaxaca: Kenoyer 1554, GH; Mickel 5078b, NY; Smith 489, MEXU, 490, UC. This last collection has been examined cytologically and found to show about 36II ⫹ 36I at meiosis. Another putative hybrid is from Michoaca´n: Arse`ne 5540, US. A putative hybrid between T. albicaulis and T. ovata is Breedlove 22172 (DS), from Chiapas. Spores of all these hybrids are malformed, and the various hybrids show morphological intermediacy between the suspected parents.

26. THELYPTERIS AUGESCENS Thelypteris augescens (Link) Munz & I. M. Johnst., Amer. Fern J. 12: 75. 1922. Figs. 305A, B. Aspidium augescens Link, Fil. Spec. 103. 1841. Dryopteris augescens (Link) C. Chr., Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 7, 10: 182. 1913. Christella augescens (Link) Pic. Serm., Webbia 31: 352. 1977. Type. Cultivated plants, presumably from Cuba, via Tumbadero, originally collected by Otto s.n. (B!).

Distribution. Wooded banks, near streams, in pine-oak forests; 1200–2200(–3050) m. Mexico. Selected Specimens Examined. Chis (Matuda 5178, LL, MEXU, US). Gro (Hinton 9921, ARIZ, LL, TEX, US; Soto N. & Martı´nez 5769, ENCB, UAMIZ). Jal (Wilbur & Wilbur 1856, MEXU, US). Me´x

Rhizomes creeping, 0.4–0.8 cm diam.; fronds (30–)65–140 cm, each with a Ⳳ distinct terminal pinna 5–17 ⫻ 3(–5) cm; stipes about as long as blades, 2–7(–9) mm diam., scaly at bases, the scales linear-lanceolate, castaneous, lustrous, setulose on the margins; blades coriaceous, generally 30–70 cm long; pinnae (4–)10–22(–28) ⫻ (0.3–)0.7–1.5 cm, incised 0.5–0.7 the way to the costae, proximal pinnae with basal segments slightly longer and narrower than more distal segments, basiscopic segment of distal pinnae adnate to rachises; segments subfalcate, the margins

THELYPTERIS CRETACEA

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revolute; veins (3–)5–8(–10) pairs per segment, the proximal pair from adjacent segments connivent at the sinuses; indument abaxially of castaneous, setulose scales mostly 0.6–1 mm long along costae; also with hairs 0.2–0.4 mm long costae, costules, and tissue between veins, adaxially the blades glabrous except along costae, eglandular on both sides; sori medial, indusia with hairs 0.2–0.4 mm long, paraphyses absent; 2n⫽144 (Fla).

Distribution. Terrestrial in forested ravines, especially on calcareous substrates; 2300–2400 m. Mexico; Guat. This is distinguished from other species of subg. Cyclosorus by the spreading, lustrous, castaneous scales at the bases of the stipes, glabrous or very sparsely hairy blades abaxially, and by the dense long hairs on the indusia. Distribution. Terrestrial or on limestone rocks in sun or partial shade; 50–100 m. USA (s Fla); Mexico; Guat; Bah, Cuba. Specimens Examined. QR (Cabrera 4536 et al., ENCB, MO, XAL, 16053, CAS; Calzada et al. 7100, XAL; Torres 529, 533, UAMIZ).

Palacios-Rios (2002c) cited a hybrid between this species and T. kunthii from Quintana Roo: Palacios-Rios et al. 3669 (XAL); we have not seen this specimen. Hybrids between these same two species are known from southern Florida (Smith, 1971).

27. THELYPTERIS BLEPHARIS Thelypteris blepharis A. R. Sm., Proc. Calif. Acad. Sci, ser. 4, 40: 227, f. 7F–H. 1975. Type. Mexico. Chiapas: Mpio. La Independencia, road from Las Margaritas to Campo Alegre, Breedlove 33605 (DS!; isotype DS!). Figs. 305C–E. Rhizomes creeping; fronds 70–110 cm long; stipes 20–40 cm ⫻ 4–6 mm, at bases with castaneous, lustrous, spreading, glabrous or minutely glandular scales to 8 ⫻ 1 mm; blades 50–70 cm long, gradually attenuate to the pinnatifid apices; pinnae to 19 ⫻ 2.1 cm, incised nearly to the costae; segments subfalcate, the basal ones of proximal pinnae only slightly or not at all enlarged; veins to 13 pairs per segment, the proximal pair from adjacent segments meeting the margins at or just above the sinuses; indument abaxially absent or costae and veins sparsely hairy, sometimes with scattered, yellowish, stipitate glands ca. 0.1 mm, adaxially with hairs to ca. 0.5 mm on rachises and costae, tissue between veins glabrous on both sides; sori medial, with indusia reddish brown, glandular, densely long-hairy, hairs to ca. 1 mm long.

28. THELYPTERIS CRETACEA Thelypteris cretacea A. R. Sm., Univ. Calif. Publ. Bot. 59: 92. 1971. Figs. 305G, H. Christella cretacea (A. R. Sm.) A˚. Lo¨ve & D. Lo¨ve, Taxon 26: 326. 1977. Type. Costa Rica. Guanacaste: Rı´o Piedras, 20 km SE of Liberia, Mickel 2867 (NY!; isotypes ISC!, UC!).

Rhizomes short-creeping, at apices with tan, glabrous, ovatelanceolate scales 3–5 ⫻ 1–1.5 mm, with cells Ⳳ isodiametric or slightly elongate; fronds 20–60(–100) cm long; stipes stramineous, 5–35(–50) cm ⫻ 1.5–4(–7) mm, basal scales like those of rhizomes, otherwise glabrous; blades 13–40(–56) ⫻ 10–20(–34) cm, gradually tapering distally; rachises glabrous or glabrescent, lacking scales; pinnae 5–10(–17) ⫻ (0.6–)0.9–1.5(–2.1) cm, incised (0.5–)0.7–0.8 the way to the costae, broadest at bases; segments oblique, subfalcate, 2–3(–4) mm wide; veins (4–)6–9(–12) pairs per segment, proximal pair from adjacent segments meeting margins at the sinuses or nearly so; indument abaxially absent on costae, veins, and tissue between veins, or costae with sparse to moderate hairs to 0.5 mm long, sometimes also with light yellowish, stipitate glands; sori supramedial, with indusia glabrous to moderately hairy, sometimes also with stipitate glands; 2n⫽72 (CR).

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dense, uniformly short hairs 0.1–0.2 mm long on costae, costules, and tissue between veins, occasionally with scattered longer hairs, adaxially with hairs longer and stouter except on tissue between veins, glands usually absent on blades; sori medial, with indusia uniformly pubescent, hairs 0.1–0.2 mm long; 2n⫽144 (Fla, Jam, Hisp, Trin; OW).

Distribution. Along river banks; 300–1300 m. Mexico; Guat, Salv, Nic, CR. Specimens Examined. Chis (Breedlove 35367, DS; Roe et al. 983, MICH, US). Oax (Mickel 6298, NY). Ver (Smith 2123, UC).

Thelypteris cretacea is closely related to T. patens and differs from that species chiefly by the decumbent rhizomes. In general, it is a smaller and more glabrous plant than T. patens and lacks the orangish glands along the costules that often characterize T. patens. Breedlove 35367 is atypical in having prominent auricles at the bases of the proximal pinnae and scattered setae on the stipe base scales. Roe et al. 983 lacks rhizomes and so its identity is somewhat uncertain; it could be merely a depauperate and more glabrous form of T. patens. Citation of this species by LoreaHerna´ndez & Vela´zquez M. (1998) from Guerrero, is based on a misidentification of T. patens.

29. THELYPTERIS DENTATA Thelypteris dentata (Forssk.) E. P. St. John, Amer. Fern J. 26: 44. 1936. Fig. 304K. Polypodium dentatum Forssk., Fl. Aegypt.-Arab. 185. 1775. Dryopteris dentata (Forssk.) C. Chr., Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 8, 6: 24. 1920. Cyclosorus dentatus (Forssk.) Ching, Bull. Fan Mem. Inst. Biol. 8: 206. 1938. Christella dentata (Forssk.) Brownsey & Jermy, Brit. Fern Gaz. 10: 338. 1973. Type. Yemen. Forsska˚l s.n. (C!). Aspidium molle Sw., J. Bot. (Schrader) 1800(2): 34. 1801. Dryopteris mollis (Sw.) Hieron., Hedwigia 46: 348. 1907. Type. Cultivated specimen (W!). More complete synonymy given in Smith (1971, 1983).

Rhizomes short-creeping; fronds somewhat dimorphic, the sterile shorter and with fewer, wider pinnae than the fertile; stipes darkened, often purplish, 15–45 cm ⫻ 3–5 mm, paleate at bases; blades herbaceous or chartaceous, (24–)40–92 cm long, tapering gradually toward the pinnatifid apices; pinnae 7–17 ⫻ 1.1–2.7 cm, incised 0.5–0.8 the way to costae, proximal (1–)2–6 pairs reduced, auriculate at the acroscopic bases, auricles entire or crenate; segments mostly 3–4(–5) mm wide, rounded at tips; veins 6–10 pairs per segment, the proximal pair from adjacent segments united at an obtuse angle below sinuses with an excurrent veinlet 2–4 mm long to sinuses; indument abaxially of

Distribution. Along trails, roadsides, and banks, and in cutover woods, in shade or partial sun; 50–1700 m. Se USA; Mexico; Guat, Bel (Rivero et al. 2539, UC), Hond, Salv, Nic, CR, Pan; Bah, Gr & L Ant; Col, Ven, Trin, Ec, Peru, Braz, Parag, n Arg; tropical and subtropical Africa, Asia, Pacific Islands. Selected Specimens Examined. Camp (Martı´nez S. 29489, UC). Chis (Ventura & Lo´pez 1710, UC). Col (Va´zquez 856, MEXU, UC). Gro (Lorea 4524, FCME). Mor (Trujillo E. 255, UAMIZ). Oax (Mickel 7133, NY). Pue (Oble 11, UAMIZ). Qro (Rubio 893, IEB, MEXU). QR (Torres 684, UAMIZ). Tab (Ishiki et al. 2011, CHAPA). Ver (Goconi 15142, GUADA, UC). Yuc (Garcı´a Kantun 3, UADY). Unverified, Doubtful, or Mistaken Reports. Tam (reported by Sharp, 1954, but the specimen on which this is based is no doubt T. hispidula, which see).

Thelypteris dentata differs from most other members of subg. Cyclosorus in Mexico by the veins united below the sinuses. From T. hispidula, its closest relative, it differs by the uniformly short hairs 0.1–0.2 mm long on the costae abaxially (hairs 0.3–0.8 mm long and of variable length in T. hispidula), by the short-creeping rhizomes, and by the usually darkened purplish brown stipes and rachises. This species is native to the Old World but introduced and widely naturalized in the New World. It is becoming one of the most common ferns in many areas of the Neotropics. The first collection from Mexico (Hellwig 495, NY) was in 1967, in Oaxaca; Garcı´a S. 66, ENCB, was collected in 1968, in Veracruz.

30. THELYPTERIS HISPIDULA Thelypteris hispidula (Decne.) C. F. Reed, Phytologia 17: 283. 1968. Fig. 304L.

THELYPTERIS INTERRUPTA Aspidium hispidulum Decne., Nouv. Ann. Mus. Hist. Nat. 3: 346. 1834. Christella hispidula (Decne.) Holttum, Kew Bull. 31: 312. 1976. Type. Timor. Guichenot s.n. (P). Nephrodium quadrangulare Fe´e, Me´m. Foug. 5: 308. 1852. Dryopteris quadrangularis (Fe´e) Alston, J. Bot. 75: 253. 1937. Thelypteris quadrangularis (Fe´e) Schelpe, J. S. African Bot. 30: 196. 1964. Type. French Guiana. Leprieur 182 (isotypes NY!, P!). More complete synonymy given in Smith (1971, 1983).

Rhizomes suberect to erect (short-creeping to long-creeping in some parts of range), caudices usually 1.5–3 cm diam.; stipes stramineous, darkened only at bases, 10–40(–55) cm ⫻ 1.5–5 mm, hairy; blades herbaceous to chartaceous, 18–55(–75) cm long, tapering evenly to the pinnatifid apices; pinnae 3.5–16 ⫻ 0.8–2(– 2.6) cm, incised (0.5–)0.6–0.8 to costae, 0–4 pairs of proximal pinnae gradually reduced at blade bases, sometimes with small crenately lobed auricles at superior bases; segments suboblique, mostly straight, acute or rounded; veins (4–)6–9(–11) pairs per segment, the proximal pair from adjacent segments united below sinuses with an excurrent veinlet 1–2 mm long (ours); indument abaxially of numerous, often stout hairs 0.3–0.8 mm long on costae, costules, veins, and tissue between veins, adaxially usually also hairy on costae, costules, and veins, stipitate yellow glands often present on both surfaces; sori medial, indusia usually with dense hairs less than 0.3 mm long; 2n⫽72 (Fla, La, Trin; OW).

Distribution. Along streams in moist woodlands; 20–1550 m. Se USA; Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol, Parag, n Arg, Uru; tropical Africa and Asia. Selected Specimens Examined. Chis (Matuda 18156, F, MEXU, US). Col (Lott 2989 et al., UC). Jal (Pringle 8793, BH, CAS, CU, ENCB, NY, UC, US). Mich (Hinton 16256, DS, LL, MICH, NY, UC). Nay (Jones 23492, UC). Oax (Mickel 6087, NY, US). Sin (Lamb 494, NY, US). SLP (Rzedowski 9672, ISC). Son (Gentry 1646, ARIZ, NY, UC). Tab (Matuda 3363, F, LL, MEXU, US). Tam (Sharp et al. 52156, US). Ver (Martı´nez 2182, MEXU, MO).

This species is most closely related to T. dentata, which see for differences. From other Mexican Thelypteris, T. hispidula differs in the veins being united below the sinuses, with an excurrent vein to the sinuses. Some specimens from western Mexico, e.g., Correll & Correll 28835 (LL), from Nayarit, are atypical in

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having densely yellow-glandular indusia and veins variously connivent at or united below the sinuses. Several varieties have been recognized, including var. versicolor (R. P. St. John) Lellinger, in the southeastern United States, and var. inconstans (C. Chr.) Proctor, in the Greater Antilles. Mexican and most continental tropical American specimens seem not or only weakly distinguishable from var. hispidula in the Old World.

31. THELYPTERIS INTERRUPTA Thelypteris interrupta (Willd.) K. Iwats., J. Jap. Bot. 38: 314. 1963. Fig. 305J. Pteris interrupta Willd., Phytogr. 13, t. 10, f. 1. 1794. Cyclosorus interruptus (Willd.) H. Itoˆ, Bot. Mag. Tokyo 51: 714, f. g. 1937. Type. India. Klein s.n. (BWilld. 19770, 2 sheets). Polypodium tottum Thunb., Prodr. Pl. Cap. 172. 1800. Thelypteris totta (Thunb.) Schelpe, J. S. African Bot. 29: 91. 1963. Cyclosorus tottus Thunb.) Pic. Serm., Webbia 23: 173. 1968. Type. South Africa. Thunberg Hb. 24724 (UPS!). Aspidium gongylodes Schkuhr, 24. Kl. Linn. Pfl.–Syst. 2: 193, t. 33c. 1805 [“goggilodus”]. Cyclosorus gongylodes (Schkuhr) Link, Hort. Berol. 2: 128. 1833. Thelypteris gongylodes (Schkuhr) Small, Ferns S.E. States 248. 1938. Type. British Guiana. Essequebo, received by Ga¨rtner (isotypes S, B). Nephrodium paludosum Liebm., Mexic. Bregn. 275 (reprint 133)]. 1849. Lectotype (chosen by Smith, 1971: 56). Mexico. Veracruz: Circa San Antonio Huatusco, Liebmann s.n. [Pl. Mex. 2658, Fl. Mex. 494] (C!, photos UC!, frag. US!).

Rhizomes long-creeping, black, naked, 3–6 mm diam., stipe bases mostly 2–9 cm apart; fronds 50–150(–200) cm long; stipes 17–105 cm ⫻ 3–6 mm, glabrescent; blades chartaceous to subcoriaceous, 30–80 cm long, tapering evenly toward the pinnatifid apices; pinnae 7–30 ⫻ 1–2.2 cm, incised 0.3–0.6 the way to the costae, lowermost short-stalked 1–4 mm; segments deltate, rounded to acute at tips; veins 9–18 pairs per segment, proximal pair from adjacent segments united at an obtuse angle below the sinuses with an excurrent vein usually 2–4 mm long; indument abaxially of tan, ovate ciliate costal scales mostly 0.5–1.5 ⫻ 0.2– 0.8 mm, infrequently the costae also with sparse to moderately dense hairs 0.1–0.2 mm long, tissue between veins with red or orange, sessile, lustrous glands, adaxially the blades glabrous except for sparse costal hairs; sori medial to supramedial, often appearing confluent when sporangia mature, with glabrous or hairy indusia, sporangial stalks often bearing stalked glands like those on blades; 2n⫽72 (OW), 144 (Fla, Jam).

644

THELYPTERIS

out pronounced basal auricles, or infrequently with auricles weakly developed; segments oblique, straight or subfalcate, rounded or acute at tips; veins (3–)6–11 pairs per segment, the proximal pair from adjacent segments running to or near the sinuses; indument abaxially of usually rather dense hairs mostly 0.2–0.5 mm long on costae, costules, veins, and between veins, adaxially the costae, costules, and veins also hairy, the costal hairs stouter on both sides of blades, to 0.8 mm long, tissue between veins glabrous or often with light yellowish, stipitate glands; sori medial, with indusia sparsely to densely hairy, hairs 0.2–0.4 mm long; 2n⫽144 (USA, Jam, CR)

Distribution. Marshes and swamps, wet ditches; 0–1700 m. USA (Fla); Mexico; Guat, Hond, Salv, CR, Pan; Gr & L Ant; Col, Ven, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol, Parag, n Arg, Uru; tropical and subtropical Africa, Asia. Selected Specimens Examined. Camp (Ocan˜a-Nava 279, MEXU). Chis (Breedlove 29945, 33042, DS). Hgo (Moore 2463, BH). Jal (Garcı´a 4954, IBUG). Nay (Knobloch 1644, US; Pinkava et al. P12825, ASU). QR (Castillo R. 229, XAL). Tab (Cowan 2001, CAS, CSAT, MEXU, MO, NY, UAMIZ; Matuda 3338, F, LL, MICH, NY, US). Ver (Martı´nez C. 1615, CAS, K, MEXU, MO, UC).

Thelypteris interrupta is easily distinguished from other Mexican Cyclosorus by the presence of orangish or reddish glands on the blades abaxially, ovate costal scales, and very long-creeping, blackish, nearly naked rhizomes. Cyclosorus is treated by Holttum as a small genus of about two or three species, in Mexico including only T. interrupta. This species, T. serrata, and T. palustris, are also the only Thelypteris in Mexico that grow in swampy sites. Most specimens of T. interrupta from Mexico lack hairs on the rachises and costae abaxially; exceptions are Matuda 3338, from Tabasco, and Liebmann 492, 493, 494 (C) and Martı´nez C. 1615, from Veracruz.

32. THELYPTERIS KUNTHII Thelypteris kunthii (Desv.) C. V. Morton, Contr. U.S. Natl. Herb. 38: 53. 1967. Figs. 304A, B, G. Nephrodium kunthii Desv., Me´m Soc. Linn. Paris 6: 258. 1827. Lectotype (designated by Morton, Contr. U. S. Natl. Herb. 38: 53. 1967). [Venezuela.] Cumanacoa, Caripe, Humboldt & Bonpland s.n. (P-Humb.!). Dryopteris normalis C. Chr., Dansk Bot. Ark. 9(11): 31. 1910. Thelypteris normalis (C. Chr.) Moxley, Bull. S. Calif. Acad. Sci. 19: 57. 1920. Christella normalis (C. Chr.) Holttum, Webbia 30: 193. 1976. Lectotype (chosen by Smith, 1971: 78). Jamaica. Jenman s.n. (NY!). More complete synonymy given in Smith (1971), under T. normalis.

Rhizomes short- to long-creeping; fronds mostly 50–150 cm long; stipes (5–)20–80 cm ⫻ (1–)3–6 mm, with basal scales castaneous, lustrous, lanceolate, hairy; blades herbaceous to chartaceous, (9–)30–65(–80) cm long, tapering evenly toward the pinnatifid apices; pinnae (2–)8–15(–20) ⫻ (0.6–)0.9–2.5 cm, proximal pair the largest, incised 0.6–0.9 the way to costae, with-

Distribution. In woods, often along streams and trails; 350– 1600 m. Se USA; Mexico; Guat, Bel, Hond, Salv, Nic, CR; Bah, Gr & L Ant; Col, Ven, n Braz. Selected Specimens Examined. Camp (Lundell 1224, DS, MICH, US). Chis (Breedlove 38102, DS). Gro (Lorea 2869, FCME). Hgo (Arreguı´n S. 1037, MEXU). Oax (Mickel 5095, NY, UC). Qro (Servı´n 559, IEB, UC). QR (Gaumer 1341, MEXU, NY). SLP (Whittier 23, MICH). Tam (Rutten et al. 497, U). Ver (Palmer 316a, CAS, MO, NY, US). Yuc (Steere 1571, MICH, US).

This species, which is very common in the Greater Antilles and southeastern United States, is uncommon or rare everywhere in Mexico, except locally in Quintana Roo, Chiapas, and Oaxaca. Thelypteris kunthii is distinguished from T. hispidula by having the proximal pair of veins from adjacent segments running to the sinuses. From T. patens, it differs by having creeping rhizomes. Other close relatives (T. ovata, T. puberula) differ in having costules and veins glabrous adaxially. Generally, this species grows at lower elevations than T. puberula. Orcutt 6626 (DS), said to be from Berta, Veracruz, is very unusual in having two pairs of greatly reduced proximal pinnae and resembles in blade shape T. conspersa (Schrad.) A. R. Sm., known largely from South America but with populations also in Panama and Hispaniola. We wonder if the Orcutt specimen is wrongly labelled.

33. THELYPTERIS LANOSA Thelypteris lanosa (C. Chr.) A. R. Sm., Univ. Calif. Publ. Bot. 59: 70. 1971. Figs. 305M–O.

THELYPTERIS OVATA VAR. LINDHEIMERI Dryopteris patens (Sw.) Kuntze var. lanosa C. Chr., Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 7, 10: 180. 1913. Christella lanosa (C. Chr.) A˚. Lo¨ve & D. Lo¨ve, Taxon 26: 326. 1977. Lectotype (chosen by Smith, 1971: 70). Mexico. [Veracruz:] Bourgeau s.n. (B!). Aspidium pallidum E. Fourn. var. majus E. Fourn., Mexic. Pl. 1: 96. 1872, nom. nud. Based on Mexico. [Veracruz:] Valle´e de Cordova, Bourgeau 1643 (A!, MICH!, P!, US!).

Rhizomes short-creeping; fronds mostly 100–170 cm long; stipes stramineous, darkened at bases, to 60 cm ⫻ 9 mm, scaly at bases, the scales persistent, white or faintly pinkish when young, light brown when mature, ovate-lanceolate, to 14 ⫻ 6 mm, glabrous; blades chartaceous to subcoriaceous, dark greenish gray when dried, to 110 ⫻ 70 cm, tapering evenly toward the pinnatifid apices; pinnae to 35 ⫻ 3.7 cm, sessile, incised 3⁄4–4⁄5 the way to costae, proximal pinnae slightly or not at all reduced, with basal acroscopic segments often greatly enlarged and incised, the auricles to 7 ⫻ 2 cm; segments oblique, subfalcate to falcate, the margins frequently revolute; veins 13–19 pairs per segment, the proximal pair from adjacent segments connivent at the sinuses or distal vein of a pair reaching the margin just above the sinuses; indument abaxially of dense hairs on costae, costules, veins, and usually tissue between veins, costae lacking scales and glands, or the latter inconspicuous, stipitate, obscured by hairs, adaxially the costae densely hairy, hairs mostly 0.5–1 mm; sori medial, with indusia pale brown, densely hairy, hairs 0.5–1 mm, sometimes nearly obscuring the indusia; 2n⫽72 (Ver).

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34. THELYPTERIS OVATA VAR. LINDHEIMERI Thelypteris ovata R. P. St. John var. lindheimeri (C. Chr.) A. R. Sm., Amer. Fern J. 61: 30. 1971. Figs. 304C–E. Dryopteris normalis C. Chr. var. lindheimeri C. Chr., Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 7, 10: 182. 1913. Type. U.S.A. Texas: Lindheimer 742 (B!; isotypes IJ, MO!, UC!, US!). More complete synonymy given in Smith (1971).

Rhizomes creeping; fronds (30–)55–135(–165) cm long; stipes about as long as blades, (1.5–)2–6 mm diam., the scales at bases linear-lanceolate, tan to brown, ciliate; blades (18–)30–75 ⫻ 10– 50 cm, tapering evenly toward the pinnatifid apices; rachises with scattered, persistent, narrow scales 1–3 mm long, moderately hairy to glabrescent; pinnae (5–)10–25 ⫻ 0.8–2.2 cm, incised more than 0.8 the way to costae; proximal ones the largest, often somewhat narrowed toward their bases, proximal segments of medial pinnae slightly longer than more distal segments, often parallel to rachises; segments oblique, mostly 2–4 mm wide, subfalcate, usually acute at tips; veins 6–13 pairs per segment, proximal pairs from adjacent segments reaching margins at or just above sinuses; indument abaxially of linear tan costal scales 0.5– 1.5 mm long and moderate to dense hairs 0.2–0.5 mm long on rachises and costae, veins and tissue between veins usually moderately to densely hairy, adaxially the blades glabrous except along rachises and costae; sori supramedial to submarginal, indusia with hairs mostly 0.2–0.4 mm long, paraphyses absent; 2n⫽72 (Tex).

Distribution. Terrestrial in wet montane forests; 300–1450 m. Mexico. Selected Specimens Examined. Ver (Bourgeau 2367, A, US; Finck 49, MEXU; Purpus 2934, MO, NY, UC, US, 4338, UC, 16484, US, 16606, US; Reeves R5866, ASU; Smith 405, 411, 413, UC; Ventura A. 2717, LL).

Distribution. Along trails; 50–1500(–2100) m. USA (Tex); Mexico; Guat, Bel; Jam.

This is most similar to T. patens, and, without rhizome, the two are difficult to distinguish. They differ especially by the rhizome habit (creeping in T. lanosa, erect in T. patens), stipe base scales (narrower and pubescent in T. lanosa), and the denser hairs on the indusia and abaxial surfaces of the blades. From T. kunthii, also very similar, T. lanosa differs by the denser and longer hairs on the indusia and abaxial blade surfaces, and the more auriculate proximal pinnae.

Selected Specimens Examined. Camp (Steere 1784, MICH). Chih (Wendt & Lot 99, CHAPA, MEXU, TEX). Chis (Davidse 29580 et al., UC). Coah (Villareal 3998, IEB, MEXU). Gro (Rhymes & Rowell 3874B, MICH, US). Hgo (Croat & Hannon 65980, UC). Mor (Lorence 5035, MEXU, TEX). Nay (McVaugh 18808, IEB, MICH). NL (Dorr 2591, TEX, UC). Oax (Yatskievych 85-125 et al., UC). Pue (Sa´nchez 655, MEXU, US). Qro (Rzedowski 48438, IEB, XAL). SLP (Duncan 2409, UC). Tab (Gonza´lez 17140, UC). Tam (Palmer 183, NY, UC, US). Ver (Harriman 15662, UC).

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Unverified, Doubtful, or Mistaken Reports. Me´x (Luja´n 5 and Tejero-Dı´ez 2670, 2755, IZTA, cited by Tejero-Dı´ez & Arreguı´nSa´nchez, 2004; the specimens are, however, T. puberula var. puberula).

McVaugh 18809 (MICH) from Nayarit, was cited by Mickel (1992) as T. puberula var. puberula, but this number at IEB is T. ovata var. lindheimeri. It is unusual in having large (3 ⫻ 1 cm) pinnatifid auricles at the bases of the proximal pinnae. Thelypteris ovata var. lindheimeri is most similar to T. puberula, but differs in the more elongate proximal segments of the proximal pinnae, longer and stiffer hairs on costae abaxially, lighter-colored and longer scales on the rachises and costae abaxially, and submarginal sori. Apparently, it also differs in chromosome number. Variety ovata is known from the southeastern United States and Bahamas; it is based on Thelypteris ovata R. P. St. John in Small, Ferns S.E. States 239, cum tab. 1938 [syns.: Dryopteris ovata (R. P. St. John) Broun, Ind. N. Amer. Ferns 76. 1938. Christella ovata (R. P. St. John) A˚. Lo¨ve & D. Lo¨ve, Taxon 26: 326. 1977. Type. U.S.A. Florida: Citrus Co., Lecanto, R. St. John [394] (NY!)].

35. THELYPTERIS PATENS Thelypteris patens (Sw.) Small, Ferns S.E. States 243. 1938. Fig. 304H. Polypodium patens Sw., Prodr. 133. 1788. Dryopteris patens (Sw.) Kuntze, Revis. Gen. Pl. 2: 813. 1891. Christella patens (Sw.) Holttum, Webbia 30: 193. 1976. Lectotype (chosen by Smith, 1971: 74). Jamaica. Swartz s.n. (S!). Complete synonymy given in Smith (1971).

Rhizomes erect, caudices 2–4 cm diam., apices and stipe bases covered with large, ovate, tan (whitish when young) scales to 1.5 ⫻ 0.5 cm, with cells isodiametric; fronds mostly 50–125(–250) cm long; stipes (5–)15–50(–95) cm ⫻ (1–)2.5–9(–12) mm; blades (8– )25–75(–100) cm long, tapering evenly toward apices; pinnae (3– )10–32 ⫻ (0.5–)1.2–3(–4) cm (excluding proximal segments, those sometimes elongate parallel to rachises or pinnatifid and auriculate to 5 cm long), the largest proximal or subproximal, usually deeply incised 0.7–0.9 the way to the costae; segments oblique, mostly 3–5 mm wide, subfalcate to falcate, acute; veins (6–)8–14(–21) pairs per segment, the proximal pair from adjacent segments running to sinuses; indument abaxially usually of numerous hairs mostly 0.2–0.4 mm long on costae, veins, and tissue between veins (occasionally hairs sparse or absent), also often with short-stipitate glands, scales absent on costae, blades adaxially glabrous except along costae; sori medial, with indusia tan, often stipitate-glandular, usually hairy; 2n⫽72 (Chis, Trin), 144 (CR, Jam).

Distribution. Along trails, river banks, and in shady canyons; 100–1500 m. USA (Fla); Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Peru, Braz, Bol, Parag, nw Arg. Selected Specimens Examined. Camp (Martı´nez S. 28084, XAL). Chis (Case et al. 198, NY, UC). Gro (Lorea 1853, FCME). Hgo (Gimate 947, CHAPA, NY, UAMIZ). Mich (Rufino del Llano 86, IEB, UC). Oax (Hallberg 1677, NY, UC). Qro (Dı´az B. & Carranza 6517, IEB, UC). QR (Chan 2605, CICY, XAL). Tab (Zamudio R. 203, IEB). Ver (Copeland herb. 26, MEXU, MICH, UC, US). Unverified, Doubtful, or Mistaken Reports. NL (reported by Aguirre-Clavera´n and Arreguı´n-Sa´nchez, 1988, but not verified, specimen is probably misidentified). Tres Marı´as (reported by Lenz, 1995, but not verified, most likely misidentified).

The erect rhizomes and large, ovate, glabrous scales at rhizome apices and stipe bases distinguish this species from all other Mexican members of subg. Cyclosorus. Mexican specimens are referable to var. patens; var. smithiana Ponce occurs in the Antilles, Central America, and South America, and var. dissimilis (Schrad.) A. R. Sm. is confined to southern Brazil and perhaps adjacent countries.

36. THELYPTERIS PUBERULA Thelypteris puberula (Baker) C. V. Morton, Amer. Fern J. 48: 138. 1958. Aspidium puberulum Fe´e, Me´m. Foug. 10: 40. 1865, hom. illeg., non Gaudich., 1827. Nephrodium puberulum Baker in Hooker & Baker, Syn. Fil., ed. 2, 495. 1874. Dryopteris puberula (Baker) Kuntze, Revis. Gen. Pl. 2: 813. 1891. Dryopteris feei C. Chr., Index Filic. 264. 1905, nom. superfl. Dryopteris augescens (Link) C. Chr. var. puberula (Baker) C. Chr., Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 7, 10: 184. 1913. Thelypteris feei (C. Chr.) Moxley, Bull. S. Calif. Acad. Sci. 20: 34. 1921. Christella puberula (Baker) A˚. Lo¨ve & D. Lo¨ve, Taxon 26: 326. 1977. Type. Mexico. Veracruz: Near Huatusco, Schaffner 247, p.p. (not found at P). Complete synonymy given in Smith (1971).

Rhizomes long-creeping, stipe bases (0.5–)1–3 cm apart; fronds 35–120(–160) cm long; stipes equaling blades, 15–50(–80) cm ⫻ (2–)3–7 mm, paleate at bases, the scales hairy, not persistent;

THELYPTERIS PUBERULA

blades chartaceous to subcoriaceous, 20–55(–85) cm long, tapering evenly toward the pinnatifid apices, or abruptly reduced just below the somewhat elongate apices; rachises lacking scales or infrequently with a few persistent castaneous scales less than 1 mm long, also usually moderately hairy; pinnae 7–20(–26) ⫻ 0.9–2(–3) cm, proximal pair the largest, incised 0.5–0.8 the way to the costae, proximal several pairs narrowed toward their bases except for slightly enlarged entire acroscopic basal segments, basal basiscopic segment of distal pinnae strongly adnate to rachises; segments oblique, mostly 3–5 mm wide, subfalcate, acutish; veins 7–12(–15) pairs per segment, the proximal 1–2(–3) pairs from adjacent segments connivent at sinuses; indument abaxially on costae of a few minute castaneous scales less than 0.5 mm long, the costae, veins, and tissue between veins pubescent with hairs ca. 0.2 mm long, blades eglandular or sparsely glandular, adaxially the blades glabrous except along costae; sori medial, with indusia hairy, rarely glabrous or glabrescent; 2n⫽144, var. puberula (Oax, Chis). This species hybridizes with T. albicaulis, which see. Va´zquez T. 370 (XAL), from Veracruz, has strongly malformed spores and is, apparently, a hybrid involving this species; the specimen has glabrous indusia and sparsely hairy costae abaxially. Ventura A. 7781 (MEXU), also from Veracruz, has collapsed or empty sporangia or extremely malformed spores and is probably also of hybrid origin. Breedlove 33725 (DS), from Chiapas, has malformed spores and is the putative hybrid between T. puberula and T. tuerckheimii. Mickel 6204 (NY) has spores that are greater than 50 percent malformed, but its parentage (other than T. puberula) is not obvious. Key to the Mexican Varieties of Thelypteris puberula 1. Adaxial blade surface glabrous, except for costae. .... 36a. var. puberula. 1. Adaxial blade surface bearing short, appressed hairs 0.1– 0.2 mm long. .............................................................. 36b. var. sonorensis.

36a. Thelypteris puberula var. puberula

Fig. 304F.

Differing from var. sonorensis in having the adaxial blade surfaces, except for costae, glabrous.

Distribution. Oak and pine-oak woods, open stream banks, roadbanks; (100–)500–2600 m. Mexico; Guat, Hond, Salv, Nic, CR; Hisp.

647

Selected Specimens Examined. Chis (Palacios-Rios 2919, UC). Col (Sanders 10646, UC). Dgo (Benı´tez 1156, CHAPA, IEB). Gro (Mexia 8893, MICH, NY, UC, US). Gto (Rzedowski 41560, ENCB, IEB). Hgo (Rzedowski 25425, MICH). Jal (Sahagun 17263, UC). Me´x (Matuda 27540, CAS, US). Mich (Dı´az Luna 3219, UC). Mor (Yatskievych 86-315 & Gastony, UC). Nay (McVaugh 18809, MICH). Oax (Krueger & Gillespie 21, UC, US). Pue (Yatskievych 83-376, ARIZ, UC). Qro (Ferna´ndez N. & Acosta 2086, 2147, IEB). Sin (Rose et al. 14081, NY, US). Tam (Bartlett 10419, MEXU, MICH, US). Ver (Seaton 67, NY, US). Unverified, Doubtful, or Mistaken Reports. Ags (De la Cerda & Garcı´a 15, 280, HUAA, cited by Siqueiros-Delgado & Gonza´lezAdame, 2004, but not verified). NL (reported by Aguirre-Clavera´n and Arreguı´n-Sa´nchez, 1988, but not verified and perhaps misidentified).

Thelypteris puberula var. puberula differs from other members of subg. Cyclosorus in Mexico by the combination of veins (1–3 pairs) running to the sinuses, presence of small costal scales abaxially, costae also hairy, tissue between veins glabrous adaxially (costae excepted), and pubescent indusia. A few specimens, especially from Veracruz, appear to be this species but have glabrous indusia and glabrous or sparsely hairy costae abaxially: Herna´ndez A. & Figueroa N. 47 (XAL) and Galeotti 6311 (P, BR), both from Puente Nacional, the latter a syntype of Aspidium albicaule Fe´e; Nee & Taylor 29509 (XAL); Robles H. 47 (XAL). Mostly, these glabrous forms are from low elevation, 100–500 m. A similar specimen from Chiapas is Croat 40367 (MEXU), which has very large, glabrous indusia and moderately hairy costae; curiously it does not seem to produce sporangia! Another aberrant specimen from low elevation in Chiapas appears to lack costal scales: Davidse et al. 30094 (MEXU). From T. grandis A. R. Sm., of the Antilles, Central America, and South America, T. puberula differs mainly by the shorter and narrower pinnae and more densely hairy blades abaxially. 36b. Thelypteris puberula var. sonorensis A. R. Sm., Univ. Calif. Publ. Bot. 59: 91, f. 125e. 1971. Type. U.S.A. Arizona: Pinal Co., Aravaipa Canyon, Galiuro Mts, Phillips 2877 & Reynolds (US!; isotypes MICH! MO! NY! UC! US!). Figs. 305K, L. Differing from var. puberula in having the adaxial blade surfaces between veins with short, appressed hairs 0.1–0.2 mm long.

648

THELYPTERIS

Distribution. Terrestrial in moist ravines in pine-oak forests or tropical subdeciduous forests; 300–2000 m. Sw USA (Calif, Ariz); Mexico.

Distribution. Fissured limestone slopes, sink holes, banks, margins of montane rain forests, pine-oak forests; 400–2700 m. Mexico; Guat.

Selected Specimens Examined. BCN (Charlton 1338, UC). BCS (Moran 18194, UC). Chih (Palmer 161, US). Col (Orcutt 4590, US). Dgo (Palmer 1241⁄2, NY, US). Gto (Kennoyer 2245, US). Jal (Dı´az L. 5479, UC). Mich (Hinton 16284, LL, MICH, NY, UC). Nay (Jones 23491, NY, UC). SLP (Schaffner s.n., in 1879 and 1881, NY). Sin (Rose et al. 14864, US). Son (Sanders 13084 et al., UC).

Selected Specimens Examined. Chis (Breedlove 22336, 27938, 29577, 32150, 33426, 34680, DS, 48263, 57110, CAS; Carlson 1836, US; Riba 1501, UAMIZ). Oax (Geith 95-M034, NY).

McVaugh & Koelz 1756 (MEXU), from Jalisco, 300 m (unusually low for this species and variety), is aberrant in having thin blade texture, deeply incised pinnae (nearly to costae), and absence of costal scales abaxially. Assignment to var. sonorensis is somewhat uncertain.

37. THELYPTERIS TUERCKHEIMII Thelypteris tuerckheimii (Donn. Sm.) C. F. Reed, Phytologia 17: 321. 1968. Fig. 305F. Nephrodium tuerckheimii Donn. Sm., Bot. Gaz. 12: 133. 1887. Dryopteris tuerckheimii (Donn. Sm.) C. Chr., Index Filic. 299. 1905. Lectotype (chosen by Smith, 1971: 69). Guatemala. Alta Verapaz: Near Coba´n, Tu¨rckheim s.n. [Donn. Sm. 704] (US!; isotypes B!, GH!, K!, NY!, P!, UC!, US).

Rhizomes creeping, stout; fronds 60–175 cm long; stipes 30–85 cm ⫻ 4–10 mm, bearing dense brown or light brown, ovatelanceolate, glabrous scales to ca. 10 mm long; blades coriaceous to subcoriaceous, (30–)45–90 ⫻ (16–)25–55 cm, tapering evenly toward the short-pinnatifid apices; pinnae (8–)12–28 ⫻ 1.3–3 cm, proximal pair the largest, incised ca. 0.7–0.8 the way to the costae, often strongly auriculate at bases; segments subfalcate, mostly 3–5 mm wide, the margins often strongly revolute; veins 10–18 pairs per segment, proximal pair from adjacent segments connivent at sinuses; indument abaxially of numerous ovate to lanceolate scales on rachises, costae, and costules, also rather densely hairy on rachises, costae, costules, veins, and tissue between veins, hairs 0.5–1 mm long; sori medial, with hairy indusia, hairs mostly 0.3–0.5 mm; 2n⫽72 (Chis).

Usually, this is easily distinguished from all other species of subg. Cyclosorus by its densely and persistently scaly stipes and rachises. However, some specimens from Chiapas are less scaly than the Guatemalan type and approach T. ovata var. lindheimeri. A putative hybrid between T. tuerckheimii and T. puberula is mentioned under the latter species. Unplaced Name Aspidium conspersoides Fe´e, Me´m. Foug. 8: 105. 1857. Syntypes. Mexico. [Veracruz:] Orizaba, Schaffner 335, 463; Hispaniola. de Tussac s.n. (not found).

THELYPTERIS SUBG. GONIOPTERIS Thelypteris subg. Goniopteris (C. Presl) Duek, Adansonia ser2. 11: 720. 1971. Goniopteris C. Presl, Suppl. Tent. Pterid. 181. 1836. Dryopteris Adans. subg. Goniopteris (C. Presl) C. Chr., Index Filic. XXII. 1906. Thelypteris sect. Goniopteris (C. Presl) C. V. Morton, Amer. Fern J. 53: 154. 1963. Lectotype (chosen by J. Smith, Hist. Fil. 191. 1875): Polypodium crenatum Sw. [ Goniopteris crenata C. Presl] ⫽ Thelypteris poiteana (Bory) Proctor.

Rhizomes with numerous thick prop-like roots ca. 1 mm diam., these seemingly glabrous; scales of rhizome apices and stipe bases usually with minute forked or stellate hairs ca. 0.1 mm long; fronds rarely simple, mostly pinnate or pinnatepinnatifid, gradually reduced toward the pinnatifid apices or with terminal pinnae conform; blades dark green or gray-green, sometimes pustulate (warty), aerophores absent, proliferous buds sometimes present in axils of distal (rarely proximal) pinnae; proximal pinnae the longest or nearly so, rarely reduced; veins connivent at sinuses, or with one to several pairs of veins united below sinuses, rarely veins free; indument of stipes, rachises, costae, costules, and sometime veins and tissue between veins, es-

THELYPTERIS SUBG. GONIOPTERIS

pecially abaxially, with few to many stellate or forked hairs, also with acicular hairs, rarely with only acicular hairs; indusia present or absent, often minute and easily overlooked; sporangia glabrous or often setose with acicular or stellate hairs; x⫽36. Subgenus Goniopteris is restricted to the New World tropics and subtropics, from Florida, the Antilles, and central Mexico to Bolivia and northeastern Argentina. Twenty-four species have been found in Mexico, and three additional ones are known from Guatemala and Belize and so might eventually be expected in southern Mexico: T. pellita (Willd.) Proctor & Lourteig [syn. T. asterothrix (Fe´e) Proctor], T. schippii (Weath.) A. R. Sm., and

649

T. skinneri (Hook.) C. F. Reed. Altogether, there are about 80 species in the subgenus. Goniopteris is primarily a group of lowto middle-elevation rain forests, with species becoming rare above 1500 m and non-existent above ca. 2000 m. Some species grow in semi-disturbed areas, such as cafetals and cacao plantations. Subgenus Goniopteris differs from all other subgenera of Thelypteris in the presence of forked or stellate hairs on some part of the blades. A few species lack such hairs (in Mexico, only T. ghiesbreghtii and T. hondurensis) but are placed in the subgenus because of their strong morphological similarity and undoubted relationship to stellate-hairy members of the group.

Key to the Mexican Species of Thelypteris subg. Goniopteris 1. Veins meniscioid, i.e., 3 or more pairs of veins from adjacent segments anastomosing with an excurrent vein that ends before reaching the next pair of anastomosing veins. 2. Rachises zig-zag; sporangia setulose, the hairs ca. 0.1 mm long. ................................................................................................. 57. T. rhachiflexuosa. 2. Rachises straight or nearly so; sporangia glabrous or setulose. 3. Sporangia each with several setae 0.1–0.3 mm long; costae abaxially moderately hairy, hairs mostly 0.3–0.5(–1) mm long. ................................................................................................................................................................................................................ 53. T. poiteana.

3. Sporangia glabrous (uncommonly with a few minute setulae in T. meniscioides); costae abaxially glabrous to variously hairy, hairs 0.1–1 mm long. 4. Costae, veins, and tissue between veins moderately to densely hairy abaxially, the hairs to 1.5 mm long, with many hairs ca. 1 mm long. .......................................................................................................................................................................... 40. T. ghiesbreghtii. 4. Costae, veins, and tissue between veins glabrous, or nearly so, to short-pubescent abaxially, hairs, if present, less than 0.2 mm long. 5. Costae, veins, and tissue between veins with short hairs 0.1–0.2 mm long; pinnae 3–4 lateral pairs, mostly 5–6 cm wide, crenate along margins. ........................................................................................................................................... 43. T. hondurensis. 5. Costae, veins, and tissue between veins glabrous, or near so; pinnae mostly 4–12 lateral pairs, mostly 2–5 cm wide, more or less entire. ........................................................................................................................................................... 47. T. meniscioides. 1. Veins free, or connivent at sinuses, or the lower 1–2(–3) pair(s) united with an excurrent vein to sinuses. 6. Blades lobed to pinnatifid, separate pinnae (if any, 1–3 pairs maximally) at blade bases only. ...............................................41. T. guadalupensis. 6. Blades fully pinnate except near apices. 7. Fronds arching, radicant at or near the tips and producing new fronds; pinnae mostly 3 cm long or less, entire to lobed ca. 1⁄3 the way to the costae. ........................................................................................................................................................................ 55. T. reptans. 7. Fronds not rooting at the tips (a few species producing proliferous buds back from the tips); pinnae greater than 2.5 cm long, shallowly to deeply lobed more than 3⁄4 the way to the costae. 8. Blades each without a distinct terminal pinna, distally gradually narrowed into pinnatifid apices; sporangia glabrous. 9. Costae, veins, and tissue between veins glabrous or with unbranched hairs abaxially. 10. Pinnae shorter than 7.5 cm, often shorter than 5 cm; indusia absent. ..................................................................... 39. T. blanda. 10. Pinnae usually longer than 7 cm; indusia present. 11. Indusia densely short-pubescent; proliferous buds absent at bases of distal pinnae; fronds subdimorphic, the fertile taller than the sterile and with contracted pinnae. ........................................................................ 49. T. muenchii. 11. Indusia short-setose on margins; proliferous buds present at bases of distal pinnae; fronds monomorphic, the fertile and sterile similar. .................................................................................................................................. 56. T. resiliens. 9. Costae and sometimes veins and tissue between veins abaxially with rather dense stellate or furcate hairs. 12. Fronds small, mostly less than 45 cm long. 13. Indusia present, small and hidden, bearing acicular hairs; stalked, stellate hairs present on tissue between veins. ............................................................................................................................................................................ 55. T. reptans. 13. Indusia absent; stellate hairs absent on tissue between veins, stalked on costae and rachises. ................... 59. T. stolzeana. 12. Fronds usually more than 45 cm long; indusia absent or present; stellate hairs on blades often sessile. 14. Indusia absent; stellate hairs absent on blades (both sides) between veins; sporangia with furcate hairs. ................................................................................................................................................................................ 45. T. leptocladia.

14. Indusia small but usually present, occasionally absent; stellate hairs present on both sides of blades, between veins; sporangia glabrous or with a few stellate or furcate hairs. 15. Anchor-shaped hairs, i.e., with 2–4 short recurved branches at tips present on costae and tissue between veins abaxially. ................................................................................................................................... 38. T. biolleyi. 15. Anchor-shaped hairs absent abaxially. ........................................................................................................... 42. T. hatchii.

650

THELYPTERIS 8. Blades each with a terminal pinna resembling the lateral pinnae; sporangia with or without minute setulae or furcate hairs. 16. Indusia persistent and relatively large; sori near the margins. 17. Blades subcoriaceous; costae glabrous or nearly so abaxially; pinnae incised 1⁄3 the way to the costae, proximal ones cuneate at their bases; proliferous buds often present in axils of distal pinnae; indusia glabrous. ..... 52. T. paucipinnata. 17. Blades chartaceous; costae hairy abaxially; pinnae incised ca. 1⁄2–1⁄3 the way to costae, proximal ones Ⳳ truncate at their bases; proliferous buds absent; indusia setulose on margins. .................................................... 58. T. schaffneri. 16. Indusia absent or minute and easily overlooked; sori subcostular to medial. 18. Pinnae serrulate to lobed 1⁄3 the way to the costae, cuneate at the bases; proximal veins of segments upcurved and anastomosing at acute angles, the next pairs usually also anastomosing. ...................................................... 51. T. obliterata. 18. Pinnae incised 1⁄3–4⁄5 the way to the costae, truncate or cuneate at the bases; proximal 1–2(–3) pairs of veins anastomosing at an acute or obtuse angle, or veins connivent at sinuses. 19. Rachises, costae, costules, and veins glabrous abaxially. ................................................................................... 61. T. toganetra. 19. Rachises, costae, costules, and often veins sparsely to densely hairy abaxially. 20. Pinnae long-stalked, proximal ones with stalks greater than 10 mm; blades on both sides moderately to densely setose with unbranched hairs on costae, veins, and tissue between veins. ....................... 46. T. martinezii. 20. Pinnae sessile or short-stalked less than 5 mm; blades sparsely to moderately hairy with unbranched hairs on costae and veins abaxially, glabrous adaxially. 21. Proximal pair of veins from adjacent segments united at an obtuse angle with an excurrent vein to the sinuses; sporangia setulose. ....................................................................................................... 60. T. tetragona. 21. Proximal pair of veins from adjacent segments connivent at the sinuses, or if connivent just below sinuses, then running together at an acute angle; sporangia glabrous or setulose. 22. Sporangia with minute stellate or furcate hairs; costal hairs abaxially of two Ⳳ distinct size classes: short (0.1–0.3 mm), stellate or furcate hairs and long (0.5–1 mm), stout, simple or minutely furcate hairs. ............................................................................................................... 54. T. praetermissa. 22. Sporangia glabrous or setulose with unbranched hairs; costae hairs abaxially not of two distinct size classes, mostly simple, 0.1–0.3 mm long. 23. Pinnae elliptic, gradually narrowed in their proximal third; pinnae relatively few, often 3– 8(–10) lateral pairs. ........................................................................................................... 50. T. nicaraguensis. 23. Pinnae linear-lanceolate, truncate or abruptly narrowed at their bases; pinnae usually more than 8 lateral pairs. 24. Proximal pinnae not greatly narrowed at their truncate bases; rhizomes shortcreeping. ......................................................................................................................... 44. T. imbricata. 24. Proximal pinnae abruptly narrowed at their bases, usually with two greatly reduced rounded lobes; rhizomes suberect to erect. .................................................................... 48. T. minor.

38. THELYPTERIS BIOLLEYI Thelypteris biolleyi (Christ) Proctor, Bull. Inst. Jamaica, Sci. Ser. 5: 58. 1953. Fig. 306K.

costules, and tissue between veins, also with a few sessile, appressed stellate hairs (4 arms), adaxially the tissue between veins with numerous appressed stellate hairs; sori inframedial to medial, exindusiate or indusiate; sporangia glabrous or with a few minute stellate hairs; 2n⫽72 (Jam).

Aspidium biolleyi Christ in Pittier, Prim. Fl. Costaric. 3(1): 31. 1901. Dryopteris nephrodioides (Klotzsch) Hieron. var. biolleyi (Christ) C. Chr., Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 7, 10: 248. 1913. Goniopteris biolleyi (Christ) Pic. Serm., Webbia 31: 251. 1977. Type. Costa Rica. Tuis pre`s Turrialba, Pittier 11243 (P?; isotypes CR, US!). Complete synonymy given in Smith (1983).

Rhizomes short-creeping to suberect; fronds monomorphic, or nearly so; stipes to 50 cm long or more, brownish, stellate-hairy; blades dark green, herbaceous, pustulate, to 75 cm long with pinnatifid apices; pinnae 10–20 pairs, 12–22(–28) ⫻ 2–3(–4) cm, sessile or short-stalked to 1 mm, incised ca. 0.5–0.7 the way to the costae, proximal pinnae often narrowed at their bases; segments suboblique, subfalcate, rounded at apices, ca. 4–6 mm wide; veins to 16 pairs per segment, the lowermost pair from adjacent segments anastomosing at a 60–90⬚ angle or connivent at sinuses; indument abaxially of numerous anchor-shaped (2 recurved branches at tips) hairs 0.2–0.5 mm long on costae,

Distribution. In tropical and montane rain forests; 250–1150 (–1600) m. Mexico; Guat, Bel, Hond, Nic, CR, Pan; Jam; Col, Ven, Fr Gui, Ec, Peru, n Braz.

THELYPTERIS GHIESBREGHTII

651

Specimens Examined. Chis (Breedlove 32861, DS, 32990; 58017, CAS, ENCB; Fisher 35395, F, US; Matuda 17959, F, MEXU, US; Purpus 7256, UC, US). Oax (Arreguin 703, ENCB; Mickel 1031, ENCB, MEXU, NY). Ver (Va´zquez T. 3350, XAL; Va´zquez et al. V-846, UAMIZ).

Thelypteris biolleyi differs from all other Mexican species of Goniopteris except T. hatchii in having sessile, appressed, stellate hairs on both sides of the blades, between the veins. From T. hatchii it differs in having some anchor-shaped hairs on the costae and laminae abaxially. The two Oaxacan collections have relatively large indusia and proximal pairs of veins connivent at sinuses. In these two characters they match better T. nephrodioides (Klotzsch) Proctor from the Antilles and South America. However, they bear abundant anchor-shaped hairs characteristic of T. biolleyi.

39. THELYPTERIS BLANDA Thelypteris blanda (Fe´e) C. F. Reed, Phytologia 17: 264. 1968. Figs. 306A–C. Phegopteris blanda Fe´e, Me´m. Foug. 8: 91. 1857. Dryopteris blanda (Fe´e) C. Chr., Index Filic. 254. 1905. Type. Mexico. Veracruz: Mirador, Schaffner 222 (not found at P; isotype K!). Phegopteris caespitosa E. Fourn., Mexic. Pl. 1: 89. 1872. Dryopteris caespitosa (E. Fourn.) C. Chr., Index Filic. 256. 1905. Type. Mexico. Veracruz: Co´rdoba, Bourgeau 2005 (P!; isotypes C, GH!, K!, MO!, NY!, P!, UC!, US!). Dryopteris polyphylla Copel., Univ. Calif. Publ. Bot. 19: 288, t. 37. 1941. Thelypteris polyphylla (Copel.) C. F. Reed, Phytologia 17: 305. 1968. Type. Mexico. Veracruz: below Sumidero, Copeland herb. 33 (MICH; isotypes MEXU!, UC!, US!).

Rhizomes erect or suberect; fronds monomorphic, or nearly so; stipes stramineous or darkened, 10–20(–25) cm ⫻ ca. 1 mm, glabrous; blades herbaceous, thin, dark green, pinnate-pinnatifid, ovate-lanceolate, 12–25 ⫻ 6–10(–15) cm, gradually reduced to pinnatifid apices, without proliferous buds; pinnae 7–12 pairs, sessile, to 7.5 cm ⫻ 1.5(–2) cm, the proximal pair somewhat shortened and deflexed, incised to within 1 mm of costae; segments 7–15 pairs per pinna, oblique, obtuse or acutish at apices; veins 5–6 pairs per segment, reaching margins above sinuses; indument abaxially and especially adaxially in the grooves of a few stellate hairs ca. 0.1 mm long along rachises, or rachises glabrescent, costae on both sides of blades with sparse acicular hairs 0.2–0.4 mm long, tissue between veins glabrous on both sides; sori inframedial to medial, exindusiate; sporangia glabrous; 2n⫽72 (Chis).

Distribution. Along stream banks and trails, limestone outcrops; 200–1500(–2150) m. Mexico; Guat, Bel, Hond, CR. Selected Specimens Examined. Chis (Breedlove 32991, DS, 68716, CAS). Hgo (Moore 2723, BH, GH). Oax (Hallberg 1466, NY, UC; Herna´ndez G. 2503, UAMIZ). Pue/Ver (Go´mez-Pompa s.n., MEXU). Qro (Rzedowski 46541, 46794, IEB, XAL). SLP (Cottam 10522, US). Ver (Purpus 5947, UC, US).

From other Mexican Goniopteris, T. blanda differs in having small fronds with pinnae often shorter than 5 cm long, exindusiate sori, blades each without a distinct terminal pinna, and in the blades glabrous abaxially or with a few simple hairs on the costae. Stellate or furcate hairs are found scattered primarily along the rachises, especially in the adaxial grooves.

40. THELYPTERIS GHIESBREGHTII Thelypteris ghiesbreghtii (Hook.) C. V. Morton, Contr. U.S. Natl. Herb. 38: 45. 1967. Figs. 309D, E. Polypodium crenatum Sw. var. ghiesbreghtii Hook., Sp. Fil. 5: 3. 1864. Dryopteris ghiesbreghtii (Hook.) C. Chr., Index Filic. 267. 1905 [“gheisbeghtii”]. Type. Mexico. Tabasco: Linden 1499 (K, frag. US!; isotypes B, BR!, P!). Goniopteris mollis Fe´e, Me´m. Foug. 5: 252. 1852. Dryopteris mollis (Fe´e) Maxon, Contr. U.S. Natl. Herb. 13: 18. 1909, hom. illeg., non (Jacq.) Hieron., 1907, nec Thelypteris mollis (Mett.) R. M. Tryon, 1967. Type. Mexico. Tabasco: Linden 1499 (RB, not found at P; isotypes B, BR!, K, P!).

Rhizomes creeping; fronds monomorphic or nearly so; stipes 30–40 cm long, scaleless or sparingly scaly; blades (15–)30–50 ⫻ 20–30 cm, 1-pinnate, without buds in axils of pinnae; pinnae 2–5 sessile lateral pairs and a conform terminal one, 20–30 ⫻ 4.5–8 cm, elliptic, with acuminate tips, subentire to irregularly serrulate or lobed; segments essentially absent, the pinna margins subentire to very shallowly lobed less than 1⁄10 the distance to costae; veins 10–12 pairs, 7–12 pairs meniscioid, distalmost pairs anastomosing at obtuse angles; indument abaxially of dense hairs on rachises, costae, veins, and tissue between veins, the hairs often 1–1.5 mm long, furcate and stellate hairs apparently absent throughout; sori in a double row between costules, exindusiate or each with very small indusial fragment; sporangia glabrous but intermixed with long hairs from receptacle; 2n⫽72 (Chis, CR).

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THELYPTERIS

costae, and veins, also with a few on leaf issue between veins, adaxially glabrous except for relatively sparse hairs on rachises and costae; sori inframedial, indusia small, stellate-puberulous; sporangia glabrous; 2n⫽144 (Jam).

Distribution. Wet lowland forests, often in cacao plantations; 200–1000 m. Mexico; Guat, Bel, Hond, Nic, CR, Pan. Selected Specimens Examined. Chis (Breedlove 22449, 32836, 38682, DS; Purpus 6713, F, UC). Oax (Herna´ndez G. 1009, CHAPA, MEXU, NY, 1967, CAS, CHAPA, MEXU, 1200, 1489, CHAPA). Tab (Dı´az L. 13278, UC). Ver (Va´zquez 874, XAL).

This is the only Mexican species of subg. Goniopteris that completely lacks stellate hairs, but it is clearly allied to T. poiteana, from which it also differs in having glabrous sporangia and meniscioid veins 7–12-seriate (vs. 5-seriate). From T. meniscioides it differs in the much hairier blades.

41. THELYPTERIS GUADALUPENSIS Thelypteris guadalupensis (Wikstr.) Proctor, Bull. Inst. Jamaica, Sci. Ser., 5: 60. 1953. Figs. 307A–C. Polypodium guadalupense Wikstr., Kongl. Vetensk. Acad. Handl. 1825: 435. 1826. Dryopteris guadalupensis (Wikstr.) C. Chr., Index Filic. 291. 1905, hom. illeg., non Kuntze, 1891. Type. Guadeloupe. Forsstro¨m s.n. (S). Polypodium domingense Spreng., Syst. Veg., ed. 16, 4: 51. 1827. Dryopteris domingensis (Spreng.) Maxon, Pteridophyt. Porto Rico 474. 1926. Goniopteris domingensis (Spreng.) Pic. Serm., Webbia 31: 251. 1977. Type. Hispaniola. Bertero s.n. (?).

Rhizomes short-creeping to suberect, 4–6 mm diam. (excluding stipe bases), bearing an apical tuft of pale brown, lanceattenuate, minutely stellate-puberulous scales 5–9 mm long; fronds often or usually somewhat dimorphic, the sterile ones shorter and spreading or prostrate, the fertile ones longer, longstiped, and rigidly erect; stipes of fertile fronds to 15 cm, minutely stellate-puberulous; blades lanceolate, narrowly oblanceolate, or linear-attenuate, 10–40 ⫻ 2–7(–9) cm, pinnate only at bases, with 1–3 pairs of greatly reduced, more distant proximal pinnae, blades otherwise lobed or pinnatifid; pinnae entire to crenulate or shallowly lobed, 1–5.5 ⫻ 0.5–1.5 cm, confluent at the bases with adjacent pinnae; segments deltate-oblong to linear-oblong, obtuse or somewhat acute, entire to crenate or lobed; veins 8–18 pairs, mostly pinnately branched, the proximal veins joined and forming a series of areoles along the rachises and costae; indument abaxially of moderate to dense stellate hairs on rachises,

Distribution. Epipetric on limestone rocks, associated with Tectaria heracleifolia, in low subdeciduous riparian forests; 50– 100 m. Mexico; Gr & L Ant. Specimens Examined. QR (Ceballos 1, CIQRO not seen; PalaciosRios 3695, XAL; Torres et al. 605, CIQRO not seen, UAMIZ).

This is another of the Antillean calciphiles that is known only from the Yucatan Peninsula in Mexico (Riba, 1993). Other examples with this distribution pattern are Sphenomeris clavata and Thelypteris reptans. Thelypteris guadalupensis differs from other species of subg. Goniopteris in Mexico in having the blades merely lobed to pinnatifid, or 1-pinnate only at the bases (1–3 pairs of free proximal pinnae). In blade dissection, it is similar to T. skinneri (Hook.) C. F. Reed, known from Guatemala and Belize, which differs in having the rachises and veins bearing some stout acicular hairs and in having setose (vs. stellate-hairy) indusia. A putative hybrid between T. guadalupensis and T. reptans (T. darwinii form) has been found in Quintana Roo, growing with T. guadalupensis (Palacios-Rios 3691, XAL). The hybrid is intermediate in morphology between the suspected parents.

42. THELYPTERIS HATCHII Thelypteris hatchii A. R. Sm., Amer. Fern J. 63: 118, f. 1–3. 1973. Figs. 306G, H, J. Goniopteris hatchii (A. R. Sm.) A˚. Lo¨ve & D. Lo¨ve, Taxon 26: 326. 1977. Type. Guatemala. Alta Verapaz: Senahu, Cerro Sillab, Hatch & Wilson 159 (US!).

Rhizomes erect or suberect, caudices to 25 cm long; fronds monomorphic, or nearly so; stipes brownish, 20–45 cm long, puberulous, the hairs stellate, ca. 0.1 mm long; blades dark green, 30–50 cm long, with a hastate subconfluent terminal segment; pinnae 9–14 pairs, sessile or the proximal ones with a stalk to 1 mm, to 20 ⫻ 2.7 cm, incised ca. 0.5–0.6 the way to costae,

THELYPTERIS IMBRICATA

653

proximal 1–3 pairs deflexed and slightly narrowed towards their bases, also slightly shortened; segments suboblique, subfalcate, ca. 4–6 mm wide, rounded at tips; veins 8–12 pairs per segment, the proximal pair from adjacent segments connivent at sinuses, or anastomosing at an acute angle below sinuses, less often united at oblique angles and with an excurrent vein to sinuses; indument abaxially of stellate or furcate hairs 0.1–0.2 mm long along rachises, costae, veins, and tissue between veins, hairs densest along costae, also with a few longer acicular hairs, especially along the costae and veins, on both sides with appressed sessile stellate hairs between the veins; sori medial to supramedial, with indusia small, persistent, bearing simple or furcate hairs 0.1 mm long; sporangia glabrous; 2n⫽72 (Oax). Distribution. In lowland rain forests; 100–200 m; Mexico; Bel, Hond. Specimens Examined. Tab (Acosta G. C.40, UAMIZ; Croat 40097, US). Ver (Dorantes 3932, XAL).

Thelypteris hondurensis is most closely related to T. ghiesbreghtii and T. meniscioides, and differs from those by the characters given in the diagnosis.

44. THELYPTERIS IMBRICATA Thelypteris imbricata (Liebm.) C. F. Reed, Phytologia 17: 284. 1968. Figs. 306Q, R Distribution. In wet forests, along trails and streams; 200– 1700 m. Mexico; Guat, Hond, CR. Selected Specimens Examined. Chis (Breedlove 56974, CAS). Oax (Hallberg 1599, NY, UC; Herna´ndez G. 1613, CHAPA, UAMIZ, 1640, 1706, 1748, CAS, CHAPA; Mickel 6576, MEXU, NY; Wendt et al. 3554, CHAPA). Ver (de la Sota 4017, ENCB, US; Go´mez-Pompa 5456, XAL; Ventura A. 14851, ENCB).

Thelypteris hatchii is closely allied to T. biolleyi, agreeing with that species in having appressed sessile stellate hairs on both sides of the laminae between the veins. The two species differ in the absence of anchor-shaped hairs on the costae abaxially in T. hatchii.

Polypodium imbricatum Liebm., Mexic. Bregn. 310 (reprint 158). 1849. Goniopteris imbricata (Liebm.) A˚. Lo¨ve & D. Lo¨ve, Taxon 26: 326. 1977. Type. Mexico. Veracruz: Papantla, Liebmann s.n. [Pl. Mex. 2414, Fl. Mex. 170] (C!).

Similar to T. tetragona except: fronds monomorphic or only slightly dimorphic; pinnae more numerous, mostly 10–17 lateral pairs; veins from adjacent segments connivent at sinuses, or sometimes arcuately and acutely united just below the sinuses; sporangia glabrous or infrequently sparingly setose; spores relatively small, ca. 25–40 ␮m long (perispore excluded); 2n⫽72 (Oax).

43. THELYPTERIS HONDURENSIS Thelypteris hondurensis L. D. Go´mez, Phytologia 50: 458. 1982. Type. Honduras. Go´mez 7012 (CR!). Figs. 309F, G. Differs from T. ghiesbreghtii in the short hairs on the costae, veins, and tissue between veins abaxially, the hairs 0.1–0.2 mm long; from T. meniscioides, it differs in the generally fewer (3–4 pairs), broader crenate-margined pinnae 5–6 cm wide; from T. poiteana it differs in the glabrous sporangia, shorter blade pubescence (0.1–0.2 mm), and lack of buds in the axils of proximal pinnae.

Distribution. In woods and along streams, trails, and roadsides, sometimes associated with limestone, also often in un-

654

THELYPTERIS

derstory of cafetals; 50–900(–2000) m. Mexico; Guat, Hond, CR. Selected Specimens Examined. Chis (Palacios-Rios 2831, UC). Col (Palmer 1129, NY, US). Gro (Hinton 14704, ENCB, GH, LL, NY, US; Lorea 1402, FCME). Jal (Mexia 1310, CAS, DS, GH, MICH, NY, UC, US). Nay (Lehto 24202A, ASU, ENCB). Oax (Herna´ndez G. 1862, CHAPA; Smith 404, UC). Tab (Cowan 1675, CHAPA; Gonza´lez 17141, UC). Ver (Carlson 1302, US).

This species is closely related to T. tetragona and agrees with that in having a more or less conform terminal pinna on each blade. It differs especially in having veins connivent at the sinuses or acutely united just below the sinuses, in the greater number of lateral pinnae, and usually glabrous sporangia. However, a significant number of specimens from Veracruz (e.g., Va´zquez T. 2200, XAL; Ventura A. 14710, IEB), Oaxaca (e.g., Morton & Makrinius 2333, US), and Chiapas (see Smith, 1981: 230) have sparingly to sometimes densely setose sporangia. Some of these even have the lowermost veins occasionally and seemingly sporadically uniting below the sinuses (e.g., Va´zquez T. 2200, XAL). Thelypteris imbricata is the more common species in southern Mexico, and it is likely that hybridization is taking place at some localities. Chromosome counts indicate that T. imbricata is diploid, while T. tetragona is tetraploid, so any hybrids would be sterile triploids. Additional study is needed.

45. THELYPTERIS LEPTOCLADIA Thelypteris leptocladia (Fe´e) Proctor, Bull. Inst. Jamaica Sci. Ser. 5: 61. 1953. Figs. 310A–D. Goniopteris leptocladia Fe´e, Me´m. Foug. 11: 60, t. 16, f. 1. 1866. Dryopteris leptocladia (Fe´e) Maxon, Pteridophyt. Porto Rico 476. 1926. Type. Guadeloupe. 1861, L’Herminier s.n. (P?).

Rhizomes short-creeping; fronds slightly dimorphic, 40–80 cm long; stipes ca. 15 cm ⫻ 1.5–3 mm, stellate-pubescent, hairs ca. 0.1 mm long; blades ca. 30–50 cm long, abruptly narrowed distally, each blade with a subconfluent, subhastate, pinnatifid terminal segment ca. 10 cm long; pinnae ca. 10(–17) pairs, sessile, to 8(–11) ⫻ 1.2(–2) cm, lobed or incised ca. 0.2–0.3 the way to the costae, proximal 1–3 pairs slightly deflexed, not or only very slightly narrowed at their bases, shortened, the basalmost pair ca. 1⁄4–1⁄3 the length of the longest pair; segments suboblique, ca. 2–3(–4) mm wide, rounded or blunt at tips; veins 4–5(–9) pairs per segment, the proximal pair from adjacent segments united below the sinuses, with an excurrent vein to sinuses; indument abaxially of very short-stalked-stellate or furcate hairs mostly 0.1– 0.2 mm long on rachises, costae, and veins, hairs densest along costae, acicular hairs absent or rare, tissue between veins glabrous or nearly so on both sides; sori inframedial to medial, exindusiate; sporangia setose with usually furcate hairs ca. 0.1 mm long; 2n⫽72 (Jam).

Distribution. Terrestrial in gallery forests; 0–100 m. Mexico; Gr & L Ant. Specimen Examined. QR (Palacios-Rios et al. 3692, UADY not seen, XAL-photocopy UC).

This is known from Mexico from only a single very recent collection (Oct 1997), and represents one of several Antillean species of Thelypteris that are known from Mesoamerica only from the Yucatan Peninsula. Others with this distribution include T. augescens and T. guadalupensis. Extraterritorial measurements are included parenthetically in the description. Thelypteris leptocladia differs from all other Goniopteris in Mexico by the pinnatifid blade apices, lack of hairs on the tissue between veins, exindusiate sori, and furcate hairs on the sporangial capsules. The closest relative in Mexico is probably T. reptans.

46. THELYPTERIS MARTINEZII Thelypteris martinezii A. R. Sm., sp. nov. Type. Mexico. Chiapas: Mpio. Ococingo, a 2 km al W de Crucero Corozal, camino Palenque Boca Lacantum, 180 m, 13 Feb 1985, Martı´nez S. 10123 (holotype MEXU!). Figs. 308L, M. A T. minori, T. tetragona, et T. nicaraguensi pinnis longestipitatis, petiolulis pinnarum proximalium 10–18 mm longis, setis aliquantum densioribus in costis, venis, et laminis inter venas utrinque laminarum, sporangiis dense setulosis differt; a T. praetermissa pilis non ramosis (vs. stellatis) in sporangiis differt; a T. obliterata pinnis plus incisis, petiolulis pinnarum longis differt. (Dedicated to Esteban Martı´nez S., tireless collector of ferns and other plants in southern Mexico and Mesoamerica, and co-author of a floristic list for the Lacandona region in Chiapas.)

Rhizomes not known; fronds monomorphic or nearly so, mostly 60–90 cm; stipes light brownish, 3–5 mm diam., equaling or to 2 times longer than blades; blades dark green, chartaceous, 30–40 ⫻ 30–45 cm; pinnae 4–10 lateral pairs and a nearly conform terminal one, 13–25 ⫻ 1.5–4 cm, incised 0.3–0.6 the way to costae, the proximal several pairs stalked 10–18 mm, the distal pairs sessile or nearly so; segments oblique, subfalcate, 5–7 mm wide, acutish to obtuse at tips, basal 1–4 pairs on proximal pinnae shortened and the pinna bases cuneate; veins 13–18 pairs per

THELYPTERIS MENISCIOIDES

segment, proximal 2–3 pairs from adjacent segments united below sinuses or connivent with a common vein leading to the sinuses; indument abaxially of rather dense hairs 0.15–0.2 mm long on costae and veins, hairs mostly acicular but a few furcate, especially at bases of costae, tissue between veins with erect acicular hairs 0.2–0.5 mm, adaxially with scattered to dense hairs 0.1–0.3 mm on costae and veins; sori inframedial or subcostular, exindusiate, receptacle with a few stout setae to 0.5 mm; sporangia densely setulose, with hairs 0.1 mm.

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47. THELYPTERIS MENISCIOIDES Thelypteris meniscioides (Liebm.) C. F. Reed, Phytologia 17: 292. 1968. Fig. 309M. Polypodium meniscioides Liebm., Mexic. Bregn. 211 (reprint 59). 1849. Dryopteris meniscioides (Liebm.) C. Chr., Index Filic. 277. 1905, hom. illeg., non (Willd.) Kuntze, 1891. Dryopteris liebmannii Maxon & C. V. Morton, Bull. Torrey Bot. Club 65: 348. 1938. Lectotype (chosen by Christensen, 1913: 266). Mexico. Oaxaca: Teotalcingo, Liebmann s.n. [Pl. Mex. 2407, Fl. Mex. 171] (C!).

Rhizomes short-creeping; fronds monomorphic or nearly so; stipes 30–60 cm ⫻ 2–4 mm, glabrous; blades chartaceous to subcoriaceous, 25–35 cm long, nearly as broad; pinnae (2–)4–12 lateral pairs plus a conform terminal pinna, narrowed on both sides of the middle, 15–25 ⫻ (2–)3–5(–6) cm, the margins entire, crenate, or very shallowly lobed, long-acuminate at tips; veins ca. 8–10 pairs from costules, these regularly united and meniscioid; indument on both sides absent or the costae abaxially with a few minute furcate hairs (0.1 mm long) toward bases; sori in double rows between costules, exindusiate; sporangia glabrous, rarely minutely setulose, the hairs less than 0.1 mm long; 2n⫽72 (Oax, Chis). Distribution. Terrestrial in lowland rain forests; 100–200 m. Mexico; Guat. Specimens Examined. Camp (Uca´n et al. 6915, UADY). Chis (Martı´nez S. 10110, MEXU, UAMIZ, 11527, XAL, 15332, MEXU, 15358, MEXU, UC). QR (Chater 60, MEXU, CR; Uca´n & Flores 936, CICY). Guat (Croat 24736, MO, UC).

The type and four paratypes from Chiapas were all collected from the same locality, on different dates, growing with T. obliterata (Martı´nez S. 11816, MEXU) and T. praetermissa vel aff. (Martı´nez 15360, MEXU). Another possible collection of T. martinezii from the same general area is Martı´nez S. 6864 (MEXU, UAMIZ). Davidse 20176 et al. (MEXU, MO), from Quintana Roo, is most likely a hybrid, possibly involving T. martinezii (Chater 60 collected at same locality, on same date) and T. obliterata; Torres 42 (UAMIZ), also from Quintana Roo, is similar. These collections have 1–2 pairs of veins united below the sinuses, cuneate pinna bases, proximal pinnae stalked to 5 mm and incised ca. 1⁄3 of the distance toward the costae, seven pairs of lateral pinnae and a conform terminal one, and densely but simply setose costae abaxially. Thelypteris martinezii differs from related species of subg. Goniopteris (T. minor, T. tetragona, T. nicaraguensis) by the very long-stalked pinnae, the stalks of proximal pinnae 10–18 mm long; by the rather dense setae on the costae, veins, and tissue between veins on both sides of the blades; and by the densely setulose sporangia. From T. praetermissa, T. martinezii also differs by the unbranched (vs. stellate) hairs on the sporangia. From T. obliterata, T. martinezii differs by the slightly more deeply incised pinnae, long pinna stalks, and fewer pairs of veins united below the sinuses.

Distribution. Common in lowlands along roadsides, trails, and in woods, especially near streams, often in partially disturbed sites such as understory of cafetals; 0–1250 m. Mexico; Guat. Selected Specimens Examined. Chis (Breedlove 55425, CAS; Martı´nez S. 18856, UAMIZ; Matuda 3603, GH, MEXU, MICH, NY, US). Oax (Croat & Hannon 65435, UC; Herna´ndez G. 1641, UAMIZ; Mickel 6669, MEXU, NY, UC). Tab (Cowan 3434, CAS, UC, 2993, CHAPA). Ver (Croat 40057, MEXU, MO; Dressler & Jones 98, UC).

This species differs from T. poiteana, T. ghiesbreghtii, and T. hondurensis, the most similar Mexican species, in the nearly or quite glabrous blades abaxially. In addition, T. meniscioides differs from T. poiteana in the usually glabrous sporangia and lack of buds in axils of pinnae. From T. ghiesbreghtii it also differs in having scattered stellate or furcate hairs, especially along the rachises and at bases of costae.

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THELYPTERIS

Mexican specimens are all referable to var. meniscioides; var. ternata A. R. Sm. is restricted to Guatemala.

48. THELYPTERIS MINOR Thelypteris minor (C. Chr.) A. R. Sm., Phytologia 34: 232. 1976. Figs. 307D, E. Dryopteris nicaraguensis (E. Fourn) C. Chr. var. minor C. Chr., Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 7, 10: 252. 1913. Lectotype (chosen by Smith, Phytologia 34: 232. 1976). Guatemala. Alta Verapaz: Secanquin, Maxon & Hay 3196 (US!).

Rhizomes suberect to erect, caudices to 10 ⫻ 1.5–2 cm; fronds monomorphic or nearly so, mostly 60–80(–100) cm; stipes brownish, 2–4 mm wide, as long as or slightly longer than blades; blades dark green, chartaceous, 30–45 ⫻ 20–30(–40) cm; pinnae 8–10 pairs and a nearly conform terminal pinna, 12–16(–20) ⫻ 2–2.8(–3.5) cm, incised 0.5–0.6(–0.7) the way to costae, all but the distal several pairs stalked 1–3 mm; segments oblique, subfalcate, 4–6 mm wide, acutish to obtuse at tips, basal 1–3 pairs on proximal pinnae rather abruptly shortened (to a sinuate wing on proximal 1–2 pinna pairs); veins 10–12(–15) pairs per segment, proximal pair from adjacent segments connivent at sinuses or (less frequently) united at an acute angle just below sinuses; indument abaxially of scattered to dense hairs 0.1 mm long on rachises and costae, hairs mostly acicular but a few furcate, tissue between veins glabrous on both sides; sori inframedial, exindusiate; sporangia usually setulose, with hairs 0.1 mm, occasionally glabrous.

the proximal pinnae not so greatly narrowed at the bases (pinnae abruptly narrowed in T. minor), while the latter has fewer, more elliptic, and less deeply lobed pinnae. Breedlove 42003 (DS), from Chiapas, may be a hybrid involving T. minor; the spores and sporangia seem malformed. Citation of this species from Guerrero (Lorea-Herna´ndez & Vela´zquez M., 1998) is based on a misidentification of Lorea 1402 (FCME), which is T. imbricata, and Diego 3913 (FCME), which is T. tetragona.

49. THELYPTERIS MUENCHII Thelypteris muenchii A. R. Sm., Amer. Fern J. 63: 120, f. 9– 10. 1973 [“munchii”]. Type. Mexico. Chiapas: Mu¨nch s.n. (DS!; isotype DS!). Figs. 307L, M. Rhizomes erect or suberect, apices bearing castaneous, glabrous scales; fronds subdimorphic, the fertile longer and more erect, with longer stipes, to ca. 55 cm long; stipes brownish, those of fertile fronds ca. 30 cm, sterile ca. 15–20 cm, stellatepuberulous with hairs ca. 0.1 mm, glabrescent; blades apparently subdimorphic, pustulate abaxially, the fertile ovate-lanceolate, ca. 25 ⫻ 10 cm, pinnate-pinnatifid, apices unknown but probably pinnatifid, sterile blades ovate, ca. 30 ⫻ 10 cm; pinnae sessile, proximal ones arcuate, 3–4 lowermost pairs reflexed, fertile pinnae ca. 1–1.2 cm wide, incised ca. 0.8 the way to the costae, sterile pinnae to 2.3 cm wide, incised ca. 0.8–0.9 the way to the costae; segments 2–3 mm wide, basal basiscopic segment of each proximal pinna reduced, acroscopic not reduced; veins simple, to 7 pairs per segment on fertile pinnae (to 10 pairs on sterile pinnae), proximal pairs from adjacent segments meeting margins at or just above sinuses; indument abaxially on fertile fronds of mostly simple hairs ca. 0.1 mm long on rachises, costae, costules, veins, and tissue between veins, indument absent on sterile fronds, adaxially the blades of both fertile and sterile fronds glabrous; sori medial, with indusia densely hairy, hairs 0.1 mm; sporangia glabrous.

Distribution. Terrestrial in lowland and montane rain forests; 100–1000 m. Mexico; Guat, Hond, Salv, Nic, CR, Pan. Specimens Examined. Chis (Breedlove 41979, CAS, MEXU, 42002, DS, 56576, CAS, ENCB, 57738, CAS; Chater 133, MEXU; Martı´nez S. 13615, MEXU, UAMIZ). Unverified, Doubtful, or Mistaken Reports. Tab (reported by Magan˜a, 1992, but not verified, possibly based on a misidentification). QR (Chater 1, CR, cited by Palacios-Rios, 2002c, but not verified).

Thelypteris minor is most closely related to T. imbricata and T. nicaraguensis, agreeing with these in having lowermost veins from adjacent segments connivent at the sinuses; the former has

Distribution. Habitat and elevation unknown. Mexico. Known only from the type.

THELYPTERIS OBLITERATA

This poorly known species seems to be related and most similar to T. resiliens; Thelypteris muenchii differs in lacking buds in axils of distal pinnae and in having densely short-pubescent indusia.

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obtuse angle with an excurrent vein to the sinus; perhaps it represents a hybrid with T. tetragona.

51. THELYPTERIS OBLITERATA 50. THELYPTERIS NICARAGUENSIS Thelypteris nicaraguensis (E. Fourn.) C. V. Morton, Contr. U.S. Natl. Herb. 38: 55. 1967. Figs. 307F, G. Phegopteris nicaraguensis E. Fourn., Bull. Soc. Bot. France 19: 252. 1872. Dryopteris nicaraguensis (E. Fourn.) C. Chr., Index Filic. 279. 1905. Type. Nicaragua. Chontales, Le´vy 460bis (P, frag. BM, photo UC!; isotypes BM, P!).

Rhizomes short-creeping to suberect; fronds monomorphic or slightly dimorphic, (35–)50–100(–130) cm; stipes tan to brownish, to 80 cm ⫻ 3–5 mm; blades dark green, chartaceous, 30–50 cm long, with 3–8(–10) pairs of lateral pinnae plus a conform terminal pinna; pinnae 11–20(–24) ⫻ 2.5–4 cm, proximal pair stalked to 2 mm, incised ca. 0.5 the way to the costae or less; segments falcate, rounded to subacute, entire, 6–8 mm wide, basal ones of proximal pinnae gradually reduced; veins (8–)12–18 pairs, proximal 2–3 pairs from adjacent segments connivent to sinuses; indument abaxially of mostly acicular hairs 0.1 mm long on rachises and costae, or costae glabrescent, tissue between veins glabrous on both sides; sori inframedial, exindusiate; sporangia glabrous or minutely setulose.

Thelypteris obliterata (Sw.) Proctor, Bull. Inst. Jamaica, Sci. Ser. 5: 62. 1953. Figs. 307H, J, K. Polypodium obliteratum Sw., Prodr. 132. 1788. Goniopteris obliterata (Sw.) C. Presl, Suppl. Tent. Pterid. 183. 1836. Dryopteris obliterata (Sw.) C. Chr., Index Filic. 280. 1905. Type. Jamaica. Swartz s.n. (S; possible isotype B-Willd. 19680). Polypodium faucium Liebm., Mexic. Bregn. 209 (reprint 57). 1849. Lectotype (chosen by Smith, 1981: 232). Mexico. Veracruz: “Baranca de la Hacienda de Jovo, pr. Sta. Marı´a Tlapacoyo,” Liebmann s.n. [Pl. Mex. 2416, Fl. Mex. 736] (C!).

Rhizomes short- to long-creeping; fronds monomorphic or slightly dimorphic; stipes stramineous to tan, 30–50 cm long, Ⳳ glabrous; blades chartaceous to subcoriaceous, 1-pinnate, to 50 ⫻ 40 cm, each with a conform terminal pinna, lacking buds in axils of distal pinnae; rachises with stellate hairs ca. 0.1 mm long, glabrescent; pinnae (4–)5–10 pairs, stalked to 5 mm, alternate to subopposite, 10–20 ⫻ (1.3–)2–3.5 cm, serrate (teeth oblique, acute) or shallowly lobed to 1⁄3 the way to the costae, the proximal pinnae narrowed towards their entire cuneate or rounded bases; segments 3–5 mm wide, rounded to truncate; veins 6–8 (–11) pairs, the proximal 2 pairs from adjacent segments anastomosing with an excurrent branch to sinuses, connivent with the next 2–3 pairs of veins; indument abaxially of numerous simple (a few stellate) hairs 0.1 mm long on costae, tissue between veins glabrous on both sides; sori usually inframedial to subcostular on distal part of segments, exindusiate, in 2 convergent rows; sporangia with a few simple setae 0.1 mm long; 2n⫽ca. 108, with pairs and univalents at first metaphase (Jam), possibly apogamous (Walker 1966).

Distribution. Along stream banks in montane rain forests, pineoak-Liquidambar forests; 150–1300 m. Mexico; Hond, Nic, CR, Pan. Specimens Examined. Camp (Martı´nez S. 29321, MEXU, UC, 29485, MEXU, UC). Chis (Breedlove 22378, DS; Martı´nez S. 12172, 15092, MEXU).

Thelypteris nicaraguensis is related to T. imbricata, T. minor, and allies, but differs from these by its more shallowly incised pinnae (incised about halfway to the costae or less) and fewer lateral pinna pairs. Breedlove 22378 is atypical in its rather densely acicular-hairy stipe bases and rhizome scales (densely stellatehairy in specimens from Nicaragua and further south). Martı´nez S. 29321 is atypical in having the basal pair of veins united at an

Distribution. In woods, especially near streams and in coffee understory; 100–1200 m. Mexico; Guat, Bel, Hond, Nic, CR (Sanders et al. 17618, UC); Cuba, Jam, Hisp. Selected Specimens Examined. Camp (Ortega & Uca´n 1379, UC). Chis (Breedlove 67034, CAS; Martı´nez S. 11816, MEXU). Gro (Urbina M. 8, 36, FCME). Oax (Mickel 6436, 6837, NY). Pue (Lo´pez-Forment

658

THELYPTERIS

C. 227, MEXU; Meza P. 215, XAL). Ver (Croat 44020, MO, UC; Orcutt 6627, DS).

This species most resembles and is allied to T. tetragona, T. toganetra, and T. imbricata and agrees with these in having blades with conform terminal pinnae and exindusiate sori. Thelypteris obliterata differs in having the pinnae merely serrulate or shallowly lobed 1⁄3 of the distance to the costae and in having 3–4 pairs of veins anastomosing below the sinuses with a common excurrent vein to the sinuses. It is also close to T. martinezii (q.v.). Specimens cited from Guerrero are atypical in having pinnae more deeply incised, 2⁄5–1⁄2 the way to the costae in the middle. A specimen from Veracruz (Smith s.n., UC) is atypical in having glabrous sporangia. Mickel 7180 (NY) (Mickel & Beitel, 1988: figs. 79G, H) is probably T. obliterata or a hybrid involving this species. The shorter, narrower, more deeply incised pinnae, the more truncate pinna bases, and the somewhat reduced distal pinnae suggest that it may be a hybrid with T. tetragona or T. imbricata. Both this collection and Mickel 6436 lack sporangial setae, which are found in most collections of T. obliterata. Breedlove 49555 (CAS), from Chiapas, has malformed spores, unusually long-stalked pinnae, and numerous hairs on sporangial capsules. It may be T. obliterata or a hybrid involving that. Two other Chiapas specimens (Breedlove 57382, CAS; Martı´nez 8128, MO) may also be hybrids of T. obliterata; the Breedlove collection has decidedly malformed spores. Linden s.n. (P), from Tabasco, has unusually long pinnae to 24 ⫻ 4 cm, pinnae incised ca. 2⁄5 the way to the costae, 2 or 3 pairs of veins anastomosing below the sinuses, dense hairs 0.3–0.5 mm long on costae abaxially, and glabrous sporangia; it may also be a hybrid involving T. obliterata.

52. THELYPTERIS PAUCIPINNATA Thelypteris paucipinnata (Donn. Sm.) C. F. Reed, Phytologia 17: 302. 1968. Figs. 306L–N. Nephrodium fendleri (D. C. Eaton) Hook. var. paucipinnatum Donn. Sm., Bot. Gaz.12: 134. 1887. Dryopteris paucipinnata (Donn. Sm.) Maxon, Contr. U.S. Natl. Herb. 13: 19. 1909. Lectotype (chosen by Maxon, op. cit.: 20). Guatemala. Alta Verapaz: Petet, Tu¨rckheim s.n. [Donn. Sm. 767] (US!; isolectotypes UC!, US!).

Rhizomes creeping to suberect; fronds monomorphic or nearly so; stipes darkened, ca. 35–65 cm ⫻ 3–6 mm, glabrous except for a few deciduous basal scales; blades subcoriaceous, pustulate abaxially, 45–50(–75) ⫻ ca. 20–30 cm, usually with buds in axils of distal pinnae; pinnae (3–)5–8(–13) lateral pairs and a conform terminal one, 15–30 ⫻ ca. 3(–5) cm, obliquely pinnatifid and incised ca. 1⁄3 the way to the costae, tapering gradually to narrow entire long-acuminate apices, at bases narrowly cuneate, stalked to 8 mm; segments subfalcate; veins ca. 10–14 pairs per segment, 4–6 proximal pairs from adjacent segments connivent at sinuses, or the lowest pairs united below sinuses; indument on both sides of blades absent, abaxial surfaces verrucose; sori submarginal,

with whitish, reniform, persistent, glabrous indusia; sporangia glabrous.

Distribution. Shaded limestone cliffs, streambanks and ravines in lowland and montane rain forests; 300–1300(–1600) m. Mexico; Guat, Bel. Selected Specimens Examined. Chis (Breedlove 32160, DS, 68655, CAS; Carlson 2121, US; Ton 5687, UAMIZ). Oax (Campos V. 837, MEXU; Hallberg 1331, NY; Martı´nez 2809, IEB, TEX). Ver (Bourgeau 3156, US; Finck 44, MEXU; Riba 1304, CHAPA, ENCB, UAMIZ, XAL). Unverified, Doubtful, or Mistaken Reports. Tab (reported by Magan˜a, 1992, not verified).

This species is easily distinguished from other Goniopteris, except T. schaffneri, by the combination of a conform terminal pinna on each blade, submarginal sori, and persistent indusia. From T. schaffneri it differs by the presence of a bud in the axil of a distal pinna, thicker blade texture, nearly glabrous costae abaxially, and glabrous indusia.

53. THELYPTERIS POITEANA Thelypteris poiteana (Bory) Proctor, Bull. Inst. Jamaica, Sci. Ser. 5: 63. 1953. Figs. 309K, L. Lastrea poiteana Bory, Dict. Class. Hist. Nat. 9: 233. 1825. Dryopteris poiteana (Bory) Urb., Symb. Antill. 4: 20. 1903. Goniopteris poiteana (Bory) Ching, Sunyatsenia 5: 239. 1940. Type. “Guiane.” Comm. Poiteau (P? not found). Polypodium crenatum Sw., Prodr. 132. 1788, hom. illeg., non Forssk., 1775. Type. Goniopteris crenata C. Presl, Tent. Pterid. 183, t. 7, f. 10. 1836. Jamaica. Swartz s.n. (S; isotype B-Willd. 19682-2). Complete synonymy given in Smith (1983).

Rhizomes short-creeping, ca. 1 cm diam.; fronds monomorphic or nearly so, 50–100 cm long; stipes glabrescent, 15–50 cm ⫻ 2–5 mm; blades 20–40 ⫻ 18–30 cm, 1-pinnate; pinnae 2–6 lateral pairs plus a conform terminal pinna, 12–22 ⫻ 3–6 cm, subentire or crenate or coarsely serrate, rarely shallowly lobed, the proximal ones sometimes gemmiparous in the axil, proximal ones mostly short-stalked, subcuneate at the bases, distal ones sessile and truncate or rounded; segments absent, or the shallow lobes 4–6 mm wide; veins meniscioid, with 6–9 pairs of second-

THELYPTERIS REPTANS

ary veins, the lower 3–5 pairs upcurved and acutely united with a free excurrent veinlet, usually the next few pairs alternately united in a common veinlet running to the sinuses; indument abaxially of mostly acicular hairs ca. 0.3–0.5(–1) mm long on rachises, costae, veins, and tissue between veins, adaxially the blades with similar hairs; sori in a double row between the costules, exindusiate; sporangia with 4–6 acicular setae 0.1–0.3 mm long; 2n⫽144 (Jam, Oax, Trin).

659

segments oblique, subfalcate, 4–6 mm wide, acutish to obtuse at tips, basal 1–3 pairs on proximal pinnae abruptly shortened and the pinna bases sinuate-cuneate; veins 10–16 pairs per segment, proximal 1–3 pairs from adjacent segments connivent at the sinuses; indument abaxially of two kinds of hairs along costae: long stout hairs 0.5–1 mm long, these unbranched or minutely furcate or with recurved branches at the tips, and short stellate or furcate hairs mostly ca. 0.1–0.3 mm long, tissue between veins glabrous or nearly so on both sides of blades; sori inframedial or subcostular, exindusiate, receptacle lacking setae; sporangia with minute furcate and/or stellate hairs less than 0.1 mm.

Distribution. Shaded wet banks along stream; 0–550 m. Mexico; Guat, Bel, Hond, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, n Braz, Bol. Specimens Examined. Camp (Ortega & Uca´n 1377B, UADY). Chis (Breedlove 22530, 33367, 33980, 34537, DS, 58018, CAS; Va´zquez B. & Avendano R. 1622, XAL). Oax (Mickel 5799, NY; Torres C. 11024, MEXU). Ver (Marquez R. 261, 287, XAL). Yuc (Castillo Rivero 94, UADY).

Distribution. Terrestrial in lowland rain forests; ca. 200 m. Mexico; Guat, Bel, Hond, Nic. Specimen Examined. Chis (Martı´nez 15360, MEXU).

54. THELYPTERIS PRAETERMISSA

The sole Mexican specimen is not quite typical for the species, and may in fact be a hybrid involving T. praetermissa. The sporangia appear malformed and collapsed. This specimen was collected at the same locality as several collections of T. martinezii (q.v.) and differs from that in the veins from adjacent segments being connivent at the sinuses (vs. several pairs anastomosing); minute furcate hairs on the sporangia (vs. sporangia simply setulose); the mixture of long, stout, simple hairs and short, stellate hairs on the costae abaxially; more deeply incised and shorter-stalked pinnae; and more or less glabrous tissue between the veins on both sides of the blades. Other close relatives are T. minor and T. nicaraguensis, both of which lack furcate hairs on the sporangia and more uniform pubescence on the costae abaxially.

Thelypteris praetermissa (Maxon) A. R. Sm., Phytologia 34: 232. 1976. Figs. 308H, J, K.

55. THELYPTERIS REPTANS

Unverified, Doubtful, or Mistaken Reports. QR (Castillo R. & Tun G. 220, UADY, XAL, cited by Palacios-Rios, 2002c, but not verified).

Thelypteris poiteana differs from T. ghiesbreghtii by the presence of buds at bases of proximal pinnae, fewer vein areoles between costae and pinna margins, somewhat shorter hairs on the blades abaxially, and setose sporangia. Castillo Rivero 94, the sole collection from Yucata´n, differs from most members of the species in having the costae and blades sparsely hairy; otherwise it is typical.

Dryopteris praetermissa Maxon, Proc. Biol. Soc. Wash. 57: 20. 1944. Type. Belize. El Cayo Distr., Bartlett 13104 (MICH).

Rhizomes suberect, caudices 2–3 cm diam.; fronds monomorphic or nearly so, to 130 cm long; stipes light brownish, 3–5 mm diam., nearly equalling the blades; blades dark green, chartaceous, mostly 30–60 ⫻ 30–45 cm; pinnae 8–12 pairs plus a nearly conform terminal pinna, 15–25 ⫻ 1.7–4 cm, incised 0.6–0.7 to costae, the proximal pairs stalked 2–5 mm, the distal pairs sessile;

Thelypteris reptans (J. F. Gmel.) C. V. Morton, Fieldiana, Bot. 28: 12. 1951. Figs. 308C, D. Polypodium reptans Sw., Prodr. 132. 1788, hom. illeg., non Augl., 1775. Type. Jamaica. Swartz s.n. (S?; isotypes B-Willd. 19673, microfiche UC!, photo US, UPS-THUNB 24564, microfiche UC!). Polypodium reptans J. F. Gmel., Syst. Nat. 2(2): 1309. 1791. Goniopteris reptans (J. F. Gmel.) C. Presl, Suppl. Tent. Pterid. 182. 1836. Dryopteris reptans (J. F. Gmel.) C. Chr., Index Filic. 288. 1905. Type. Jamaica. Swartz s.n. (S).

660

THELYPTERIS

Thelypteris darwinii L. D. Go´mez, Phytologia 60: 369. 1986. Type. Mexico. Yucata´n: Calcehtok, near Opiche´n, Darwin et al. 2144 (NO! isotypes CR, MEXU!).

56. THELYPTERIS RESILIENS Thelypteris resiliens (Maxon) A. R. Sm., Amer. Fern J. 63: 121. 1973. Figs 308E–G.

Rhizomes small, decumbent to suberect; fronds usually numerous, laxly arching or procumbent, subdimorphic, the sterile often rooting at the attenuate blade tips or along the rachises, mostly (10–)15–45 cm long, the fertile more erect and longer stalked, not rooting at tips; stipes 1–10(–25) cm ⫻ 0.5–1(–2) mm; blades mostly 10–30 ⫻ 3–6(–10) cm, pinnate in the proximal half or throughout; pinnae mostly 1.5–3 cm long, stalked ca. 1 mm or sessile, entire to crenate to shallowly lobed ca. 1⁄3 the way to the costae, linear to oblong or oval, truncate or sometimes subcordate at bases (fertile pinnae usually entire and more linear than sterile ones); segments 2–3 mm wide, rounded at the tips; veins 2–5(–7) pairs, the proximal pair from adjacent segments usually united with an excurrent branch to the sinuses, or veins free; indument on both sides of stellate or furcate hairs on rachises, costae, veins, and tissue between veins, the blades also with longer acicular hairs to 1 mm abaxially; sori medial, with indusia very small or seemingly sometimes absent, bearing a few long simple and/or forked hairs to 0.5 mm; sporangia glabrous; 2n⫽72 (Jam), 144 (Fla).

Rhizomes creeping to suberect, ca. 1 cm diam., with rhizome scales few, tan, 3–5 mm long, glabrous; fronds monomorphic or subdimorphic, the fertile longer and more erect, 40–70 cm long; stipes about as long as blades, stramineous, stellate-hairy; blades herbaceous, pustulate abaxially, pinnate-pinnatifid, 20–35 ⫻ 12– 20 cm, lanceolate to ovate-lanceolate, gradually reduced toward the pinnatifid apices; pinnae 15–19 pairs, sessile, 2–3 proximal pairs deflexed, these the largest, 8–12 ⫻ 1.4–3 cm, pinnatifid ca. 0.7 the way to costae, one or more distal pinnae viviparous at bases; segments oblique, subfalcate, 2–4 mm wide, acutish at apices; veins 10–13 pairs, proximal pair from adjacent segments running to sinuses or nearly so, but not connivent; indument abaxially of mostly simple (a few stellate) hairs ca. 0.1 mm long on rachises, costae, and costules, tissue between veins glabrous on both sides; sori medial, with indusia small, tan, ciliolate, persistent; sporangia glabrous.

Distribution. Usually epipetric on limestone rocks, cenotes and grutas; 50–1350 m. USA (Fla); Mexico; Guat, Bel; Bah, Gr & L Ant; Ven, Trin.

Distribution. Wet lowland forests, along streams and on limestone rocks; 200–1200 m. Mexico; Guat, Bel (Sundue 301, NY), Hond.

Selected Specimens Examined. Chis (Breedlove 21558, 32948, DS; Davidse 29730 et al., UC; Dressler 1616, GH, MEXU). Oax (Conzatti 3528, US). Ver (Croat 39442, UC; Lorence 4992, TEX). Yuc (Calzada et al. 6686, CICY, XAL, 6709, CICY; Schott 779, US).

Selected Specimens Examined. Chis (Calzada 9766, XAL; Croat 40563, MO). Oax (Herna´ndez G. 828, CAS, MEXU, NY, 1381, 1635, 1690, 1790, 1817, 1965, CAS, MEXU, 1993, 2548, MEXU). Tab (Cowan 2905, CAS, CSAT, MEXU, NY). Ver (Ramı´rez 493, 1397, XAL; Schatz & Nee 242, UC, XAL; Va´zquez 947, IEB).

This is easily distinguished from all other Goniopteris in Mexico by the arching narrow fronds often less than 5 cm wide that are usually radicant at or near the blade tips and the entire or shallowly lobed, oblong, blunt pinnae. It is a member of a predominantly Antillean complex that often grows on calcareous rocks. The type of T. darwinii and also Calzada et al. 6686 differ primarily in having non-radicant fronds (both sterile and fertile), but this condition occurs, seemingly sporadically, nearly throughout the range of the species.

Dryopteris resiliens Maxon, Field Mus. Nat. Hist., Bot. Ser. 17: 302. 1938. Type. Honduras. Atlantida: Lancetilla Valley near Tela, Standley 52893 (US!).

This is easily distinguished from all other Goniopteris in Mexico by the combination of blades lacking distinct terminal pinnae, presence of buds in the axils of distal pinnae, indument abaxially of scattered, mostly simple (a few furcate) hairs on costae, tissue between veins glabrous, and small, ciliate indusia.

57. THELYPTERIS RHACHIFLEXUOSA Thelypteris rhachiflexuosa Riba, Amer. Fern J. 79: 122, f. 1. 1989. Type. Mexico. Veracruz: Mpio. San Andre´s Tuxtla, along

THELYPTERIS SCHAFFNERI

trail from Estacio´n de Biologı´a Tropical Los Tuxtlas to Laguna Escondida, Riba 1683 et al. (UAMIZ-22623!; isotypes ENCB!, F, GH!, MEXU!, MO, UAMIZ!, UC!, XAL!). Figs. 309H, J. Rhizomes short-creeping, 5–8 mm diam., with dark brown scales, these furcate-hairy at the margins; fronds monomorphic, 40–66 cm long; stipes 23–44 cm ⫻ 1.5–3 mm, bearing furcate trichomes at the bases and glabrous or glabrescent in the distal third; blades papyraceous to chartaceous, 1-pinnate, each with a conform terminal pinna, 18–44 ⫻ 17–28 cm, with rachises slightly to evidently flexuous; pinnae alternate, 3–6 pairs plus the terminal one, 9–23 ⫻ 1.9–5 cm, the proximal ones short-stalked, distal pinnae sessile or slightly adnate basiscopically, bases unevenly cuneate with acroscopic side excised, margins entire, crenate, or very shallowly lobed, long-acuminate, costules 2.5–5 mm apart; segments, if any, 2–3.5 mm wide, rounded at tips; veins meniscioid, with 7–13 pairs of secondary veins, the proximal 4–8 pairs united with a free excurrent veinlet and the next few pairs united in common veinlets running to the margins or the sinuses; indument abaxially absent on rachises, costae, veins, and tissue between veins, or with minute simple hairs along costae; sori medial to inframedial, exindusiate; sporangia with simple setae 0.1 mm or less.

661

58. THELYPTERIS SCHAFFNERI Thelypteris schaffneri (Fe´e) C. F. Reed, Phytologia 17: 312. 1968. Figs. 309A–C. Nephrodium schaffneri Fe´e, Me´m. Foug. 8: 108. 1857. Dryopteris schaffneri (Fe´e) Index Filic. 291. 1905. Type. Mexico. [Veracruz:] Mirador, Schaffner 244 (P?, not seen). Aspidium varians Mett. ex Kuhn, Linnaea 36: 114. 1869. Type. Mexico. San Luis Potosı´: Virlet d’Aoust 82 (B; isotype P).

Rhizomes short-creeping; fronds monomorphic or nearly so, 65–110 cm long; stipes stramineous to brownish, ca. 35–65 cm ⫻ 2.5–5 mm, with scattered to dense furcate and stellate hairs ca. 0.1 mm long, also with a few basal scales; blades chartaceous, pustulate on abaxial surfaces, 1-pinnate-pinnatifid, 25–45 ⫻ ca. 25–35 cm, lacking buds in axils of distal pinnae; pinnae 3–7 lateral pairs and a conform terminal one, 15–25 ⫻ ca. 2.5–4 cm, incised ca. 1⁄2–2⁄3 the way to the costae, stalked 3–15 mm, somewhat narrowed to truncate at the bases, tapering gradually to entire long-acuminate apices; segments subfalcate, acute or obtuse; veins ca. 15–18 pairs per segment, 3–4 proximal pairs from adjacent segments connivent at sinuses, or the lowest 1–2 pairs acutely united just below sinuses; indument abaxially of spreading simple hairs 0.1–0.5 mm on costae, costules, and veins, tissue between veins glabrous or nearly so, rachises with stellate hairs 0.1–0.2 mm and a few longer unbranched hairs; sori submarginal, with indusia whitish, persistent, reniform to athyrioid, marginally setose; sporangia glabrous.

Distribution. Terrestrial in wet lowland forests; 0–800 m. Mexico. Specimens Examined. Chis (Pe´rez Farrera 1967, CHIP, MEXU, UAMIZ, not seen, photo UC). Ver (Calzada 337, CAS, UAMIZ; de la Sota 4019, ENCB; Dressler & Jones 98, MEXU, NY, UC, US; Ibarra Manriquez 1015, MEXU, 3300, ARIZ, CAS, MEXU, UAMIZ; Lorence 3277, MEXU; Lyonnet 500900004, MEXU; Palacios-Rios 1, UAMIZ; Riba 441, UAMIZ).

This species is very close to T. meniscioides, differing primarily in the zig-zag rachises and in the copiously setulose sporangia. Some specimens of T. meniscioides from the area of sympatry (e.g., Dressler & Jones 98, UC, from Mpio. San Andre´s Tuxtla) tend towards T. rhachiflexuosa in that they have very slightly flexuous rachises and setulose sporangia. The differences between the two species may be controlled by only a few alleles that have become locally fixed in a relatively small area of Veracruz.

Distribution. Terrestrial in lowland and montane rain forests; 800–1650 m. Mexico. Selected Specimens Examined. Qro (Rubio 1234, IEB, UC, XAL; Rzedowski 42842, 46405, IEB, XAL; Servı´n 883, IEB, UC). SLP (Copeland herb. 28, GH, MEXU, MICH, UC, US; Rzedowski 23127, ENCB). Tam (Palmer 298, F, GH, MO, NY, US). Ver (Go´mez-Pompa & Riba 291, 296, MEXU; Purpus 5944, BM, MO, NY, UC, US).

Thelypteris schaffneri is readily distinguished by its submarginal sori, relatively large, persistent, often subathyrioid, setosemargined indusia, abaxially hairy costae and veins, and lack of buds in axils of distal pinnae. It is most closely related to T. paucipinnata, with which it may be sympatric in Veracruz. A specimen from Veracruz, Scheinvar et al. 678 (MEXU), is some-

662

THELYPTERIS

what intermediate between the two species in pinna dissection, but agrees better with T. schaffneri in lacking buds in axils of distal pinnae, having pubescent costae abaxially, sparingly setulose-margined indusia, thinner blade texture, and truncate pinna bases.

59. THELYPTERIS STOLZEANA Thelypteris stolzeana A. R. Sm., Phytologia 34: 231. 1976. Type. Guatemala. Alta Verapaz: Between Coba´n and San Pedro Carcha´ along Rı´o Carcha´, Standley 90107 (F!). Figs. 308A, B. Rhizomes erect, caudices ca. 1–1.5 cm; fronds monomorphic, 25–45 cm long; stipes brownish, 10–20 cm ⫻ 1–2 mm, with 2–3cleft and a few simple hairs ca. 0.1–0.2 mm, glabrescent; blades dark gray-green, chartaceous, not pustulate, 15–25 cm long, gradually reduced distally to confluent, pinnatifid, somewhat elongate apices, lacking buds in axils of distal pinnae; pinnae 8–9 pairs, sessile, to ca. 5 ⫻ 1.5 cm, incised ca. 0.5–0.7 the way toward the costae, proximal 1–2 pairs slightly reduced and deflexed, narrowed at their bases; segments suboblique, not or only weakly falcate, 2–3 mm wide, rounded to truncate at tips; veins 4–6 pairs per segment, proximal pair from adjacent segments meeting margins within 0.3 mm of sinuses; indument abaxially of 2–4cleft and some simple, stalked hairs mostly 0.1–0.2 mm long on rachises and costae, veins with short, simple hairs, tissue between veins glabrous; sori inframedial to medial, exindusiate; sporangia glabrous.

Distribution. Terrestrial in montane rain forests, along river banks; 1300 m. Mexico; Guat. Specimens Examined. Chis (Breedlove 22355, DS, MEXU).

This is most closely related to T. blanda, but differs from that in having much more obvious furcate and stellate hairs on the rachises and costae abaxially (a few furcate hairs along the rachises only in T. blanda). In addition, the two can be distinguished by the deeper cutting of the pinnae (resulting in the lowermost veins meeting the margins well above the sinuses) in T. blanda.

60. THELYPTERIS TETRAGONA Thelypteris tetragona (Sw.) Small, Ferns S.E. States 256. 1938. Figs. 306D–F. Polypodium tetragonum Sw., Prodr. 132. 1788. Goniopteris tetragona (Sw.) C. Presl, Suppl. Tent. Pterid. 183. 1836. Dryopteris tetragona (Sw.) Urb., Symb. Antill. 4: 20. 1903. Type. Jamaica. Swartz s.n. (S; isotype B-Willd. 19708-3, microfiche UC!, photo US). Complete synonymy given in Smith (1983).

Rhizomes short-creeping, caudices ca. 1 cm diam.; fronds subdimorphic, the fertile ones erect, to 100 cm long, long-stipitate, the sterile ones shorter, arching, mostly 50–100 cm long; stipes stramineous to tan, to 60 cm ⫻ 2–5 mm, stellate-pubescent; blades herbaceous, pinnate-pinnatifid, each with a subconform terminal pinna, 25–45(–55) ⫻ 12–25(–30) cm, lacking buds in axils of distal pinnae; pinnae 6–10(–12) pairs, subopposite to alternate distally, sessile to stalked 1 mm, proximal pairs somewhat narrowed at bases, 7–15(–18) cm long, sterile ones mostly 2–3 cm wide, fertile ones 1–2(–2.5) cm wide, pinnatifid 0.5–0.7 to the costae; segments oblique, subfalcate, rounded at apices, ca. 4–6 mm wide; veins 6–10 pairs, the proximal pair from adjacent segments usually united at an obtuse angle with an excurrent vein to sinuses; indument on both sides of blades of mainly acicular hairs 0.1–0.3 mm long and a few minute furcate and/or stellate hairs 0.1 mm long on rachises and costae, veins and tissue between veins glabrous; sori inframedial, exindusiate; sporangia with simple hairs 0.1 mm long; 2n⫽144 (Fla, Jam, SLP, Ver, Chis, Trin).

Distribution. In woods and along streams, trails, and roadsides, sometimes associated with limestone, also sometimes in coffee understory; 0–1000 m. USA (Fla); Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Guy, Sur, Ec, Peru, Braz, Bol. Selected Specimens Examined. Camp (Lundell 1223, US). Chis (Breedlove 42847, DS). Col (Palmer 1129, US). Gro (Morales O. 3, FCME, IEB). Hgo (Clark 6881, MO, NY). Mich (Lorea 1758, IEB). Nay (Te´llez 12485, MEXU). Oax (Mickel 5802, UC). Pue (Ramı´rez C. 566, MEXU). Qro (Rubio 2012, IEB, UC). QR (Bye 12002, UC). SLP (Pringle 3959, MEXU, UC). Sin (Lamb 493, DS, GH, NY). Tab (Cowan 1675a et al., CAS, CHAPA, CSAT, MEXU, MO). Tam (Yatskievych 83336, ARIZ, UC). Ver (Smith 412, UC). Yuc (Rico-Gray 668, XAL).

THELYPTERIS SUBG. MENISCIUM

This species occupies habitats similar to those of T. imbricata, its closest relative (which see). In addition to the differences in the key, T. tetragona usually has fewer lateral pinna pairs and the proximal pinnae are often slightly shortened and deflexed. The lowermost veins from adjacent segments are usually (but not always) united and produce an excurrent vein to the sinus. Even on a single frond there is often considerable variation (e.g., Lorea 1758, from Michoaca´n, and Pringle 3959, from San Luis Potosı´), with some vein pairs united below the sinuses and some free or connivent at the sinuses, on the same frond. Most Mexican specimens have at least some setose sporangia. Morales O. 3 (FCME, IEB), from Guerrero, and Ortega O. et al. 857 (XAL), from Tabasco, are unusual in having small, setose, spatulate indusia. Mickel 5786 (NY) and Torres C. 10137 (MEXU), both from the same general area in Oaxaca, may be hybrids involving this species. In blade cutting (subopposite pinnae, distinct terminal pinna) and venation (one pair of veins united below each sinus, with an excurrent vein to sinus) they look most like T. tetragona, but they lack stellate hairs on all parts of the plant (including rachial grooves and rhizome scales), or have only very few furcate hairs in the rachial grooves. The Torres collection has about half its spores well formed, the other half shriveled or empty. Sousa 1486 (MEXU), from Tuxtepec, Oaxaca, has the appearance of a hybrid between T. tetragona and perhaps T. obliterata, judging by the intermediate venation and depth of cutting of the pinnae; the sporangia seem empty. Two even more problematic specimens are: Acevedo R. & Castillo C. 268 (IEB), from Edo. Veracruz, Mpio. Atoyac, Ejido La Esperanza, 750 m; and Martı´nez Caldero´n 576 (NY), from Edo. Oaxaca, Chiltepec, Tuxtepec. In blade shape (fertile only, both with subconform terminal pinnae) and cutting they appear likely to belong to subg. Goniopteris, but the blades completely lack stellate or furcate hairs. There are no indusia and the sporangia are very densely setose and appear almost acrostichoid, more so than any other species of Goniopteris known to us. Of known species, these two collections most closely approach T. tetragona, but the pinnae are subentire to shallowly lobed, and only ca. 5 mm wide. The spores appear well formed and similar to those of other species of Goniopteris (and unlike those of subg. Stegnogramma). Perhaps they represent an undescribed species, but the material at hand is inadequate and/or bizarre (i.e., outside the norm of expectation within the genus Thelypteris or subg. Goniopteris) for description.

61. THELYPTERIS TOGANETRA Thelypteris toganetra A. R. Sm., Amer. Fern J. 63: 118. 1973. Figs. 306O, P. Goniopteris toganetra (A. R. Sm.) A˚. Lo¨ve & D. Lo¨ve, Taxon 26: 326. 1977. Type. Mexico. Veracruz: Atoyac River, Copeland herb. 34 (UC! isotypes BM, GH!, MEXU!-3 sheets, MICH!, US!).

Rhizomes long-creeping to suberect; fronds monomorphic or nearly so, to 70 ⫻ 30 cm, each with a conform terminal pinna; stipes 15–35 cm ⫻ 2–4 mm, glabrous on both sides, or adaxially

663

in the grooves with a few stellate hairs less than 0.1 mm long; blades 25–40 ⫻ 12–30 cm, lacking buds in axils of distal pinnae; pinnae 10–16 pairs, 12–30 ⫻ 1–3 cm, incised 0.5–0.7 to the costae, sessile, proximal ones narrowed but still lobed at the bases; segments subfalcate, acutish at apices; veins 7–15 pairs, 1–2 proximal pairs from adjacent segments connivent at or just below sinuses, not truly united; indument on both sides absent on rachises, costae, and veins, or adaxially the rachises and costae with a few simple and even fewer stellate hairs ca. 0.1–0.2 mm long, costules, veins, and tissue between veins glabrous on both sides, but sometimes with scattered sessile reddish glands along costules and veins abaxially; sori medial, exindusiate; sporangia glabrous; 2n⫽72 (Chis).

Distribution. Shaded wet banks along streams; 50–1300 m. Mexico; Guat, Bel. Selected Specimens Examined. Chis (Breedlove 34223, DS, ENCB, TEX; Martı´nez S. 14633, UAMIZ). Oax (Herna´ndez G. 712, CAS, CHAPA, ENCB, MEXU, NY; Mickel 5800, NY, UC). SLP (King 4380, LL, MICH, TEX, UC, US; Palmer 660, US). Tab (Gonza´lez L. GH145, MEXU). Tam (Harriman 10930 et al., OSH). Ver (Riba 860, ENCB, MEXU; Ventura A. 3601, ARIZ, CAS, CHAPA, ENCB, MEXU, NY).

Thelypteris toganetra is most similar and closely related to T. tetragona and T. imbricata, differing from the former in the veins connivent at the sinuses, or uniting at an acute angle just below the sinuses, and in the glabrous sporangia. From both species T. toganetra differs in the glabrous blades, with hairs absent on all parts of the blades abaxially.

THELYPTERIS SUBG. MENISCIUM Thelypteris subg. Meniscium (Schreb.) C. F. Reed, Phytologia 17: 254. 1968. Meniscium Schreb. in Linnaeus, Gen. Pl., ed. 8, 2: 757. 1791. Dryopteris subg. Meniscium (Schreb.) C. Chr., Index Filic. XXII. 1906. Thelypteris subg. Cyclosorus sect. Meniscium (Schreb.) C. V. Morton, Amer. Fern J. 53: 154. 1963. Type: Thelypteris reticulata (L.) Proctor [⬅ Meniscium reticulatum (L.) Sw.] [⬅ Polypodium reticulatum L.].

Rhizomes creeping, stout, often 1–2(–2.5) cm diam.; fronds of most species large, often 100–200 cm long, sometimes subdi-

664

THELYPTERIS

morphic; blades 1-pinnate (simple in 2 spp.), each with a terminal often conform pinna, some species bearing buds in axils of proximal or rarely distal pinnae; pinnae entire, or the margins undulate, serrate, or uncinate-serrate; lateral veins of pinnae straight and giving rise to numerous straight, arcuate, or subsigmoid regularly anastomosing secondary veins, producing an excurrent veinlet, thus creating a regular series of areoles between costae and pinna margins); indument of simple hairs, these never stellate, relatively sparse and short, sometimes curved, often confined to costae on both sides and main lateral veins abaxially; sori confluent along the secondary veins and where they join, exindusiate; sporangia glabrous or with setae arising from the stalks or receptacles; x⫽36.

Subgenus Meniscium comprises about 23 species confined to the American tropics and subtropics, from southern Mexico to northern Argentina and Paraguay, Florida, and the Antilles. Most species occur at elevations below 1500 m, along streams and wet banks. Meniscium is characterized by the regularly anastomosing venation that produces straight and very regular rows of areoles between costae and margins. A few species of subg. Goniopteris have similar venation, but most of these have stellate hairs on some part of the blades, and most have round sori in a bilateral series between main lateral veins and between the costules; the sori are elongate, often lunate along the cross-veins in subg. Meniscium.

Key to the Mexican Species of Thelypteris subg. Meniscium 1. Pinna margins uncinate-serrate, at least toward apices; plants of wet ditches and swamps. .................................................................... 65. T. serrata. 1. Pinna margins entire, undulate, or infrequently crenulate; plants of forests or streambanks. 2. Pinnae mostly less than 2 cm wide; areoles usually 4–8(–10) between costae and pinna margins; costae abaxially with adpressed hairs. ........................................................................................................................................................................................................ 62. T. angustifolia. 2. Pinnae mostly more than 2 cm wide; areoles 8 or more between costae and pinna margins; costae abaxially glabrescent or with Ⳳ spreading hairs. 3. Areoles of sterile pinnae almost as long as broad; secondary veins (arising from costules) nearly straight; pinnae Ⳳ parallelsided for most of their length. ............................................................................................................................................................... 63. T. falcata. 3. Areoles of sterile pinnae shorter than broad; secondary veins (arising from costules) curving and arcuate; pinnae broadest at the bases. 4. Receptacles and sporangial stalks bearing stiff acicular hairs. ................................................................................................. 66. T. standleyi. 4. Receptacles and sporangial stalks lacking hairs. ....................................................................................................................... 64. T. reticulata.

62. THELYPTERIS ANGUSTIFOLIA Thelypteris angustifolia (Willd.) Proctor, Bull. Inst. Jamaica, Sci. Ser. 5: 57. 1953. Figs. 311C, D. Meniscium angustifolium Willd., Sp. Pl., ed. 4, 5(1): 133. 1810. Dryopteris angustifolia (Willd.) Urb., Symb. Antill. 4: 21. 1903. Type. Venezuela. Caracas, Bredemeyer s.n. (B-Willd. 19574). Polypodium salicifolium Vahl, Eclog. Amer. 3: 51. 1807, non Thelypteris salicifolia (Wall. ex Hook.) C. F. Reed, 1968. Type. Montserrat. Ryan s.n. (C).

Rhizomes short- to long-creeping; fronds subdimorphic, the fertile with longer stipes and narrower pinnae, mostly 25–80 (–130) cm long; blades 1-pinnate, each with a Ⳳ conform terminal pinna, lacking buds in axils of pinnae; pinnae (4–)7–18 (–22) pairs, Ⳳ equal in size, the proximal ones stalked to 3 mm, (5–)7–15(–18) ⫻ (0.5–)1–1.8(–2.3) cm, widest in the middle, tapering on both sides to acuminate apices and cuneate bases, margins entire to undulate; lateral veins of fertile pinnae ca. 13–18 per 3 cm, the veins meniscioid, forming usually 4–8(–13) series of areoles between costae and margins, areoles shorter than broad, the secondary veins curved or subsigmoid; indument abaxially of adpressed, curved or crispate hairs on costae and costules, the veins and tissue between veins on both sides glabrous or glabrescent; sori linear or lunate in single rows between costules at confluence of secondary veins; sporangia and receptacles glabrous.

Distribution. Along streams in lowland rain forests; 0–600 m. Mexico; Guat, Hond, Nic, CR, Pan; Gr Ant, Montserrat; Col, Ven, Ec, Peru, Braz, Bol, Parag. Specimens Examined. Chis (Castillo C. et al. 3991, XAL). Gro (Lorea 2855, FCME, IEB). Tab (Rovirosa 826, MEXU). Ver (Pe´rez 1023, UAMIZ; Reeves R5913, ASU).

Compared to other species of subg. Meniscium, this species has narrower pinnae, with fewer vein areoles between the costae and pinna margins. More than any other species in subg. Meniscium, it is adapted to streamsides and occasional flooding, i.e., rheophytic habitats.

THELYPTERIS RETICULATA

63. THELYPTERIS FALCATA Thelypteris falcata (Liebm.) R. M. Tryon, Rhodora 69: 6. 1967. Figs. 311A, B. Meniscium falcatum Liebm., Mexic. Bregn. 183 (reprint 31). 1849. Dryopteris falcata (Liebm.) C. Chr., Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 7, 10: 270. 1913. Lectotype (chosen by Smith, 1981: 234). Mexico. Oaxaca: Dto. Chinantla, Liebmann s.n. [Pl. Mex. 2756, Fl. Mex. 680] (C!; isolectotypes C!, K!, US!). Meniscium juergensenii Fe´e, Me´m. Foug. 5: 223. 1852 [“jungersenii”]. Dryopteris juergensenii (Fe´e) Maxon & C. V. Morton, Bull. Torrey Bot. Club 65: 360. 1938. Type. Mexico. Ju¨rgensen 917 (not found at P). Complete synonymy given in Smith (1983).

Rhizomes short-creeping; fronds mostly 85–125(–200) cm long; blades 1-pinnate, each with a Ⳳ conform terminal pinna, lacking buds in axils of pinnae; pinnae 8–12(–20) pairs, Ⳳ equal in size, the proximal ones long-stalked (to 1–3 cm), often curved, 15–32(–40) ⫻ (1.8–)2–4(–5) cm, widest in the middle, tapering on both sides to acuminate apices and cuneate bases, margins entire to undulate; lateral veins of fertile pinnae ca. 8–12 per 3 cm, the veins meniscioid, forming usually 8–12(–17) series of areoles about as long as broad between costae and pinna margins, the secondary veins Ⳳ straight and uniting at an obtuse angle, an excurrent vein often dividing areole into two subequal rhomboid halves; indument on both sides of sparse hairs 0.2–0.3 mm long on costae, or costae glabrescent, veins and tissue between veins glabrous or glabrescent on both sides; sori arcuate, in a single row between costules at confluence of secondary veins; sporangia glabrous.

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is T. reticulata) Pue (reported by Smith in Mickel & Beitel, 1988, but not verified). QR (Torres P. et al. 21, CIQRO, cited by Palacios-Rios, ms., but not verified).

Thelypteris falcata differs from T. serrata by the entire or slightly undulate pinna margins, long-cuneate pinna bases, terminal pinnae nearly as long as the medial ones, and glabrous or glabrescent costae abaxially. From T. reticulata it differs in the areoles being about as broad as long, straight secondary veins, and long-cuneate pinna bases. Occasional specimens have undulate-crenate margins, e.g., Vera Santos 3394 and Martı´nez C. 544.

64. THELYPTERIS RETICULATA Thelypteris reticulata (L.) Proctor, Bull. Inst. Jamaica, Sci. Ser. 5: 63. 1953. Figs. 311E–G. Polypodium reticulatum L., Syst. Nat., ed. 10, 2: 1325. 1759. Meniscium reticulatum (L.) Sw., J. Bot. (Schrader) 1801(2): 274. 1803. Dryopteris reticulata (L.) Urb., Symb. Antill. 4: 22. 1903. Type. LINN 1251.25, from an unknown locality, possibly Jamaica (Proctor, Fl. L. Antill. 2: 309. 1977).

Rhizomes short- to long-creeping; fronds to 200 cm long or more, somewhat dimorphic, the fertile ones with longer stipes; blades 50–120 ⫻ 20–60 cm, occasionally with buds borne in the axils of proximal pinnae; pinnae to 20⫹ pairs, widest at or near bases, to ca. 30 ⫻ 2–5 cm, at the bases unequally rounded or broadly cuneate, the margins entire, wavy, or slightly crenate; lateral veins straight to usually arcuate or subsigmoid, with 12 or more pairs anastomosing and giving rise to excurrent veinlets, the areoles shorter than broad; indument abaxially of sparse hairs on costae, costules, and sometimes veins, the hairs mostly 0.3 mm long or less; sori arcuate along veins at the anastomoses; sporangia glabrous; 2n⫽144 (Jam, PR).

Distribution. Along trails and in partially cut-over, shaded woods; 50–750 m. Mexico; Guat, Bel, Hond, Nic, CR, Pan; Cuba; Col, Ec, Peru, Bol. Specimens Examined. Chis (Martı´nez S. 15913, IEB, UAMIZ). Oax (Hallberg 1614, NY, UC; Herna´ndez G. 1982, CAS, CHAPA, MEXU; Martı´nez C. 544, ENCB, MEXU, 814, CAS, ENCB, MEXU, UAMIZ, XAL; Vera Santos 3394, ENCB, MICH, TEX). Tab (Colorado 12, UAMIZ). Ver (King 1110, ENCB, TEX, US; Lot 1593, MEXU, MO). Unverified, Doubtful, or Mistaken Reports. Chis (Martı´nez S. 7675, cited by Riba et al. (Amer. Fern J. 77: 70. 1987) as this species,

Distribution. In wet woods, especially near streams; 150–300 (–800) m. USA (Fla); Mexico; Gr & L Ant; Col, Ven, Trin. Selected Specimens Examined. Chis (Martı´nez S. 7675, MEXU, MO, UAMIZ, 10079, 11111, MEXU, 17915, MEXU, UAMIZ; Va´squez & Avendeno R. 1637, XAL). Oax (Herna´ndez G. 385, CAS, CHAPA, MEXU, NY; Mexia 9202, ARIZ, F, GH, MICH, NY, U, UC, US; Solano C. 315, TEX, UAMIZ). QR (Castillo R. 218, XAL; Torres 21, MEXU, UAMIZ).

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THELYPTERIS

Thelypteris reticulata differs from T. serrata by the usually entire or slightly crenate pinnae. From T. falcata it differs by the areoles being shorter than wide and the curving (sigmoid) secondary (cross) veins. Several collections cited (e.g., Mexia 9202, Solano C. 315) are atypical of the species. They have narrower pinnae and vary toward large forms of T. angustifolia (Willd.) Proctor. However, that species generally has still narrower pinnae that are more narrowly cuneate at the bases.

65. THELYPTERIS SERRATA Thelypteris serrata (Cav.) Alston, Kew Bull. 1932: 309. 1932. Figs. 311H, J. Meniscium serratum Cav., Descr. Pl. 548. 1802. Dryopteris serrata (Cav.) C. Chr., Index Filic. 291. 1905. Type. Cuba. Havana, Guio s.n. (MA). Complete synonymy given in Smith (1983).

Rhizomes short-creeping; fronds to ca. 200 cm long or more; blades 1-pinnate, each with a lanceolate terminal pinna, distal pinnae gradually shortened, buds occasionally borne in axils of proximal pinnae; pinnae 15–25 pairs, the proximal ones shortstalked to 4 mm, largest 15–25 ⫻ (2–)2.5–3.5(–4.5) cm, the margins serrate or uncinate-serrate, fertile pinnae often somewhat narrower than sterile ones; lateral veins producing arcuate or subsigmoid secondary veins, with 10–15 pairs anastomosing and giving rise to short excurrent veinlets, the areoles much shorter than broad; indument abaxially of hairs ca. 0.2 mm long on costae, costules, and sometimes veins and tissue between veins, adaxially glabrous except for the sparsely hairy costae; sori arcuate, along veins at anastomoses; sporangia glabrous; 2n⫽72 (Fla, Trin).

Distribution. Wet roadside banks and swamps, sometimes in standing water; 0–550 m. USA (Fla); Mexico; Guat, Hond, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol, Parag, n Arg. Selected Specimens Examined. Chis (Gilly & Herna´ndez X. 163, MICH, MSC, US; Ventura A. 21352, ENCB, XAL). Gro (Fonseca J. 1076, FCME). Nay (Ortega 75, US). Oax (Mickel 6816, NY; Torres C. 8033, MEXU). Pue (Sa´nchez M. 907, MEXU). QR (Torres 636,

UAMIZ). Ver (King 997, LL, TEX, US; Purpus 8602, DS, MO, NY, UC). Tab (Cowan 3301, CAS, CHAPA).

This differs from the other Mexican species of subg. Meniscium by the uncinate-serrate pinna margins and the gradually reduced distal pinnae, each blade terminating in a relatively small narrowly deltate terminal pinna. The specimen cited from Nayarit is juvenile, sterile, and not determinable with certainty.

66. THELYPTERIS STANDLEYI Thelypteris standleyi (Maxon & C. V. Morton) R. M. Tryon, Rhodora 69: 8. 1967. Figs. 311K–N. Dryopteris standleyi Maxon & C. V. Morton, Bull. Torrey Bot. Club 65: 368. 1938. Type. Guatemala. Izabal: Near Quirigua´, Standley 24126 (US!).

Rhizomes short-creeping; fronds to 200 cm long, with stipes longer than blades; blades 1-pinnate, to 70 ⫻ 40 cm, gradually reduced in width distally, lacking buds in axils of pinnae; pinnae 12–17 lateral pairs plus a subsimilar terminal pinna, 10–22 ⫻ 2.3– 4 cm, sessile, rounded or truncate at bases, margins entire or slightly repand, acuminate at apices; lateral veins producing arcuate or subsigmoid secondary veins, these acutely anastomosing and forming a series of 9–15(–20) areoles between costae and margins, excurrent veinlets free, the areoles much shorter than broad; indument abaxially of curved to straight acicular hairs 0.2–0.3 mm long on costae, costules, veins, and sometimes tissue between veins, adaxially the blades glabrous except for scattered hairs along costae; sori oblong and confluent where secondary veins anastomose; sporangia bearing acicular solitary hairs from the sporangial stalks.

Distribution. Terrestrial in wet forested ravines; 400–1200 m. Mexico; Guat, Hond, CR, Pan; Col, Ec, Peru, Bol. Reported from Nicaragua by Go´mez (1976: 51), but no specimens have been seen. Specimens Examined. Chis (Seler & Seler 2013, GH, US). Jal (Rzedowski & McVaugh 1357, ENCB).

THELYPTERIS PILOSA

Thelypteris standleyi differs from all other species of Meniscium in Mexico by the sporangia bearing setae from the sporangial stalks. From T. falcata and T. angustifolia it differs by the rounded or truncate (vs. cuneate) pinna bases.

THELYPTERIS SUBG. STEGNOGRAMMA Thelypteris subg. Stegnogramma (Blume) C. F. Reed, Phytologia 17: 254. 1968.

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mal ones short-stalked (ca. 1 mm), incised 1⁄3–2⁄3 the way to the costae; segments oblique, oblong to deltate, mostly obtuse at tips; veins 3–6(–10)-jugate, the two proximal ones from adjacent segments running to sinuses; indument on both sides of blades of long stout hairs mostly 0.3–1.5 mm long on costae, costules, veins, and sometimes tissue between veins; sori elongate along veins, mostly 1.5–4 mm long, exindusiate; sporangia often minutely setose or glandular on capsule, hairs ca. 0.1 mm long; 2n⫽72 (Oax).

Stegnogramma Blume, Enum. Pl. Javae 172. 1828. Type: Stegnogramma aspidioides Blume [⫽ Thelypteris stegnogramma (Blume) C. F. Reed]. Leptogramma J. Sm., J. Bot. (Hooker) 4: 51. 1841. Dryopteris subg. Leptogramma (J. Sm.) C. Chr., Index Filic. XXI. 1906. Type: Leptogramma totta (Willd.) J. Sm. [⬅Polypodium tottum Willd.] ⫽ Thelypteris pozoi (Lag.) C. V. Morton.

Rhizomes short-creeping to erect; fronds pinnate, the pinnae entire to deeply pinnatifid, the proximal ones not or little reduced, distal ones broadly adnate to, and often decurrent on, the rachises and merging with apical portion of the blades, lacking buds in axils of pinnae; aerophores absent at pinna bases; veins free, meeting the margins at the sinuses (ours), or anastomosing and forming a complex network; indument of acicular (unicellular or septate) hairs on rachises, costae, veins, and tissue between veins, glands absent; sori exindusiate, running along the veins; sporangia usually setiferous; spores finely echinate; x⫽36. Subgenus Stegnogramma comprises about 15 species, and all but two are confined to the Old World (mostly eastern Asia). Stegnogramma is distinguished by the often adnate and decurrent distal pinnae, elongate exindusiate sori than run along the veins, usually setiferous sporangia, and finely spinulose spores. Two species from southern Brazil, often considered to belong here, have affinities with subg. Goniopteris.

67. THELYPTERIS PILOSA Thelypteris pilosa (M. Martens & Galeotti) Crawford, Amer. Fern J. 41: 16, t. 3a. 1951. Figs. 312A–D, 313A. Gymnogramma pilosa M. Martens & Galeotti, Me´m. Foug. Mexique 27, pl. 4, f. 1. 1842. Dryopteris pilosa (M. Martens & Galeotti) C. Chr., Index Filic. 284. 1905. Stegnogramma pilosa (M. Martens & Galeotti) K. Iwats., Amer. Fern J. 54: 141. 1964. Syntypes. Mexico. Veracruz: Pic d’Orizaba, Galeotti 6267, 6268 (BR!; isosyntypes K!, P!). Gymnogramma procurrens Fe´e, Me´m. Foug. 8: 78. 1857. Gymnogramma pilosa M. Martens & Galeotti var. major E. Fourn., Mexic. Pl. 1: 73. 1872. Dryopteris pilosa (M. Martens & Galeotti) C. Chr. var. procurrens (Fe´e) C. Chr., Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 7, 10: 197. 1913. Thelypteris pilosa (M. Martens & Galeotti) Crawford var. major (E. Fourn.) Crawford, Amer. Fern J. 41: 19. 1951. Type. Mexico. Popocate´petl, Schaffner 293 (not found at P). Dryopteris pseudototta Christ, Bull. Herb. Boissier, se´r. 2, 7: 415. 1907. Type. Mexico. Chiapas: San Cristo´bal, Mu¨nch 23 (not found at P).

Rhizomes short- to long-creeping, ca. 2–5 mm diam.; fronds mostly less than 55 cm long; stipes 0.5–1.5 times the blade length; pinnae 1–8(–10) ⫻ 1.4(–2.5) cm, often broadly adnate and decurrent to rachises (especially in distal half of blades), or proxi-

Distribution. Margins of wet pine-oak woods and rocky slopes; (900–)1500–3350 m. Mexico; Guat, Hond. Selected Specimens Examined. BCS (Brandegee s.n., UC). Chih (LeSueur 1133, ARIZ, TEX, US). Chis (Breedlove 15108, DS). DF (Rzedowski 19496, NY). Dgo (Sa´nchez S. 714, US). Gro (Mexia 9002, ARIZ, CAS, GH, MICH, NY, U, UC, US). Gto (Dı´az B. 3723, ENCB, IEB). Hgo (Moore 2086, UC). Jal (Pringle 2589, ARIZ, CAS, GH, LL, MICH, MSC, NY, U, UC, US). Me´x (Mickel 703, ISC, MICH, NY, UC). Mich (Cruden 1069, UC). Mor (Pringle 11265, GH, UC, US). Nay (Breedlove 44623, CAS, MEXU). Oax (Smith 492, UC). Pue (Nicolas 5019, MICH, NY). Qro (Ferna´ndez N. & Zamudio 979, IEB, ENCB, MEXU). SLP (Schaffner s.n., NY). Son (Gentry 2112, ARIZ, GH). Tlax (Acosta P. & Sa´nchez 2356, CIB). Ver (C. Smith 2185, MO, UC). Unverified, Doubtful, or Mistaken Reports. Tam (reported by Lof, 1980, but not verified).

This species differs from other New World Thelypteris by the elongate, exindusiate sori, setose sporangia, and adnate/decurrent pinna bases toward the blade apices. It is not closely related to other Mexican species, but is allied to species from eastern Asia. Yoko et al. (2002) included samples of the American T. pilosa and T. burksiorum (see comments below) in their molecular analysis, and both species were found to cluster with the Asiatic T. pozoi (Lag.) C. V. Morton. Thelypteris pilosa var. alabamensis Crawford (Amer. Fern J. 41: 19. 1951) has been ascribed a range in northwestern Mexico, but the Mexican specimens seem to be merely small, depauperate collections of var. pilosa, and, in terms of frond size, there are many transitional specimens from throughout Mexico. In contrast, var. alabamensis, restricted to Alabama, differs in having pinnae stalked to 2 mm, the acroscopic segment of each proximal

668

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pinna Ⳳ free, and thin blade texture. The Alabama variant has recently been considered sufficiently distinctive to recognize it at species rank as Thelypteris burksiorum J. Watkins & Farrar (Watkins & Farrar, 2002).

THELYPTERIS SUBG. STEIROPTERIS Thelypteris subg. Steiropteris (C. Chr.) K. Iwats., Mem. Coll. Sci. Kyoto Univ., Ser. B, Biol. 31: 31. 1964. Dryopteris subg. Steiropteris C. Chr., Biol. Arb. Til. Eug. Warming 81. 1911. Steiropteris (C. Chr.) Pic. Serm., Webbia 28: 449. 1973. Type. Dryopteris deltoidea (Sw.) C. Chr. [⬅ Polypodium deltoideum Sw.] ⬅ Thelypteris deltoidea (Sw.) Proctor.

Blades pinnate-pinnatifid (ours) or pinnatifid, with or without (ours) buds absent in axils of distal pinnae; pinnae shallowly to deeply pinnatifid, the proximal ones Ⳳ the largest or nearly so, distal pinnae gradually reduced or blades sometimes each with a subconform terminal pinna; aerophores often strongly developed at the bases of pinnae; veins from adjacent segments running to the margins above the sinuses or the lowermost pair connivent at the sinuses (ours), with a raised false vein extending from the sinuses toward the costae, occasionally the veins united below the sinuses and with the false vein; indument of acicular hairs, these sometimes septate; sori rounded, indusiate or exindusiate; sporangia glabrous (ours); x⫽36. Subgenus Steiropteris comprises 21 neotropical species, from southern Mexico to Bolivia and southern Brazil and the Antilles. A single species is known from Mexico, and a second, Thelypteris decussata (L.) Proctor, might be expected in Chiapas, since it occurs in Guatemala. Thelypteris decussata differs from T. glandulosa in lacking a sinus keel, veins reaching the segment margins above the sinus, and presence of sessile, resinous glands on the laminae abaxially. Species of subg. Steiropteris are usually found at middle and low elevations, below 1500 m, especially in wet primary forests and along streams. Steiropteris is usually characterized by the raised keel (false vein) extending from the sinuses toward the costae, acicular hairs, and veins from adjacent segments running toward the sinuses. It seems close to subg. Goniopteris, but species lack stellate hairs.

68. THELYPTERIS GLANDULOSA VAR. BRACHYODUS Thelypteris glandulosa (Desv.) Proctor var. brachyodus (Kunze) A. R. Sm., Phytologia 34: 233. 1976. Figs. 314A–C. Polypodium brachyodus Kunze, Linnaea 9: 48. 1834. Dryopteris glandulosa (Desv.) C. Chr. var. brachyodus (Kunze) C. Chr., Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 7, 10: 172. 1913. Thelypteris brachyodus (Kunze) Ching, Bull. Fan Mem. Inst. Biol. 6: 286. 1936. Type. Peru. Pampayaco, Poeppig s.n. (LZ destroyed; isotype W!).

Rhizomes creeping; fronds monomorphic, (35–)65–150 cm; stipes (20–)35–75 cm ⫻ (2–)4–12 mm; blades (20–)30–75 cm, the apices hastate, pinnatifid, nearly free or free; pinnae (7–)10–25 (–30) ⫻ (1.8–)2.5–4.5(–6) cm, (4–)7–16 pairs, generally stalked 1–4 mm, incised ca. 1⁄3–1⁄2 the way to the costae, with scale-like aerophores 1–22 mm at pinna bases; segments 5–9 mm wide; veins (7–)14–22 pairs per segment, the proximal pair from adjacent segments uniting or connivent below the sinuses or 3–5 pairs running alongside a keel to a sinus; indument abaxially of dense hairs 0.1–0.6(–1) mm on rachises and costae, also of linear adpressed scales along costae; sori medial, exindusiate.

Distribution. Terrestrial in lowland rain forests, especially along streams; 300 m. Mexico; Guat, Bel, Hond, Nic, CR; Col, Ec, Peru. Specimens Examined. Chis (Breedlove 34189, DS, MEXU).

This differs from congeners in Mexico by the scale-like aerophores at the pinna bases, the venation (several pairs of veins connivent below the sinuses), and the hastate blade apices. Two other varieties occur in other parts of the range: var. glandulosa, in the Lesser Antilles, Colombia, Venezuela, Trinidad, Guyana, Surinam, and French Guiana; and var. longipilosa A. R. Sm., from Costa Rica, Panama, Colombia, and Ecuador (Smith, 1980). The former is based on Polypodium glandulosum Desv., Ges. Naturf. Freunde Berlin Mag. Neusten Entdeck. Gesammten Naturk. 5: 317 (1811) [Syns.: Dryoperis glandulosa (Desv.) C. Chr., Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 7, 10: 171. 1913, hom. illeg., non (Blume) Kuntze, 1891. Thelypteris glandulosa (Desv.) Proctor, Rhodora 61: 306. 1959 (1960). Steiropteris glandulosa (Desv.) Pic. Serm., Webbia 28: 451. 1973. Type. Antilles. locality and collector not stated (P; frag. BM!)].

THELYPTERIS SUBG. THELYPTERIS Rhizomes long-creeping, slender; blades pinnate-pinnatifid, the apices gradually reduced, the base not reduced, lacking buds

TRICHOMANES

669

in axils of pinnae; aerophores absent; veins running to margins above the sinuses; indument of acicular hairs, blades also usually with sessile or stipitate glands; sori round, indusiate; paraphyses absent; x⫽35. Subgenus Thelypteris comprises 2 species, one restricted to the Northern Hemisphere, and the other, T. confluens (Thunb.) C. V. Morton, mainly in the Southern Hemisphere in Argentina, southern Africa, Madagascar, southern India, southeast Asia, Malesia, Australia, and New Zealand. The two species can be recognized by their long-creeping, slender rhizomes, nondecrescent blade bases, scattered, ovate scales on the costae abaxially, and furcate veins. The base chromosome number of 35 is also unique among New World thelypteroid ferns. Species in subg. Thelypteris occur in swamps or marshy areas. The sole Mexican species is strongly disjunct from the nearest populations in the southeastern United States.

69. THELYPTERIS PALUSTRIS VAR. PUBESCENS Thelypteris palustris Schott var. pubescens (G. Lawson) Fernald, Rhodora 31: 34. 1929. Figs. 315A–C. Lastrea thelypteris (L.) Bory var. pubescens G. Lawson, Edinburgh New Philos. J. n.s. 19: 277. 1864. Type. Canada. Ontario: Odessa, Addington Co., Lawson s.n. (presumably E). Dryopteris tremula Christ, Notul. Syst. 1: 234. 1910. Type. Mexico. Michoaca´n: La Huerta, Arse`ne 3106 (P?; isotypes GH!, MEXU–3 sheets!, NY!, U!, US!).

Rhizomes long-creeping, 1–3 mm diam.; fronds monomorphic or slightly dimorphic, often 1–3 cm apart, fertile fronds more erect, narrower and with somewhat contracted pinnae and segments, 20–90 cm long; stipes stramineous above bases, 9–45 (–60) cm ⫻ 1–3 mm, at bases sparsely set with tan, ovate, glabrous scales; blades lanceolate, 10–40(–55) cm long, proximal pinnae commonly slightly shortened, blades tapering gradually to pinnatifid apices; pinnae pinnatifid to within 1 mm of costae, 2–10 ⫻ 0.5–2 cm; segments oblong, somewhat oblique, entire; veins from adjacent segments meeting margins above sinuses, frequently forked; indument abaxially of sparsely to densely set hairs on costae and sometimes veins, costae also commonly with a few small, tan scales, tissue between veins glabrous on both sides; sori medial, with tan, often hairy indusia; sporangia glabrous; 2n⫽70 (USA).

Distribution. Along slow-moving streams and waterways, swampy areas; 1800–1850 m. E Canada, e USA; Mexico; Berm, Cuba; the distribution of the species (var. palustris) extends to Europe, Asia. Specimens Examined. DF (Schaffner s.n., MEXU). Mich (Arse`ne 2562, 5907, s.n., US; Rzedowski 39234, IEB, XAL). State unknown (Schaffner 57, NY).

Distinctive characters are the very long-creeping, narrow rhizomes, the free, frequently forked veins, and the tan scales along the costae abaxially. The sole variety recognized by Tryon et al. (1980) in temperate North America is var. pubescens and, from the meager material available, the Mexican specimens seem to be the same variety. Eurasian material is referable to var. palustris, based on Acrostichum thelypteris L., Sp. Pl. 2: 1071. 1753. [Syn.: Theylypteris palustris Schott, Gen. Fil., t. 10. 1834. Dryopteris thelypteris (L.) A. Gray, Man., ed. 1, 630. 1848. Type. Uncertain, based on plants from Europe and citations to publications by Bauhin (with illustration) and Ruppius.] Dryopteris tremula has been a puzzle since Christ first described it in 1910. It now seems likely that it is merely an outlier of the widespread north-temperate species, T. palustris, differing in no significant characters from that species. The Schaffner collection was made in 1875, from Xochimilco. The only recent collection since the early 1900’s is by Rzedowski, in 1985, from the mouth of Lago de Cuitzeo.

1 2 1 . T R I C H O M A N ES Trichomanes L., Sp. Pl. 2: 1097. 1753. Name and type conserved. Type: Trichomanes crispum L. Didymoglossum Desv., Me´m. Soc. Linn. Paris 6: 330. 1827. Lectotype (chosen by C. Chr., Index Filic. XIV. 1906): Didymoglossum muscoides (Sw.) Desv. [⬅ Trichomanes muscoides Sw.] ⫽ Trichomanes hymenoides Hedw. For additional synonymy, see Morton, 1968.

Mostly epiphytic, terrestrial (few in Mexico) or epipetric; rhizomes usually long-creeping, erect in some species, very slender, with stiff hairs; roots usually present, absent in a few species; fronds minute to small, occasionally medium-sized, generally monomorphic, rarely dimorphic, distant; stipes slender, glabrous or hairy; blades undivided to decompound, membranaceous,

670

TRICHOMANES

mostly only one cell thick, glabrous or with simple to stellate hairs (on veins and blade margins); segments mostly adnate and decurrent to next axis; sori marginal, in conical or tubular involucres, each with straight or flared, entire to bilobed mouth, immersed in laminae or exserted; receptacles growing from base, with age generally resulting in a long-exserted trichome; spores tetrahedral-globose, green, lacking conspicuous markings; gametophytes filamentous, often gemmiferous; x⫽32, 33, 34, 36. Trichomanes is a pantropical genus of over 300 species, of which about 100 are American, with 26 in Mexico. The genus was divided by Copeland (1938) into many genera, but splitting seems to compound the problem of generic delimitation in the filmy ferns, rather than alleviate it.

The genus is distinct by its mostly one-cell-thick blades, tubular or conical involucres, and strongly exserted receptacles. References Copeland, E. B. 1938. Genera Hymenophyllacearum. Philipp. J. Sci. 67: 1–110; Dubuisson, J.-Y., S. Hennequin, E. J. P. Douzery, R. B. Cranfill, A. R. Smith and K. M. Pryer. 2003. rbcL phylogeny of the fern genus Trichomanes (Hymenophyllaceae, with special reference to neotropical taxa. Int. J. Pl. Sci. 164: 753– 761; Morton, C. V. 1968. The genera, subgenera, and sections of the Hymenophyllaceae. Contr. U.S. Natl. Herb. 38: 153–214; Pacheco, L. 1994. Pterido´fitas: Familia Hymenophyllaceae. Flora de Me´xico 6(2): 1–56, Consejo Nacional de la Flora de Me´xico, Me´xico, D.F.; Pryer, K. M., A. R. Smith, J. S. Hunt, and J.-Y. Dubuisson. 2001. rbcL data reveal two monophyletic groups of filmy ferns (Filicopsida: Hymenophyllaceae). Amer. J. Bot. 88: 1118–1130; Wessels Boer, J. G. 1962. The New World species of Trichomanes sect. Didymoglossum and Microgonium. Acta Bot. Neerl. 11: 277–330.

Key to the Mexican Species of Trichomanes 1. Blades bipinnatifid to quadripinnate, never simple or merely lobed; segments without marginal hairs; false veins absent. 2. Rhizomes erect or ascending, 5–7 mm diam., the fronds somewhat clumped; stipes non-alate; plants terrestrial. ..................... 25. T. rigidum. 2. Rhizomes long-creeping, 0.1–2 mm diam., the fronds distant; stipes alate to non-alate; plants epiphytic or epipetric (occasionally terrestrial in T. pyxidiferum and T. hymenophylloides). 3. Fronds 16–36 cm long; rhizomes 1 mm or more diam., with dense hairs; apparently lacking true roots on climbing stems. 4. Involucres with erect lips, slightly flaring or straight, 0.5–0.8 mm wide; fronds with stipes 15–55 mm long; fronds broad, ca. 3 times as long as wide, 22–30 ⫻ 6–10 cm; (600–)1250–2450 m. ...................................................................... 23. T. radicans. 4. Involucre lips flaring out to form a collar 1 mm wide, lips 0.5 mm broad; fronds sessile or stipes 4–12 mm long, (rarely to 40 mm); fronds narrow, ca. 5 times as long as wide, 16–36 ⫻ 3–6 cm, narrowed at bases; 200–850 m. .................5. T. collariatum. 3. Fronds 15 cm or less long; rhizomes 0.5–0.8 mm diam. or less, creeping but not robust, lacking dense hairs; roots present. 5. Stipes non-alate, 0.1–0.5 mm diam.; blades 3–4-pinnate, segments not coalescing at bases, glabrous; rhizomes 0.1–0.3 mm diam., with sparse brown hairs 0.3 mm long; often epiphytic on tree ferns. 6. Ultimate segments filiform, mostly 0.1–0.4 mm wide. ................................................................................................ 4. T. capillaceum. 6. Ultimate segments linear, mostly 0.5–0.7 mm wide. .................................................................................................... 1. T. angustatum. 5. Stipes alate 1⁄2 or entire length, 0.5–0.8 mm diam.; blades bipinnatifid to tripinnate; segments with 2-celled clavate hairs on laminae, veins, and bases of involucres; rhizomes 0.5–0.8 mm diam. with dense hairs 0.5 mm long, epiphytic or terrestrial, not usually on tree ferns. 7. Rhizomes with black hairs; stipes blackish with dense black hairs at bases; ultimate segments 0.8–1 mm wide, with long narrow folds giving the appearance of “false veins” parallel to blade margins. .............................................. 22. T. pyxidiferum. 7. Rhizomes with spreading thin reddish brown hairs 1 mm long and tightly appressed, light brown, hairs 0.5 mm long; stipes green for the most part, with sparse light brown hairs at bases; ultimate segments 0.3–0.5 mm wide, plane. .......................................................................................................................................................................... 14. T. hymenophylloides. 1. Blades simple to not more than pinnatifid or pinnate-pinnatifid, sometimes simple or merely lobed, if bipinnatifid, then with false veins or densely hairy; segments with or without marginal hairs; false veins present or absent. 8. Fronds dimorphic, sterile fronds pinnatifid, 3–5 cm wide, with anastomosing true veins; false veins absent; rhizomes erect, 4–5 mm diam.; sterile blades often radicant at apices. ............................................................................................................................... 8. T. diversifrons. 8. Fronds monomorphic; false veins present or absent; rhizomes erect or suberect, 2–12 mm diam., or long-creeping, 0.1–1 mm diam.; sterile blades never radicant at apices. 9. Blades tightly appressed to tree substrate by hairs along rachises, veins, and margins; involucres strongly exserted; plants epiphytic; rhizomes long-creeping; .............................................................................................................................................. 26. T. tuerckheimii. 9. If epiphytic, blades not adhering to substrate; involucres exserted or immersed; plants epiphytic, epipetric, or terrestrial; rhizomes long-creeping to erect. 10. Fronds 4–60 cm long, pinnatifid, 1-pinnate, or 2-pinnatifid, never simple; false veins absent or if present, then perpendicular to veins, not parallel to veins or blade margins; plants terrestrial or epiphytic; rhizomes erect or suberect, 2–12 mm diam. (or long-creeping, 0.3–0.5 mm diam. in T. polypodioides). 11. False veins present, connecting true veins at right angles; veins and blade margins glabrous; involucres exserted, on stalks. .................................................................................................................................................................................... 20. T. pinnatum. 11. False veins absent; veins and sometimes blade margins with hairs; involucres immersed in blade tissue. 12. Marginal hairs forked or stellate, sessile; hairs on veins forked or stellate, also with scattered, short, black, clavate hairs 0.1 mm; rhizomes 0.3–0.5 mm diam., long-creeping; blades pinnatifid. .................................. 21. T. polypodioides.

TRICHOMANES ANGUSTATUM

671

12. Marginal hairs simple or absent; hairs on veins simple, 1–3 mm long, jointed; rhizomes 3–12 mm diam., erect or suberect, rarely short-creeping; blades pinnatifid to bipinnatifid. 13. Fronds pinnatifid; stipes alate in distal 1⁄2–3⁄4; stipes alate. 14. Stipes alate only in distal 1⁄2(–3⁄4); blades attenuate at apices. ................................................................... 10. T. galeottii. 14. Stipes alate to bases; blades rounded at apices. ...................................................................................... 12. T. holopterum. 13. Fronds pinnate to bipinnatifid, proximal pinnae stalked; stipes non-alate. 15. Blades bipinnatifid; marginal hairs absent; hairs on stipes, rachises, and veins 1–3 mm long, reddish brown, dense. ...................................................................................................................................................... 16. T. lucens. 15. Blades pinnate proximally; marginal hairs simple, 1–2 mm; hairs on stipes, rachises, and veins 1–2 mm long, dark brown, sparse. ....................................................................................................................... 6. T. crispum. 10. Fronds 1–8 cm long, simple to bipinnatifid but not truly pinnate; false veins running parallel to, and sometimes perpendicular between, the true veins, sometimes also in a submarginal position; plants epiphytic or epipetric; rhizomes long-creeping, 0.1–0.8 mm diam. 16. Blade margins glabrous; a submarginal row of cells slightly different from the others present (submarginal false vein), also with false veins parallel to the true veins; involucre lips not dark-edged; blades round to ovate. 17. Occasional false veins perpendicular to veins and false veins forming netted venation; blades round to ovate, undivided; fronds 1–2.5 cm long. ............................................................................................................................... 11. T. godmanii. 17. Veins free, false veins present but parallel to the true veins and mostly unconnected; juvenile blades more or less round; mature blades oblong or elongate, the upper part irregularly lobed. ................................................... 9. T. ekmanii. 16. Blade margins with dark, forked or stellate hairs or orbicular marginal scales 1–1.5 mm diam.; submarginal and perpendicular false veins absent, false veins running parallel between true veins; involucres with two distinct, darkedged lips (or non-bilabiate and concolorous in T. petersii); blades ovate, simple to bipinnatifid; fronds 1–8 cm long. 18. Blades with gray-brown, orbicular marginal scales 1–1.5 mm diam. ........................................................... 17. T. membranaceum. 18. Blades with dark, 2–4-rayed marginal hairs. 19. Involucres without dark-edged lips; blades simple; growing on walls of calcareous sinkholes. .................... 19. T. petersii. 19. Involucres with dark-edged lips; blades simple, pinnatifid, or bipinnatifid; mostly epiphytic, if growing on wet rocks then blades pinnatifid, not simple. 20. Blades linear, 1–3 mm wide; fronds 5–15 cm long. ................................................................................. 2. T. angustifrons. 20. Blades oblong to ovate or obovate, 2.5–35 mm wide; fronds 2–35 cm long. 21. Blades mostly entire with rounded apices, obovate or flabellately lobed, 1 cm or less long; epiphytic. 22. Sori one per frond; fronds 2–8 mm long. ......................................................................................... 18. T. ovale. 22. Sori (3–)6–13 per frond; fronds 16–30 mm long. .............................................................................. 7. T. curtii. 21. Blades pinnatifid to bipinnatifid, 1.5–8 cm long; epiphytic or on wet rocks. 23. Marginal hairs simple, forked, or stellate; main veins appearing almost palmate; fronds (0.9–)1.5–3.5 cm long, blades broadest at base, with deep sinuses and overlapping segments. ........................................................................................................................................................ 13. T. hymenoides.

23. Multi-rayed hairs present along margins and/or at bases of sinuses; main veins distantly pinnate; fronds 1.5–8 cm long; blades lanceolate to broad, narrowed proximally, with shallow to deep sinuses, segments often of irregular lengths, rarely overlapping. 24. Blades linear with shallow lobes to mixture of short and long segments, the longer segments mostly near blade apices; involucres with stalks 0.5–1 mm long, with wide lips 1 mm long; blade margins with both forked and stellate hairs, many near pinna tips. .................................................................................................................................................. 3. T. bucinatum.

24. Blades with both short and long segments; involucres somewhat exserted (only to base of involucre) to immersed, with lips 0.5 mm long; blade margins with sparse forked hairs and stellate hairs restricted to based of sinuses. 25. Involucres immersed (at least alate to lips) to exserted; false veins absent between true veins, somewhat parallel to and near blade margins; 0–500(–1070) m elevation. ................................................................................................................................................ 15. T. krausii.

25. Involucres exserted to slightly immersed; false veins often present between true veins, also with some parallel to blade margins; 900–2500 m. .................................... 24. T. reptans.

1. TRICHOMANES ANGUSTATUM Trichomanes angustatum Carmich., Trans. Linn. Soc. London 12: 513. 1818. Figs. 317K–M. Vandenboschia angustata (Carmich.) Copel., Philipp. J. Sci. 73: 466. 1941. Type. Tristan da Cunha, Carmichael s.n. (K!; isotype BM!).

Trichomanes tenerum Spreng., Syst. Veg., ed. 16, 4: 129. 1827. Vandenboschia tenera (Spreng.) Copel., Philipp. J. Sci. 67: 53. 1938. Type. Brazil. s. coll. s.n. (LZ destroyed).

Closely related to T. capillaceum, differing mainly in the slightly broader ultimate segments ca. 0.5–0.7 mm wide (vs. 0.1– 0.4 mm); 2n⫽144 (Tristan da Cunha).

672

TRICHOMANES

Distribution. Usually epiphytic on tree ferns in wet forests; widespread in the American tropics, but exact distribution uncertain because of problems in circumscription and confusion with T. capillaceum; elevation not known for Mexican collections; 250–2000 m in Greater Antilles, 2600 m in CR. Mexico; Guat, Hond; Gr Ant; Col, s Braz, Bol, Parag, Uru, nw Arg; Tristan da Cunha.

Distribution. Epiphytic on tree trunks, in montane rain forests; 1300–1850 m. Mexico; Guat, CR; Gr & L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol, Parag. Specimens Examined. Chis (Breedlove 32179, 35257, DS).

The cited specimens are difficult to determine. The involucral lips are not dark, but these may be juvenile specimens.

Specimens Examined. Chis (Breedlove 68356, CAS; Mu¨nch 108, DS p.p.). Gro (Hinton et al. 14307, ARIZ). Ver (Correll & Correll 28768, MICH, TEX; Purpus 2411, UC).

3. TRICHOMANES BUCINATUM

The T. angustatum-capillaceum complex needs monographic attention before names can be applied with confidence. The few specimens cited here seem to have consistently wider segments and are here recognized as distinct from T. capillaceum.

Trichomanes bucinatum Mickel & Beitel, Mem. New York Bot. Gard. 46: 390. 1988. Type. Mexico. Oaxaca: Distrito Ixtla´n, trail from San Pedro Nolasco N to the Llano Verde, Mickel 4394 (NY!). Figs. 322F–H.

2. TRICHOMANES ANGUSTIFRONS Trichomanes angustifrons (Fe´e) Wess. Boer in Kramer, Fl. Neth. Antill. 1: 17. 1962. Figs. 318A, B. Didymoglossum angustifrons Fe´e, Me´m. Foug. 11: 113, t. 28, f. 5. 1866. Type. Guadeloupe, L’Herminier s.n. (isotype P!).

Rhizomes slender, long-creeping, ca. 0.2 mm diam.; rhizome hairs dark brown, 0.2 mm long; roots absent; fronds 5–15 ⫻ 1–3 mm, distant; stipes very short, 0.5–1 mm, 1⁄10–1⁄15 the frond length; blades linear or linear-oblong, mostly entire, irregularly pinnatifid distally; veins free, ascending, with false veins parallel but not connected to true veins, both types running to margins; margins with simple to stellate hairs; sori single on apical lobes; involucres totally immersed, lips spreading, semiorbicular, margin with 1–3 rows of black-walled cells; 2n⫽68 (Trin).

Rhizomes slender, 0.3 mm diam., long-creeping; rhizome hairs dense, dark brown to black, 0.5 mm long; roots absent; fronds 1.5–4 cm long, distant; stipes ca. 1⁄10–1⁄4 the frond length, 0.5 mm wide, flattened, alate to base or nearly so, with dark brown to black hairs 0.3–0.5 mm long; blades irregularly pinnatifid, some linear, 0.5–2 cm wide, with short lobes, others with pinnae of differing lengths, broader towards the blade apices, to 2 cm wide, margins entire with both forked and stellate hairs, many near pinna tips; rachises broadly adnate; veins free, many unconnected false veins parallel to true veins, with scattered short, clavate hairs 0.1 mm long; sori 1-several per blade; involucres strongly exserted, stalk 0.5–1 mm long, 3 ⫻ 0.8 mm, each with large, bilabiate mouth, lips 1 ⫻ 1 mm wide, with dark margins of several rows of cells.

TRICHOMANES COLLARIATUM

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0.5 mm wide at middle, 1.5 mm long, mouth flared, 0.8–1 mm wide, entire; 2n⫽72 (CR, Jam).

Distribution. Epiphytic in wet montane forests; 1500–2200 m. Mexico. Specimens Examined. Hgo (Gimate L. 971, NY). Oax (Mickel 982, ISC, 5105, 5387, NY). Unverified, Doubtful, or Mistaken Reports. Ver (Pringle 10809, XAL, cited by Palacios-Rios, 1992, but specimen is T. reptans).

The variable pinnatifid fronds (some narrowly lanceolate to linear with reduced proximal pinnae), some stellate hairs at the pinna tips, and the strongly exserted involucres with long lips separate this species from other members of sect. Didymoglossum (T. petersii, T. ovale, T. hymenoides, and T. reptans). It resembles T. reptans most closely.

4. TRICHOMANES CAPILLACEUM Trichomanes capillaceum L., Sp. Pl. 2: 1099. 1753. Figs. 321J, K. Vandenboschia capillacea (L.) Copel., Philipp. J. Sci. 67: 53. 1938. Lectotype (chosen by Proctor, 1985: 109.). Plumier, Traite´ Foug. Ame´r., pl. 99D, 1705, based on a plant collected in Haiti. Trichomanes tenellum Hedw., Fil. Gen. Sp., ad t. 6, text. 1799. Trichomanes trichoideum Sw., J. Bot. (Schrader) 1800(2): 98. 1801, nom. illeg. (see Morton 1973, Contr. U.S. Natl. Herb. 38: 274). Trichomanes trichodes Sw., Fl. Ind. Occid. 3: 1741. 1806 (renaming of T. trichoideum Sw.). Type. Hedwig’s figure. Trichomanes schiedeanum Mu¨ll. Berol., Bot. Zeit. (Berlin) 12: 716. 1854. Type. Mexico, Schiede s.n. (B?). Trichomanes hypnoides Christ, Bull. Herb. Boissier, se´r. 2, 5: 725. 1905. Type. Mexico. Chiapas: Zontehuitz, Mu¨nch 108 (P?, not found; isotypes DS!, US!).

Rhizomes very fine, 0.1–0.3 mm diam., long-creeping, filamentous; rhizome hairs sparse to abundant, brown, 0.3 mm long; roots present; fronds 5–10(–15) cm long, distant, glabrous; stipes ca. 1⁄4 the frond length, 0.1–0.5 mm diam., glabrous, non-alate, green to stramineous (darker at base); blades lanceolate, ovate to oblanceolate, light yellow-green, membranaceous, 3–4-pinnate, 1–5 cm wide, proximal pinna pairs slightly reduced; rachises nonalate; pinnae non-alate in proximal third, ultimate segments filiform, 0.1–0.4 mm wide, emarginate, margins entire, plane; veins free, false veins absent; sori often 1(–4) per pinna, terminating proximal veinlets; involucres stalked and free from the tissue, 0.3–

Distribution. On tree fern trunks or rarely on wet rocks by streams, wet montane forests at middle elevations; 900–2600 m. Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Gr Ant; Col, Ven, Ec, Peru. Selected Specimens Examined. Chis (Breedlove 23142, MICH, 33027, DS, MICH). Gro (Hinton 14307, NY, US; Rzedowski 18611, MICH). Me´x (Tejero-Dı´ez 2899, IZTA). Mor (Lyonnet 2860, CHAPA, ENCB, MEXU, US). Oax (Mickel 1024, MICH, NY, 6585, NY). Pue (Rzedowski 29972, NY). Tab (Cowan 2902, CAS). Ver (Pringle 8173, CAS, P, UC, US; Purpus 2411, UC).

The very finely dissected blades, filiform segments, and nonalate stipes distinguish T. capillaceum. Specimens from Oaxaca and farther north in Mexico are typical of the species, whereas in Chiapas intermediates between T. capillaceum and T. angustatum were reported by Smith (1981). Trichomanes angustatum has broader ultimate segments 0.5–0.7 mm wide and costae alate to the main rachis.

5. TRICHOMANES COLLARIATUM Trichomanes collariatum Bosch, Ned. Kruidk. Arch. 4: 368. 1858 [1859]. Type. Mexico. Tabasco: Linden 1508 (isotypes GH!, K!). Figs. 316D–F. Trichomanes martinezii Rovirosa, Pteridogr. Sur Me´xico 106. 1909. Vandenboschia martinezii (Rovirosa) Pic. Serm., Webbia 28: 472. 1973. Type. Mexico. Chiapas: Laguna de Lacanja, Martı´nez s.n. (herb. no. 1103, isotype GH).

Rhizomes stout, 1–1.5 mm diam., long-creeping; rhizome hairs thin, brown, 1 mm long, and also with scattered, thick, dark brown hairs 1–1.5 mm long; roots absent on climbing stems; fronds 16–36 cm long, distant; stipes very short, absent to 4–12 (–40) mm long, light to dark brown, alate to bases; blades linearoblong, very dark green, drying nearly black, membranaceous, 2-pinnate to 2-pinnate-pinnatifid, 3–6 cm wide, reduced proximally; rachises alate; pinnae glabrous or with occasional short hairs 0.1 mm long; veins free, false veins absent; sori one per pinnule or segment; involucres nearly free from blade tissue, elon-

674

TRICHOMANES

gate, 1.5–2 mm long, 0.5 mm at middle, 0.3 mm at base, mouth strongly flaring, lips 0.5 mm wide; 2n⫽72 (Oax).

Distribution. Epiphytic in wet montane forests of Atlantic slope; 200–850 m. Mexico; Guat, Bel, Hond, Nic, CR, Pan; Col, Ven, Ec, Peru, n Braz, Bol. Selected Specimens Examined. Chis (Breedlove 34073, 57296, MICH, NY; Martı´nez S. 12071, NY). Oax (Hallberg 1300, NY; Herna´ndez G. 1230, NY; Mickel 1558, MICH, NY, 7157, NY). Tab (Cowan 2902, NY). Ver (Ventura A. 8117, NY; Wendt et al. 2688, NY).

This species is commonly confused with T. radicans (q.v.) but can be distinguished most readily by the broadly flaring indusial mouths, tapering blade bases with short stipes, and occurrence at lower elevations, 200–850 m. (vs. (600–)1250–2450 m).

6. TRICHOMANES CRISPUM Trichomanes crispum L., Sp. Pl. 2: 1097. 1753. Lectotype (chosen by Proctor 1977: 95). Plumier, Traite´ Foug. Ame´r., pl. 86, 1705, illustrating a plant from Martinique. Figs. 320G, H. Rhizomes stout, 6–12 mm diam., short-creeping or suberect; rhizome hairs dark brown, 1–2 mm long; roots present; fronds 10– 40 cm long, clumped; stipes ca. 1⁄4 the frond length, 0.8–1 mm diam., brown, non-alate, covered with hairs 1–3 mm long; blades linear to narrowly ovate, membranaceous, pinnate, not reduced at bases, 2–5 cm wide, apices pinnatifid; rachises alate only in distal part; pinnae obtuse at apices, proximal pinnae with rounded bases, margins entire, undulate to crisped; hairs simple, 1–2 mm long, on veins and along blade margins; veins free, forked, false veins absent; involucres immersed in blade tissue, 1.5–2 mm long, 1–1.2 mm wide distally, mouths flaring but sunken, not bilabiate; 2n⫽256 (Jam, Trin).

Distribution. Terrestrial or epiphytic in wet montane forests, along Atlantic slope at middle elevations; 180–1350 m. Mexico; Guat, Bel, Hond, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Braz. Specimens Examined. Chis (Breedlove 42066, DS; Martı´nez S. 14935, XAL). Oax (Mickel 1036, ISC, MICH, US, 5725, NY, 6687, ENCB). Ver (Ramı´rez 1896, XAL). Unverified, Doubtful, or Mistaken Reports. Tab (Becerra s.n., MEXU, cited by Pacheco, 1994, but not verified).

Trichomanes crispum is similar to T. galeottii (q.v.). In T. crispum, stipes are non-alate, the proximal pinnae are stalked, and the rachises are non-alate for at least the proximal 1⁄2 of their length.

7. TRICHOMANES CURTII Trichomanes curtii Rosenst., Repert. Spec. Nov. Regni Veg. 22: 5. 1925. Type. Costa Rica. Litorale Atlanticum, Finca Gebr. Hundriesser, Brade & Brade 398a (S). Figs. 318E, F. Similar to T. ovale except: fronds 16–30 mm long (vs. 2–8 mm), sori (3–)6–13 per frond (vs. only 1).

Distribution. Epiphyte in lowland wet forests; 100 m. Mexico; Guat, Bel, Nic, CR, Pan; Col. Specimen Examined. Chis (Pacheco s.n., UAMIZ).

TRICHOMANES GALEOTTII

Trichomanes curtii also resembles T. petersii, which differs in the oblong, pinnately lobed blades with more distant veins. Trichomanes ovale is entire to sub-palmately lobed, with blades and mostly orbicular.

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Trichomanes diversifrons is distinct in its strongly dimorphic fronds with undivided fertile blades, pinnatifid sterile blades, anastomosing veins, and concave involucral mouths.

9. TRICHOMANES EKMANII 8. TRICHOMANES DIVERSIFRONS Trichomanes diversifrons (Bory) Mett. ex Sadeb. in Engler & Prantl, Nat. Pflanzenfam. 1(4): 108. 1899. Figs. 317A, B. Trichomanes elegans Rudge, Pl. Guian. 24, t. 35. 1805, hom. illeg., non Rich., 1792. Type. French Guiana. Martin s.n. (ex Herb. Rudge) (BM; isotype NY!). Hymenostachis diversifrons Bory, Dict. Class. Hist. Nat. 8: 462. 1825. Feea diversifrons (Bory) Copel., Philipp. J. Sci. 67: 74. 1938. Type. “Guiane.” Poiteau s.n. (P?).

Rhizomes stout, 4–5 mm diam., erect; rhizome hairs dark brown, 1–2 mm long; roots present; fronds strongly dimorphic, sterile fronds 10–35 cm long, fertile fronds 30–35 cm long, clumped; stipes 1⁄6–1⁄3 the sterile frond length, 1⁄2 the fertile frond length, stout, 1–1.3 mm wide, brown, alate in distal 1⁄4, bases with scattered dark brown hairs 1–2 mm long, glabrous otherwise; sterile blades pinnatifid, 3–5 cm wide, reduced proximally, apices pinnatifid, often with naked, extended rachises ending in proliferous tips; pinnae 12–36 pairs, 5–6 mm wide, glabrous except for rare, brown hairs 0.3 mm long, on rachises, margins serrate with teeth 0.3–0.5 mm; veins 2–3-forked, anastomosing, ending in teeth at margins, false veins absent; fertile fronds 5–9 mm wide, glabrous, veins simple to 1-forked, margins appearing denticulate; involucres immersed, contiguous, 1.5–2 mm long, 0.5 mm wide in middle, mouths 0.8 mm wide, concave.

Trichomanes ekmanii Wess. Boer, Acta Bot. Neerl. 11: 319. 1962. Type. Dominican Republic. Sto. Domingo: Cordillera Central, La Cumbre, Ekman H-14342 (U; isotypes F, G, GH!, MO, S, UC!, US!). Figs. 318G, H. Similar to T. godmanii in frond form and distinct submarginal row of cells (false vein), but differs in lacking transverse false veins; small blades nearly round, but mature blades oblong or elongate, the distal part irregularly lobed.

Distribution. Epiphytic in rain forests; 0–1000 m. Mexico; Bel, Hond, CR, Pan; Cuba, Hisp; Col, Ven, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol. Specimens Examined. Chis (Espejo 1440, CHAPA, ENCB, MEXU, UAMIZ; Martı´nez 25596, MEXU). Oax (Mickel 5849, ENCB).

Mickel 5848 (NY) (⫽ T. godmanii) has some transverse false veins, but 5849 does not. The coincidence of the two species cooccurring suggests that the character of transverse false veins may be a weak one, at least in Mexico. However, we treat them here as distinct, pending further study.

10. TRICHOMANES GALEOTTII

Distribution. Terrestrial in wet forests at low elevations; 150– 350 m. Mexico; Guat, Bel, Hond, Nic, CR, Pan; Col, Ven, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol. Selected Specimens Examined. Chis (Breedlove 34109, DS; Ghiesbreght 442, K, NY, PH; Martı´nez S. 15901, MEXU). Oax (Croat & Hannon 63268, UC; Herna´ndez G. 822, 1177, NY). Tab (Cowan 3990, NY, UC). Ver (Lorence et al. 3940, MEXU; Riba 804, UAMIZ; Wendt et al. 3845A, UC, 5305, NY).

Trichomanes galeottii E. Fourn., Bull. Soc. Bot. France 15: 147. 1868. Lectotype (chosen by Smith, 1981: 242). Mexico. [Oaxaca:] Galeotti 6530 (P!–3 sheets, further lectotypification needed; isolectotypes BR, LE!, photos BM!, GH!). Figs. 317E, F, N, O. Rhizomes stout, 3–6 mm diam., ascending or suberect; rhizome hairs dark brown, 1 mm long; roots present; fronds 5–25 cm long, clumped; stipes ca. 1⁄3 the frond length, green to dark green, alate 1⁄2(–3⁄4) their length, with hairs 1 mm long; blades linear to narrowly ovate, pinnatifid, membranaceous, 1.5–4 cm wide, proximal pinnae slightly reduced, adnate, apices attenuate, pin-

676

TRICHOMANES

natifid; rachises alate; pinnae oblong, apices obtuse, margins entire to slightly undulate, hairs simple, jointed, 1–2 mm long on veins and along margins; veins free, 1–2-forked, false veins absent; involucres immersed in blade tissue, 1.5–2 mm long, mouths slightly flaring, 1 mm wide, bilabiate.

Distribution. Epiphytic in wet montane forests on Atlantic slope; 120–600 m. Mexico; Guat, Bel, Hond, Nic, CR, Pan; Cuba.

Distribution. Epiphytic to epipetric in wet montane forests on Atlantic slope; 300–1500 m. Mexico; Guat, Bel, Hond, Nic, CR, Pan; Cuba, PR; Col, Ec. Selected Specimens Examined. Chis (Breedlove 21893, MICH, 57684, CAS; Martı´nez S. 16165, IBUG, 18892, MEXU). Oax (Herna´ndez G. & Gonza´lez L. 1665, NY; Liebmann s.n., P; Mexia 9282, CAS, GH, MICH, NY, UC, US; Mickel 7391, NY). Ver (Nee et al. 25035, NY; Ventura A. 17041, UAMIZ).

See Trichomanes crispum and T. holopterum for comments. In T. galeottii the proximal pinnae are not stalked, the rachises are alate throughout, and the stipes are alate in the upper 1⁄2 to 3⁄4.

11. TRICHOMANES GODMANII Trichomanes godmanii Hook. in Baker, J. Linn. Soc., Bot. 9: 337, t. 8, f. A. 1866. Lectotype (chosen by Wessels Boer, 1962: 319). Guatemala. Salvin & Godman s.n. (K!, frag. US!). Figs. 322C–E.

Specimens Examined. Chis (Martı´nez S. 10095, MEXU, 11101, UAMIZ). Oax (Herna´ndez X. 631, LL; Mickel 5848, NY). Tab (Gilly & Herna´ndez S. 294, US; Sa´nchez S. 284, US). Ver (Va´zquez Torres V-2516, CHAPA, ENCB, NY).

The perpendicular false veins distinguish this taxon from T. ekmanii, the only other Mexican small, glabrous species with a submarginal vein.

12. TRICHOMANES HOLOPTERUM Trichomanes holopterum Kunze, Farrnkra¨uter 1: 185. 1845. Lectotype (chosen by Proctor, 1977: 93). Kunze, Farrnkra¨uter 1: t. 77, f. 2. 1845. Figs. 319C, D. Similar to small specimens of T. galeottii in being deeply pinnatisect (the latter fully pinnate in mature specimens) and with winged rachises, but distinct in having the stipes alate to the frond bases (vs. alate only in the distal half) and the blades rounded at the apices (vs. attenuate), even on small plants; 2n⫽128 (Jam).

Rhizomes slender, 0.1–0.2 mm diam., long-creeping; rhizome hairs dense, brown, 1 mm long; roots absent; fronds 1–2.5 cm long, distant; stipes 1⁄10–1⁄4 the frond length, with dense, whitish brown hairs 0.8–1 mm long; blades round to ovate, 1–2 cm wide, margins glabrous, subentire, undulate, bases cuneate, apices rounded, membranaceous, glabrous; major veins free, connected by a continuous submarginal vein, false veins abundant, lying between the true veins, both parallel with and perpendicular to the true veins; sori near blade apices; involucres 1–1.5 ⫻ 0.8 mm, wholly immersed, mouths flaring, not dark-margined, not bilabiate.

Distribution. Terrestrial in wet forests; 220 m. USA (Fla); Mexico; Gr & L Ant. Specimen Examined. Chis (Martı´nez S. 15009, UAMIZ).

TRICHOMANES HYMENOPHYLLOIDES

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Trichomanes holopterum is generally Caribbean in distribution, and the one Mexican collectoin may turn out to be an odd specimen of T. galeottii.

veins that appear almost subpalmate, and the broad fronds with overlapping segments. See T. bucinatum for distinguishing characters.

13. TRICHOMANES HYMENOIDES

14. TRICHOMANES HYMENOPHYLLOIDES

Trichomanes hymenoides Hedw., Fil. Gen. Sp., t. 3. 1799. Figs. 322J, K. Trichomanes muscoides Sw., J. Bot. (Schrader) 1800(2): 95. 1801, nom. superfl. illeg. Didymoglossum muscoides (Sw.) Desv., Me´m. Soc. Linn. Paris 6: 330. 1827. Didymoglossum hymenoides (Hedwig) Copel., Philipp. J. Sci. 67: 77. 1938. Type. Not found by Wessels Boer (1962), who designated Hedwig’s plate as lectotype; probably based on a Jamaican specimen collected by Swartz. Trichomanes apodum Hook. & Grev., Icon. Filic. 1: t. 117. 1831. Type. Barbados. Parker s.n. (K!). Microgonium schaffneri Bosch ex Fe´e, Me´m. Foug. 9: 38. 1857, non Trichomanes schaffneri Schltdl., 1853 [1854]. Type. Mexico. Veracruz: Huatusco, Schaffner 2 (P?, not found).

Rhizomes long-creeping, 0.1–0.3 mm diam.; rhizome hairs dark brown, 1 mm long; roots absent; fronds (0.9–)1.5–3.5 cm long, distant; stipes ca. 1⁄10–1⁄4 the frond length, brown, with dense, dark brown hairs like those of the rhizomes; blades broad, pinnatifid to bipinnatifid, 0.3–1.5 cm wide oblong or obovate, sinuses deep, segments overlapping, margins membranaceous, with simple, forked, or stellate hairs, otherwise glabrous; veins free, major veins appearing almost palmate, false veins sparse, not parallel to margins; sori 1-several near blade apices; involucres nearly free from blade tissue, 1–1.5 ⫻ 1 mm, mouth flaring, darkmargined, bilabiate.

Distribution. Epiphytic in wet montane forests; 250–1450 m. Mexico; Guat, Bel, Salv, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Col, Ec, Peru, Braz, Bol, ne Arg, Uru. Specimens Examined. Chis (Breedlove 25006, MICH, NY, 57063, ENCB). Oax (Mickel 5905, 6120, 6189b, NY). Unverified, Doubtful, or Mistaken Reports. Ver (Pringle 10809, CAS, F, UC, US; Purpus 2929, F, US, both cited by Pacheco, 1994, but not verified).

Trichomanes hymenoides is very close to T. krausii (q.v.) and can be distinguished by its small size, sessile stellate marginal hairs, lack of stellate hairs in sinuses, the pinnate main lateral

Trichomanes hymenophylloides Bosch, Ned. Kruidk. Arch. 5: 209. 1863. Figs. 321A–C. Trichomanes leptophyllum Bosch, Ned. Kruidk. Arch. 4: 363. 1858 [1859], hom. illeg., non A. Cunn., 1836. Vandenboschia hymenophylloides (Bosch) Copel., Philipp. J. Sci. 67: 53. 1938. Type. St. Vincent. Guilding s.n. (isotype GH!).

Rhizomes thin, 0.5–0.8 mm diam., long-creeping; rhizome hairs of two kinds: spreading, thin, reddish brown, 1 mm long, and also appressed, light brown, 0.5 mm long; roots present; fronds 2.5–14 cm long, distant; stipes 1⁄5–1⁄3 the frond length, flattened, 0.5–0.8 mm diam., green, dark at bases, alate halfway to bases, with sparse light brown hairs 0.5 mm long at bases; blades membranaceous, bipinnate-pinnatifid to tripinnate, 1.5–4 cm wide, not reduced proximally, apices pinnatifid; rachises alate, with short (less than 0.1 mm long), 2-celled, clavate, bicolorous hairs (black at bases, light at tips), blades otherwise glabrous; ultimate segments 0.3–0.5 mm wide, margins entire; veins free, false veins absent; sori 1 per segment; involucres 1.5–2 ⫻ 0.5 mm, alate, mouths flaring, 1–1.5 mm wide, not bilabiate, concolorous or dark-margined; 2n⫽72 (Jam, Trin).

Distribution. Epiphytic or terrestrial in wet montane forests; 1600–2000 m. Mexico; Guat, Hond, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Braz. Selected Specimens Examined. Chis (Breedlove 35234, DS, 57072, CAS; Gittens 4198, NY). Hgo (Lorence 4937, MEXU). Me´x (Tejero-Dı´ez 2256, IZTA). Mor (Lyonnet 2859, MEXU, US). Oax (Mickel 4273, 4291, 5564, 6564, NY). Ver (Ventura A. 18487, ENCB, XAL).

Trichomanes hymenophylloides is closely allied to T. diaphanum Kunth (Guat to CR; Ven and Col to Peru, Braz, and Bol), but T. diaphanum has stipes alate to the bases and broadly alate rachises. For comparison with T. pyxidiferum, see that species and the key. Trichomanes hymenophylloides occurs at generally higher elevations than does T. pyxidiferum.

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TRICHOMANES

15. TRICHOMANES KRAUSII Trichomanes krausii Hook. & Grev., Icon. Filic., t. 149. 1831. Figs. 322A, B. Didymoglossum krausii (Hook. & Grev.) C. Presl, Abh. Ko¨nigl. Bo¨hm. Ges. Wiss., ser. 5, 3: 115. 1843. Hemiphlebium krausii (Hook. & Grev.) Prantl, Unters. Morph. Gefa¨sskrypt. 1: 46. 1875. Type. Dominica. Kraus s.n. (E).

Difficult to distinguish from the much more common T. reptans, but T. krausii can be identified by its occurrence at low elevations, 0–500(–1070) m (vs. 900–2500 m in T. reptans), and the false veins being more or less parallel to the margins and generally not between the true veins; 2n⫽136 (Jam).

Rhizomes stout, 3–7 mm diam., erect; rhizome hairs reddish brown, 1.5–2 mm long; roots present; fronds 10–60 cm long, clumped; stipes ca. 1⁄8 the frond length, 0.3–0.8 mm wide, green to brown, non-alate, with hairs similar to those of rhizomes; blades lanceolate, membranaceous, bipinnatifid, 3–8 cm wide, proximal pinnae reduced, apices pinnatifid; rachises alate to basal pinnae or basal pinna stalked; pinna margins entire, flat; veins free, forked, with dense, simple, jointed, 1–3 mm long, reddish brown hairs, false veins absent; involucres immersed in blade tissue, triangular-tubular, 1.5–2 ⫻ 1–1.5 mm at apices.

Distribution. Epiphytic in montane pine-oak woods; 1800– 2150 m. Mexico; Hond (Hawkins 917, MO), CR, Pan; Jam, Guad; Col, Ven, Ec, Peru, se Braz, Bol. Distribution. 0–500 m. USA (Fla); Mexico; Guat, Hond, Salv, Nic, CR, Pan; Gr & L Ant; Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol, Parag, Arg. Selected Specimens Examined. Chis (Breedlove 32824, DS, 57304, CAS). Oax (Mickel 5770, 7390a, NY, UC). Pue (Pacheco 3214, UAMIZ). SLP (Mickel 595, ISC). Tam (Pacheco 3027, IEB, UAMIZ, 3028, MEXU, UAMIZ). Ver (Copeland herb. 11, UC, US; Riba et al. 1155, UAMIZ). Unverified, Doubtful, or Mistaken Reports. QR (Castillo R. & Tun G. 290, UADY, XAL, cited by Palacios-Rios, 1992, but not verified).

Trichomanes krausii is distinct by its occurrence at low elevations, some false veins parallel to the blade margins (virtually none between true veins), and immersed sori. These characters work best for specimens below 500 m, and character states of T. reptans work best for specimens above 1500 m. At intermediate elevations specimens combine the characters, e.g., few false veins and exserted sori. Generally the most false veins are found in specimens collected above 2000 m and fewest below 500 m. More work is needed to resolve the taxonomic status of these two taxa.

16. TRICHOMANES LUCENS Trichomanes lucens Sw., Prodr. 136. 1788. Type. Jamaica. Swartz s.n. (S; isotypes B, BM, photo NY!). Figs. 320E, F.

Specimens Examined. Oax (Mickel 5588, NY; Rzedowski 33351a, ENCB, IEB).

This is an unusual range for this distinctive species, whose nearest stations are in Jamaica and Honduras. It is distinguished by its bipinnatifid, densely hairy blades.

17. TRICHOMANES MEMBRANACEUM Trichomanes membranaceum L., Sp. Pl. 2: 1097. 1753. Figs. 317G, H, J. Lecanium membranaceum (L.) C. Presl, Gefa¨ssbu¨ndel Farrn 104. 1843. Hemiphlebium membranaceum (L.) Prantl, Unters. Morph. Gefa¨sskrypt. 1: 46. 1875. Lecanolepis membranacea (L.) Pic. Serm., Webbia 28: 449. 1973. Type. “America.” Without exact locality, LINN 1253.1; Linnaeus also cited Sloane and Plumier references pertaining to material from Jamaica and Hispaniola.

Rhizomes slender, 0.5(–0.8) mm wide, long-creeping; rhizome hairs dark brown, 0.8–1 mm long; roots present; fronds 2–6 cm long, distant; stipes short, 1–1.5 mm long, 1 mm diam., with dense, dark brown hairs 0.8–1 mm long; sterile blades orbicular and entire to divided into ragged, cuneate segments, 1.5–4 mm long and wide, laminar surfaces glabrous, abaxial and adaxial margins with overlapping, orbicular, grayish brown scales 1–1.5 mm diam.; fertile blades broadly to narrowly cuneate, often divided into ragged lobes, glabrous, margins with scattered orbicular scales like those of sterile fronds or glabrous; veins free, forked, false veins parallel to true veins, unconnected; involucres

TRICHOMANES PETERSII

679

immersed to stalked, 0.5 mm wide at middle, 1.5–1.8 mm long, mouths 0.8 mm wide, dark-margined, bilabiate; 2n⫽68 (Jam).

Distribution. Epiphytic, epipetric or terrestrial on banks in wet forests; 350–600 m. Mexico; Guat, Bel, Hond, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Bol. Specimens Examined. Chis (Espejo & Herna´ndez 1441, UAMIZ). Tab (Gilly & Herna´ndez X. 284, UAMIZ, US). Ver (Pe´rez Garcı´a 35, UAMIZ; Wendt et al. 3844A, NY).

The diverse nature of the fronds, from orbicular to narrowly cuneate proximally, marginal orbicular scales, and unconnected, parallel false veins distinguish this species.

18. TRICHOMANES OVALE Trichomanes ovale (E. Fourn.) Wess. Boer, Acta Bot. Neerl. 11: 296. 1962. Figs. 317C, D. Didymoglossum ovale E. Fourn., Bull. Soc. Bot. France 19: 240. 1872. Lectotype (chosen by Wessels Boer, 1962: 296). Venezuela. Colonia Tovar, Fendler 25 (P; isotypes B, G, GH!, MO, YU!).

Rhizomes long-creeping, slender, 0.1 mm diam.; rhizome hairs dark brown, 0.1 mm long; roots absent; fronds 2–8 mm long, distant; stipes 1–4 mm long, 1⁄4–1⁄2 the frond length, with hairs similar to those of rhizomes; blades obovate to narrowly oblanceolate, entire or occasionally shallowly to deeply lobed, 2–4 mm wide, bases cuneate to cordate, apices rounded, margins membranaceous, with sparse, black or reddish brown, stellate hairs; veins pinnate to flabellate, free, false veins parallel to the true veins; sori one per blade; involucres immersed to partly exserted, with bilabiate lips, dark-margined with a few rows of darkened cells.

Distribution. Epiphytic in montane rain forests; 150 m. Mexico; Guat, Bel, Hond, Nic, CR, Pan; Gr Ant; Col, Ven, Sur, Braz. Specimens Examined. Chis (Breedlove 38177, DS; Martı´nez S. 14882, ENCB, UAMIZ, 25325, MEXU; Riba 1370, UAMIZ). Oax (Mickel 6870, NY). Tab (Cowan 3141, CAS, MEXU, NY). Unverified, Doubtful, or Mistaken Reports. QR (Castillo-Rivero 349, UADY, XAL, cited by Palacios-Rios, 2002c, but not verified).

Trichomanes ovale is closely related to Trichomanes petersii in being small, lacking submarginal veins, and possessing stellate marginal hairs, but the latter is distinct in lacking the dark involucral margins typical of T. ovale. The variable fronds (some narrowly lanceolate, some variously lobed), some stellate hairs at the tips of pinnae, and the strongly exserted sori with wide lips separate this species from other members of sect. Didymoglossum.

19. TRICHOMANES PETERSII Trichomanes petersii A. Gray, Amer. J. Sci. Arts 15: 326. 1853. Figs. 318C, D. Microgonium petersii (A. Gray) Bosch, Verh. Kon. Ned. Akad. Wetensch., Afd. Natuurk. 9(6): 7. 1861. Hemiphlebium petersii (A. Gray) Prantl, Unters. Morph. Gefa¨sskrypt. 1: 46. 1875. Didymoglossum petersii (A. Gray) Copel., Philipp. J. Sci. 67: 78. 1938. Type. U.S.A. Alabama: Hancock Co. [Winston Co.], near Sipsey River, 1 Jun 1853, Peters s.n. (not found in GH by Wessels Boer; topotypes collected by Peters in B, F, G, GH!, K!, L, MO, U, UC!, US-says Winston Co.). Trichomanes schaffneri Schltdl., Linnaea 26: 368. 1853 [1854]. Type. Mexico. [Veracruz:] Huatusco, Schaffner s.n. (B).

Rhizomes short- to long-creeping, slender, 0.1 mm diam.; rhizome hairs dense, dark brown, 0.1 mm long; roots absent; fronds 1 cm long, distant; stipes 1⁄4 to nearly 1⁄2 the frond length; blades ovate, 0.4 cm wide, bases cuneate, apices rounded, 0.3–0.5 mm long, margins membranaceous, with dark reddish brown forked or stellate hairs; veins free, false veins parallel to the true veins; sori one per blade, apical; involucres immersed in blade tissue, with mouths flaring, not bilabiate, concolorous.

680

TRICHOMANES

Distribution. Walls of calcareous sinkholes; 1400–2200 m. Se USA; Mexico; Guat, Hond, Salv, Nic, CR. Specimens Examined. Chis (Sharp 451021, MEXU, US; Xolocotzi & Sharp X-651, US). Pue (Sharp 451021, 45321, US). Ver (Delgadillo 3393, UAMIZ; Rebolledo s.n., 19 Sep 1964, US).

Distribution. Terrestrial in wet forests at low elevation in Atlantic lowlands; 200–600 m. Mexico; Guat, Bel, Hond, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, s Braz, Bol.

Trichomanes petersii is closely related to T. ovale (q.v.), which differs in having dark-lipped involucres and epiphytic habitat.

Selected Specimens Examined. Chis (Martı´nez S. 25014, 25342, MEXU). Gro (Langlasse´ 672, MEXU; Lorea 2885, F, FCME). Oax (Mexia 9133, NY; Mickel 5830, NY). Tab (Edith et al. 12, XAL; Manrı´quez L. 19, ENCB). Ver (Orcutt 3198, US; Wendt et al. 4803, NY).

20. TRICHOMANES PINNATUM

Trichomanes pinnatum is distinct in its pinnate fronds with false veins perpendicular to the true veins.

Trichomanes pinnatum Hedw., Fil. Gen. Sp., t. 4. 1799. Type. Hedwig’s plate, t. 4, f. 1. 1799, based on material from Jamaica, possibly sent to Hedwig by Swartz. Figs. 320A, B. Rhizomes stout, 3–5 mm diam., suberect; rhizome hairs dark brown, 1.5–2 mm long; roots present; fronds 12–40 cm long, clumped; stipes 1⁄4–1⁄3 the sterile fronds, 1⁄2 the fertile fronds, stout, 1–1.3 mm wide, brown, non-alate, with scattered dark brown hairs 1–1.5 mm long; blades pinnate, 8–15 cm wide, not reduced at bases, each with a conform terminal pinna, ovate to deltate, light green, membranaceous; rachises sometimes extended and proliferous, alate with hairs similar to those of stipes; pinnae 8–15 pairs, basal pair stalked, glabrous, bases truncate, apices acute, margins irregularly denticulate, sinuses between vein ends 0.3–1 mm deep; veins free, simple or 1-forked, rarely netted at base of sori, but with abundant false veins perpendicular to the true veins making them appear netted, false veins forming a submarginal vein; involucres stalked, free from blade tissue, 1 mm long, 0.3 mm wide at bases, 0.5–0.8 mm wide in middle, mouths not flaring, 0.5 mm wide, not bilabiate, concolorous, stalks 0.5 mm long; 2n⫽ca. 72 (Braz).

21. TRICHOMANES POLYPODIOIDES Trichomanes polypodioides L., Sp. Pl. 2: 1098. 1753. Neotype (chosen by Proctor, Fl. L. Antill. 2: 93. 1977). Montserrat. Proctor 19068 (A; isoneotype US!, GH!). Figs. 320C, D. Trichomanes sinuosum Rich. ex Willd., Sp. Pl., ed. 4, 5(1): 502. 1810. Type. Guadeloupe. Richard s.n. (B-Willd. 20191!).

Rhizomes slender, 0.3–0.5 mm diam., green, long-creeping; rhizome hairs sparse, 1 mm long, brown; roots present; fronds 4–15 cm long, distant; stipes less than 1⁄10 the frond length, greenish, 0.3 mm wide, with sessile, forked hairs 1 mm long; blades pinnatifid, linear-lanceolate, 1–3 cm wide, alate to bases, margins entire, with brown, sessile, stellate hairs 1 mm long on veins and margins; veins free, with scattered to rare, short, black, clavate hairs 0.1 mm long; sori 1–2 at segment apices; involucres immersed in blade tissue, 1.5 mm long, mouths slightly flaring, 1 mm wide, not bilabiate, concolorous; n⫽64 (Trin).

TRICHOMANES RADICANS

681

Distribution. Epiphytic in wet montane forests along Atlantic slope; 250–1300 m. Mexico; Guat, Bel, Hond, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol, Parag, Uru.

Distribution. Epiphytic or terrestrial in wet montane forests; 450–1400(–1700) m. Mexico; Guat, Bel, Hond, Nic, CR, Pan; Gr Ant; Col, Ven, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol; Africa.

Selected Specimens Examined. Chis (Breedlove 30907, 31522, 32121, MICH; Dressler 1603, MICH; Martı´nez S. 13494, NY). Gro (Lorea 2886, FCME, NY). Oax (Hallberg 1282, NY; Herna´ndez G. 685, NY). Ver (Purpus 2772, NY; Ventura A. 15230, NY).

Selected Specimens Examined. Chis (Breedlove 24099, NY; Martı´nez S. 20489, NY). Oax (Hallberg 1563, NY; Mickel 6469, 7335, NY). Pue (Sharp 441731 bis, US). Qro (Servin O. 957, IEB, NY). SLP (Frye & Frye 2662, NY; Pringle 3800, NY; Rzedowski 10983, NY). Ver (Mu¨ller 400, NY).

Trichomanes polypodioides is distinct by its small pinnatifid blades with stellate hairs and lack of false veins.

22. TRICHOMANES PYXIDIFERUM Trichomanes pyxidiferum L., Sp. Pl. 2: 1098. 1753. Figs. 321D–F. Vandenboschia pyxidifera (L.) Copel., Philipp. J. Sci. 67: 53. 1938. Type. Plumier, Traite´ Foug. Ame´r., pl. 50E, 1705, illustrating a plant from Haiti. Trichomanes cavifolium Mu¨ll. Berol., Bot. Zeitung (Berlin) 12: 753. 1854. Type. Mexico. Schiede s.n. (B?).

Rhizomes slender, long-creeping, 0.5–0.8 mm diam.; rhizome hairs dense, black, 0.5 mm long; roots present; fronds 3–10 cm long, distant; stipes ca. 1⁄3 the frond length, blackish, with dense, black hairs 0.5 mm long at bases, running up stipes along two margins, alate 3⁄4 to all the way to base; blades 2–3-pinnate, 1.5–1 cm wide, not reduced at bases, apices pinnatifid; rachises alate; pinnae 4–10 pairs, oblique, nearly glabrous with scattered to rare minute glandular hairs; ultimate segments linear, 0.8–1 mm wide, obtuse, margins entire, often drying with long narrow “folds” parallel to the veins, giving the appearance of false veins; veins free, false veins absent; involucres immersed in blade tissue, 1.5–2 ⫻ 1 mm, mouths flaring, 1.3–1.5 mm wide, concolorous, not bilabiate.

Trichomanes pyxidiferum is similar to T. hymenophylloides, but the ultimate segments are wider (0.8–1 mm) and the rhizomes and stipes have dense black hairs; the former is found at lower elevations (450–1400(–1700) m).

23. TRICHOMANES RADICANS Trichomanes radicans Sw., J. Bot. (Schrader) 1800(2): 97. 1801. Figs. 316A–C, 319K–M. Trichomanes scandens Hedw., Fil. Gen. Sp., t. 6. 1799, hom. illeg., non L., 1753. Vandenboschia radicans (Sw.) Copel., Philipp. J. Sci. 67: 51, 54. 1938. Type. Jamaica. Swartz s.n. (S, photos US). Trichomanes kunzeanum Hook., Sp. Fil. 1: 127, t. 39d. 1844. Syntypes. Peru. Pangoa, Matthews 1088 (K!), “Papayaco,” Poeppig s.n. (K!); Venezuela. “Caracas,” Linden 176 (K?; isosyntype BM). Trichomanes mexicanum Bosch, Ned. Kruidk. Arch. 5: 164. 1861. Syntypes. Mexico. Schiede 806 (B), Schaffner 7 (P?, not found).

Rhizomes stout, 1–2 mm diam., long-creeping; rhizome hairs dense, slender, brown, 1 mm long, and also with occasional thick, dark brown hairs 1–1.5 mm long at frond bases and on stipes; roots absent on climbing stems; fronds 22–30 cm long, distant; stipes ca. 1⁄4 the frond length, stout, flattened, 0.8–1 mm wide, alate 3⁄4 of the way to bases, with scattered, short, glandular hairs 0.1 mm long; blades bipinnate to tripinnate-pinnatifid, 6– 10 cm wide, ovate to lanceolate, barely reduced at bases; rachises alate, glabrous except for stubby blackish hairs; pinnae 12–18 pairs, lanceolate, ultimate segments 0.5–0.8 mm wide, margins entire; veins free, false veins absent; sori terminating segments; involucres free from or barely immersed in blade tissue, elongate, 1.8–2 mm long, 0.5–0.8 mm wide at middle, 0.3 mm wide at bases, lips erect, straight or only slightly flaring, concolorous; 2n⫽72 (Chis, Jam).

682

TRICHOMANES

lucres exserted to slightly immersed, 2.5 ⫻ 1.0–1.2 mm, 0.8–1 mm wide at middle, mouths flaring, bilabiate, dark-margined, lips 0.5 mm long.

Distribution. Epiphytic in wet montane forests; (600–)1250– 2450 m. Mexico; Guat, Hond, Salv, Nic, CR, Pan; Gr & L Ant; Col, Ven, Guy, Sur, Fr Gui, Ec, Peru, s Braz, Bol, Parag; Europe, Asia, Africa. Selected Specimens Examined. Chih (Correll & Gentry 23088, F, MO, NY, UC, US). Chis (Breedlove 29454, NY). Dgo (Sanders et al. 4876, UC). Gro (Hinton 14165, NY). Hgo (Moore 5345, US). Jal (McVaugh 20439, MICH). Me´x (Tejero-Dı´ez 2966, IZTA). Mich (Rzedowski 46274, XAL). Mor (Lyonnet 2861, US). Oax (Pringle 4677, NY). Pue (Barnes & Land 497, NY). Qro (Rubio 391, ENCB, MEXU). SLP (Rzedowski 10952, NY). Ver (Mu¨ller 390, NY). Unverified, Doubtful, or Mistaken Reports. Tab (Linden 1508, cited by Rovirosa, 1909, but not verified; Rovirosa 1033, s. herb., cited by Cowan, 1983, but not verified).

Trichomanes radicans closely resembles T. collariatum in its relatively large fronds and long-creeping rhizomes but is distinct by its involucres with narrow, non-flared lips, long stipes, and broad blades. Specimens from northern states of Mexico are often named T. boschianum J. W. Sturm, a species of the eastern United States, but transitional forms suggest that these small Mexican specimens are merely dwarf forms of T. radicans.

Distribution. Epiphytic and epipetric in wet montane forests; 900–2500 m. Mexico; Guat, Hond, Salv, Nic, CR, Pan; Jam, Hisp; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, s Braz, Bol, Parag, n Arg. Selected Specimens Examined. Chis (Breedlove 32612, MICH; Somers 281, NY). Gro (Lorea 1997, FCME). Hgo (Gimate L. 971, MEXU). Jal (McVaugh 20438, NY). Oax (Mexia 9267a, NY; Mickel 982, ENCB, ISC, MICH, US). Pue (Barnes & Land 500, US). Ver (Pringle 10809, LL, UC, XAL; Purpus 8333, NY, UC).

Closely resembling T. bucinatum, T. reptans is distinct by its non-stalked sori, narrow involucral lips, and marginal forked and stellate hairs restricted to the sinus bases.

25. TRICHOMANES RIGIDUM Trichomanes rigidum Sw., Prodr. 137. 1788, non Hedw., 1802. Figs. 321G, H. Selenodesmium rigidum (Sw.) Copel., Philipp. J. Sci. 67: 81. 1938. Type. Jamaica. Swartz s.n. (S?)

24. TRICHOMANES REPTANS Trichomanes reptans Sw., Prodr. 136. 1788.

Figs. 319A, B.

Didymoglossum reptans (Sw.) C. Presl, Abh. Ko¨nigl. Bo¨hm. Ges. Wiss., ser. 5, 3: 115. 1843. Hemiphlebium reptans (Sw.) Prantl, Unters. Morph. Gefa¨sskrypt. 1: 46. 1875. Type. Jamaica. Swartz s.n. (S).

Rhizomes slender, 0.3 mm diam., long-creeping; rhizome hairs dense, dark brown, 0.5 mm; roots absent; fronds 2.5–8 cm long, distant; stipes ca. 1⁄10–1⁄3 the frond length, terete to flattened, 0.8 mm wide, with dark brown hairs 0.3–0.5 mm long; blades variable in shape, broadly lanceolate, ovate, or obovate, pinnatifid to bipinnatifid, 1.5–3.5 cm wide; rachises alate in distal 1⁄2; pinnae non-overlapping, membranaceous, margins Ⳳ with sparse, black, forked hairs and stellate hairs in sinuses, scattered clavate hairs 0.1 mm long on veins and involucres; veins free, major ones strongly pinnate, false veins parallel to true veins, many unconnected, some parallel to and very near the blade margins; invo-

Rhizomes stout, 5–7 mm diam., erect or ascending, to 4 cm long; rhizome hairs dark brown, stiff, lustrous, 1–1.5 mm long; roots present; fronds 20–35 cm long, clumped; stipes 1⁄2 the frond length, stout, 0.8–1 mm diam., brown, non-alate, glabrous except with hairs at base like rhizome hairs; blades 3–4-pinnate, 10–15 cm wide, subdeltate or broadly ovate, dark green; rachises glabrous except for stubby blackish hairs; pinnae 12–20 pairs, petiolulate, ascending or spreading, with occasional stubby blackish hairs, ultimate segments linear, 0.3–0.5 mm wide, margins entire; veins free, false veins absent; involucres stalked with narrow lateral wings, 1.0–1.2 ⫻ 0.3–0.5 mm, mouths not flared, entire, slightly bilabiate, concolorous; 2n⫽66 (Jam).

VITTARIA

Distribution. Terrestrial in wet montane forests of Atlantic slope; 300–1650 m. Mexico; Guat, Bel, Hond, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, s Braz, Bol; Old World tropics, though paleotropical material may represent a distinct taxon. Selected Specimens Examined. Chis (Breedlove 32413, MICH, 32542, DS, MICH, NY). Oax (Herna´ndez G. 1719, UAMIZ; Mexia 9275, NY; Mickel 943, ISC, MICH, 1035, MICH, NY, 1559, 5692, NY; Rzedowski 30684, NY). Ver (Wendt et al. 4802, NY).

One of the few terrestrial species of Mexican Trichomanes, T. rigidum is distinct by its erect rhizomes, coarse, non-alate stipes, and finely dissected blades.

26. TRICHOMANES TUERCKHEIMII Trichomanes tuerckheimii Christ, Hedwigia 44: 361. 1905. Lectotype (chosen by Stolze, 1989: 90). Guatemala. Alta Verapaz: Cubilquitz, Tu¨rckheim 8348 (P!; isolectotypes GH!, K!, photo US!, P!). Figs. 319E–H, J. Rhizomes slender, 0.8–1 mm diam., long-creeping, green; rhizome hairs on lower side dark brown, 0.5–1 mm long; roots absent; fronds 6.5–20 cm long, distant; stipes virtually absent, 1–2 ⫻ 0.8–1 mm, with dense hairs abaxially like those of rhizomes; blades pinnatifid, 4–11 cm wide, with pinnatifid terminal pinnae, not reduced proximally, ovate to lanceolate; pinnae 10–20 pairs, glabrous except for dense hairs similar to those of stipes on abaxial surfaces of rachises, veins, and often along blade margins affixing the fronds to the substrate, margins entire; veins free, not reaching margins, with small, unconnected false veins parallel to true veins; involucres stalked, free from blade tissue; 2– 2.2 mm long, 0.5–0.8 mm wide at mouths, mouths round, concolorous.

683

Distribution. Epiphytic in wet forests at low elevations; 50– 500 m in Guat. Mexico; Guat, Bel, Hond, Nic, CR, Pan; Col, Ven, Guy, Sur, Fr Gui, Ec, Peru. Specimens Examined. Chis (Martı´nez S. 15908, CAS, IBUG, IEB, IBUG, MEXU, MO, UAMIZ). Ver (Gilly & Herna´ndez X. 44, US; Sa´nchez 44, US).

The peculiar habit of the leaves adhering to the bark surface by trichomes on veins and margins is unique among Mexican species of Trichomanes. Excluded Species Trichomanes plumosum Kunze, Linnaea 9: 104. 1834. Type. Peru. Poeppig 1107 (LZ, destroyed). Similar to Trichomanes crispum except for its long-creeping (vs. short-creeping to suberect) rhizomes. Pacheco (1994) cited a specimen from Chiapas (Linden 59, BR). We have seen the specimen, which certainly seems to be Trichomanes plumosum, but the great distance from the nearest known collections (Pan, Col, Ven, Ec, Peru) suggests that the label was mistaken, Linden having collected also in Venezuela and Colombia.

122. VITTARIA Vittaria Sm., Me´m. Acad. Roy. Sci. (Turin) 5: 413, pl. 9. 1793. Type: Vittaria lineata (L.) Sm. [⬅ Pteris lineata L.]. Epiphytic; rhizomes short-creeping, rarely erect, dorsiventral, protostelic, with strongly clathrate scales at the apices; fronds small to medium-sized, linear, monomorphic, clumped; stipes short to nearly absent, greenish or greenish brown, somewhat flattened; blades linear, entire, subcoriaceous to coriaceous, with a conspicuous to inconspicuous midrib, glabrous; veins obscure, anastomosing and forming a single row of areoles between midrib and blade margins, on each side of midrib; sori in a narrow submarginal lines, one on each side of midrib, immersed in a groove, indusia absent; paraphyses present, filiform to slightly clavate; spores tetrahedral or bilateral in one species, clear to pale yellowish, relatively smooth with scattered spherical deposits; gametophytes with paired gemmae; x⫽60.

684

VITTARIA

Vittaria, as recently redefined by Crane (1997), is primarily a genus of the Neotropics (5 spp., 4 in Mexico), plus a single species in Africa and offshore islands in the Indian Ocean. Formerly the genus included also many paleotropical species from Asia, Polynesia, and Africa, but Crane’s work has shown that the genus as traditionally construed appears to be polyphyletic. The Old World species once included in Vittaria are now placed in Haplopteris (ca. 20 spp.), with the exception of Vittaria isoetifolia Bory, in Africa and offshore islands. Vittaria sensu stricto (shoestring ferns) grows in wet forests of low to high elevation. Two of the Mexican species are widespread, the other two of narrower distribution in Mexico and Central America. The genus is distinct in its narrow, undivided fronds, linear, submarginal sori, and single row of areoles between midribs and margins. Vittaria appalachiana Farrar & Mickel, described only recently, is a species in the eastern United States, and is known

primarily from gametophytic populations; the sporophytes (embryos), if formed at all, are generally abortive, or very weak and small (Farrar & Mickel, 1991). It would not be surprising if this species were to be found eventually in northeastern Mexico, in appropriate habitats (dark, moist cavities and rock shelters in non-calcareous formations). References Benedict, R. C. 1914. A revision of the genus Vittaria J. E. Sm. I. The species of the subgenus Radiovittaria. Bull. Torrey Bot. Club 41: 391–410; Crane, E. H. 1997 [1998]. A revised circumscription of the genera of the fern family Vittariaceae. Syst. Bot. 22: 509–517; Farrar, D. R. & J. T. Mickel. 1991. Vittaria appalachiana: A name for the “Appalachian gametophyte.” Amer. Fern J. 81: 69– 75; Lellinger, D. B. 1972. Botany of the Guayana Highland. IX. Polypodiaceae subfam. Adiantoideae. Mem. New York Bot. Gard. 23: 1–23; Tryon, R. M. 1964a. Taxonomic fern notes. IV. Some American vittarioid ferns. Rhodora 66: 110– 117.

Key to the Mexican Species of Vittaria 1. Spores bilateral; paraphyses filiform, light brown; blades 2–3 mm wide at broadest point; rhizome scales 6–7 mm long with long (1⁄3–1⁄2 the scale length) hair-like tips. ............................................................................................................................................................... 4. V. lineata. 1. Spores tetrahedral; paraphyses with enlarged apical portion, reddish brown; blades 1–4 mm wide at broadest point; rhizome scales 3–7 mm long with or without long, hair-like tips. 2. Midribs stramineous on adaxial surfaces; blades 2–3(–4) mm wide at widest point; rhizome scales without hair-like tips; blades often somewhat falcate. ................................................................................................................................................................. 2. V. flavicosta. 2. Midribs not evident on adaxial surfaces; blades 1–2 mm wide at widest point; rhizome scales with or without hair-like tips; blades not or only slightly falcate. 3. Rhizome scales blackish with long uniseriate (hair-like) tips, seemingly entire or with only a few scattered teeth, the cell walls very thick, black, cell lumina very narrow, at mid-scale ca. 5 times longer than broad; scales (2–)6–12 cells wide at bases; 500–1500 m. ........................................................................................................................................................................... 1. V. bradeorum. 3. Rhizome scales reddish brown, usually lacking hair-like tips, seemingly toothed by virtue of marginal cells with end walls protruding outward, cell walls narrow, red-brown, lumina broad, at mid-scale to ca. 2 times longer than broad; scales 8– 20 cells wide at bases; (300–)1200–3050 m. ................................................................................................................................ 3. V. graminifolia.

1. VITTARIA BRADEORUM Vittaria bradeorum Rosenst., Repert. Spec. Nov. Regni Veg. 22: 18. 1925. Type. Costa Rica. Tablazo, Brade 165 (S, photo US; isotypes NY!, UC!, US). Figs. 323A–C. Rhizomes horizontal, compact, 1–1.5 mm diam.; rhizome scales clathrate, dimorphic in width and cell type, 3.5–7 mm long, from 2 cells (0.1 mm) to 12 cells (1 mm) wide at bases, apices narrowed to hair-like tip, 2 cells wide, appearing as a single hair (the central wall between the two cells) for ca. 1⁄3 the scale length, central cell walls often thicker and darker than cell walls near margins, or all walls thick and dark; fronds to 35 cm long, 1–2 mm wide at broadest point, margins recurved; midribs not visible on adaxial side, evident on abaxial side; paraphyses medium to dark red-brown, stout, 0.1 mm long, each with an enlarged apical cell (2 times or more longer than wide); spores tetrahedral.

Distribution. Epiphytic (occasionally epipetric) in rain forests and cloud forests; 500–1500 m. Mexico; Guat, CR. Cited by Moran (in Davidse et al., 1995) from Salv, Nic, and Pan, but the identities of these specimens need verification; the specimen cited by Moran from Chiapas is V. flavicosta. Selected Specimens Examined. Chis (Breedlove 29912, DS, 56508, CAS). Col (Lott 2986, NY, UC). Gro (Lorea 1867, FCME; Vela´zquez 773, FCME). Oax (Mickel 6670, 7348, NY, UC; Pringle 5951, GH, NY). Ver (Copeland herb. 160a, MICH, UC; Nee et al. 25019, MEXU; Rowell et al. 17M664, UC, TEX).

VITTARIA GRAMINIFOLIA Unverified, Doubtful, or Mistaken Reports. Pue (reported by Mickel & Beitel, 1988, probably based on Cruz Cisneros 1519, NY, which is V. graminifolia).

Vittaria bradeorum has tetrahedral spores and evident midribs, and appears to be the earliest name for this species, now that the identity of V. dimorpha (⫽ V. graminifolia) is known with more confidence. That name was synonymized by Lellinger (1972) under V. lineata, which differs in having bilateral spores. In recent years, V. dimorpha has been applied to specimens here called V. bradeorum (e.g., by Smith, 1981; Mickel & Beitel, 1988; Lellinger, 1989; Moran in Davidse et al., 1995), but in our opinion, the type of V. dimorpha has rhizome scales like those of V. graminifolia. Vittaria bradeorum and V. graminifolia have similar, tetrahedral spores. The narrow-bladed, tetrahedral-spored Vittarias still need careful monographic revision in order to understand their species limits and geographical distribution. Vittaria bradeorum appears to be scattered on both the Atlantic and Pacific slopes of Mexico at low to middle elevations, mostly below elevations at which V. graminifolia occurs. The rhizome scales have very thick black cell walls (vs. thin and reddish brown in V. graminifolia), and the scale tips are very long and uniseriate at tips. Individual cells at mid-scale are about 5 times longer than wide, in comparison with cells at mid-scale in V. graminifolia, which are only a little longer than wide. A further difference between the two species is that the scales of V. bradeorum lack or have only a few inconspicuous marginal teeth, thus appearing to be nearly entire; in V. graminifolia, the scales appear to be prominently toothed, because the end walls of the marginal cells protrude away from the scale body. The narrow fronds and lack of stramineous adaxial midribs distinguish V. bradeorum from V. flavicosta. Misidentified collections of this group abound in herbaria, and identifications, as given in citations by Smith (1981), Mickel & Beitel (1988), and Palacios-Rios (1992), have also been changed in a number of cases. For example, most of the specimens cited by Smith (1981) for V. dimorpha are now considered to be V. flavicosta; Mickel 4339 (NY, UC), Mickel 6222 (NY), and Pringle 5951 (NY), cited by Mickel and Beitel (1988) as V. dimorpha, are V. graminifolia; Mexia 9133a (see below) is not V. dimorpha but V. flavicosta; and Purpus 1966 (UC), cited by Palacios-Rios (1992) as V. dimorpha, is V. graminifolia.

2. VITTARIA FLAVICOSTA Vittaria flavicosta Mickel & Beitel, Mem. New York Bot. Gard. 46: 399–400. 1988. Type. Mexico. Oaxaca: Distrito Ixtla´n, 79 km N of Ixtla´n de Jua´rez on Rte 175, 1 km S of Campamento Vista Hermosa, ca. 1 hour hike on trail toward Tarabundı´, Mickel 5663 (NY!; isotype UC!). Figs. 323D–F. Rhizomes horizontal, compact, 1–1.5 mm diam.; rhizome scales clathrate, uniform in width, 3–5 mm long, 0.3–0.8 mm wide (4– 10 cells at bases), apices narrowed to hair-like tip, cell walls somewhat dimorphic, those of central part of each scale thicker and darker than cell walls near scale margins; fronds to 25 cm long,

685

2–3(–4) mm wide at widest point, 0.8–1 mm wide proximally, often somewhat falcate, margins not recurved; midribs stramineous, visible on upper surface, prominent on lower surface; paraphyses medium to dark red-brown, stout, 0.1 mm long or less, with enlarged apical cell (2 times or more longer than wide); spores tetrahedral.

Distribution. Epiphytic in wet montane forests, dense woods with stream, on limestone; 150–1400 m. Mexico; Guat, CR. Selected Specimens Examined. Chis (Breedlove 21674, DS, 31523, DS, NY, 32266, 32928, DS; Dressler 1601, MEXU, NY, US). Oax (Mickel 5838, 6414, 7140, 7278, NY, UC; Mexia 9133a (CAS, GH, NY, UC, US). Ver (Valdivia Q. 121, 938, 1530, 1817, 1819, XAL; Va´zquez T. 59, XAL).

This species has the broadest blades of any species in the V. graminifolia complex and occurs along the Atlantic slope from Oaxaca and Chiapas to Costa Rica at low to middle elevations. The width of the blades (2–4 mm) and stramineous adaxial midribs distinguish V. flavicosta from V. graminifolia and V. bradeorum. The tetrahedral spores and lack of paraphyses with funnelform apical cell separate it from species of Radiovittaria. Most of the collections cited by Smith (1981) as V. dimorpha are instead V. flavicosta.

3. VITTARIA GRAMINIFOLIA Vittaria graminifolia Kaulf., Enum. Filic. 192. 1824. Type. “Brasil. Otto comm.” (LZ, destroyed; isotype E). Figs. 323G, H, J, K. Vittaria filifolia Fe´e, Me´m. Foug. 3: 20, pl. 3, f. 6. 1852. Lectotype (chosen by Tryon, 1964a: 114). Guadeloupe. L’Herminier s.n. (P-CO!, frag. NY!; isolectotype NY!). Vittaria dimorpha Mu¨ll. Berol., Bot. Zeitung (Berlin) 12: 547. 1854. Type. Mexico. [Veracruz:] Huitamalco, Schiede 791 (B!, frag. NY!; isotype NY!).

Rhizomes horizontal, compact, 1–1.5 mm diam.; rhizome scales clathrate, 4–5 mm long, 0.8–1.3 mm wide, uniform in size and cell wall type, 2 cells wide only at terminal 3–4 cells of tip, 8– 20 cells wide at scale bases; fronds to 30 cm long, 1.5–2 mm broad at broadest point, 1–1.5 mm wide proximally, margins not recurved; midribs not visible on either surface (lower surface flat between sori); paraphyses medium to dark red-brown, stout, 0.1

686

WOODSIA

mm long, with enlarged apical cell (ovate, 2 times longer than wide); spores tetrahedral-globose; 2n⫽ca. 240 (unknown source).

Rhizomes horizontal, compact, 1–1.5 mm diam.; rhizome scales clathrate, 6–7 mm long, 0.3–0.8 mm wide, uniform in size and cell wall thickness, apices narrowed to hair-like tip, 2 cells wide appearing as a single hair (the central wall between the two cells) for 1⁄3–1⁄2 the scale length, 3–6 cells wide at scale bases; fronds to 40 cm long, 2–3 mm wide, margins recurved; midribs evident abaxially only; paraphyses tan, slender, curved slightly, filiform, 0.1 mm long, the apical cell of each paraphysis not enlarged; spores bilateral.

Distribution. Epiphytic in wet montane forests and cloud forests, rarely at low elevations; (300–)1200–3050 m. Mexico; Guat, Bel, Hond, Salv, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Sur, Ec, Peru, Braz, Bol. Selected Specimens Examined. Chis (Purpus 6731, NY, UC). Gro (Martı´nez S. 888, MEXU). Hgo (Rzedowski 28696, ENCB). Jal (McVaugh 21332, NY). Me´x (Hinton 6162, ENCB, NY). Mich (Arse`ne 5252, UC). Nay (Gonza´lez 867, MEXU, UAN). Mor (Lyonnet 778, MEXU, NY). Oax (Mickel 4106, NY, UC). Pue (Cruz Cisneros 1519, ENCB, NY). Qro (Dı´az Barriga & Carranza 6554, MEXU). Sin (Breedlove 44951, CAS). Ver (Copeland herb. 160, MICH, UC). Unverified, Doubtful, or Mistaken Reports. SLP (Schaffner 53, NY, provenance in doubt, see Rzedowski, 1959). Tab (reported by Magan˜a, 1992, possibly based on a misidentification). Tam (reported by Mickel & Beitel, 1988, but not verified; also reported by Sharp, 1954, and by Lof, 1980, from Rancho del Cielo).

This is the commonest and most widespread species in the genus in Mexico, and it is especially common in middle and higher elevation montane rain forests of the Atlantic slope in Oaxaca and Chiapas. The Vittaria graminifolia complex (narrow fronds, tetrahedral spores, paraphyses with reddish brown, elongate, enlarged apical cells) needs careful study. The presence in V. bradeorum of dimorphic V. lineata-type rhizome scales and V. graminifolia-type tetrahedral spores has led to numerous misidentifications of specimens of these three species, especially if only the rhizome scales are examined. Vittaria graminifolia has wide, dark brown to reddish brown rhizome scales of uniform size without any part of the cell walls thickened, as well as a flattened lower blade surface without an evident midrib. See V. bradeorum for additional comments.

Distribution. Epiphytic in lowland and lower montane forests, often on palms; 50–500 m. USA (Fla); Mexico; Guat, Bel, Hond, Nic, CR, Pan; Gr & L Ant; Col, Ven, Trin, Guy, Sur, Fr Gui, Ec, Peru, Braz, Bol, Parag, Uru. Selected Specimens Examined. Camp (Martı´nez S. 27880-A, XAL, MEXU). Chis (Breedlove 33174, DS, 57331, CAS; Martı´nez S. 24991, MEXU, XAL). QR (Castillo R. 195, XAL; Palacios-Rios 3650, XAL). SLP (Pringle 4078, CAS, DS, UC). Tab (Croat 40070, MO, UC; Ortı´z 2156, MEXU). Ver (Croat & Hannon 63106, MO; Nee & Taylor 29183, XAL).

This species is abundant in the West Indies (at low and middle elevations) and perhaps South America, but in Mexico and Central America it is relatively uncommon (V. graminifolia and V. bradeorum are much more common). Most reports of this species in Mexico are based on misidentifications of V. graminifolia or V. bradeorum. No other species of Vittaria in Mexico or Mesoamerica has bilateral spores and filiform paraphyses. In addition, V. lineata occurs at elevations below about 500 m, whereas V. graminifolia is almost always at higher elevations. Lellinger (1972) synonymized V. deppeana under V. graminifolia, but the type of the former has bilateral spores, and thus agrees with V. lineata.

1 2 3. WO O D SI A 4. VITTARIA LINEATA Vittaria lineata (L.) Sm., Me´m. Acad. Roy. Sci. (Turin) 5: 421, t. 9, f. 5. 1793. Figs. 323L–O. Pteris lineata L., Sp. Pl. 2: 1073. 1753. Lectotype (chosen by R. Tryon 1964a: 213). Hispaniola. Plumier, Traite´ Foug. Ame´r., pl. 143. 1705. Vittaria deppeana Mu¨ll. Berol., Bot. Zeitung (Berlin) 12: 547. 1854. Type. Mexico. [Veracruz:] Jalapa, Schiede 790 (B!, frag. NY!; isotype NY!).

by John T. Mickel and Michael D. Windham

Woodsia R. Br., Prodr. 158, Obs. 4. 1810. Type: Woodsia ilvensis (L.) R. Br. [⬅ Acrostichum ilvense L.]. Terrestrial or epipetric; rhizomes compact to short-creeping, horizontal to ascending or erect; rhizome scales linear-lanceolate, orange-tan at rhizome apices, later bicolorous, each with black central stripe; fronds generally small, clumped, monomorphic;

WOODSIA CANESCENS

stipes stramineous, sometimes dark at base, vascular bundles 2, round or oblong in cross section; blades 1–2-pinnate-pinnatifid (rarely tripinnate), linear to lanceolate or ovate, membranous to subcoriaceous; pinnae not articulate to rachis, segment margins entire to dentate, sessile, bases usually nearly equilateral; costae often shallowly grooved adaxially, grooves more or less continuous from rachis to costae; indument of glandular (occasionally non-glandular) hairs on both surfaces, rarely absent; veins free, simple or forked, ending before the blade margins, the distal 0.5 mm of the veins swollen, pale, evident, occasionally sunken; sori in one row between costae and margin on ultimate segments, round; indusia basal, dissected into several to numerous filamentous or scale-like segments encircling the sori, persistent but often obscure in mature sori; spores bilateral, brownish, cristate or rugose; x⫽38, 39, 41.

687

Species about 35 (8 in the flora), mostly north temperate regions and higher elevations in the tropics. Of the Mexican species, only one (W. mollis) is widespread, the rest comprising a difficult complex of hybridizing taxa, mostly in northwestern Mexico. Woodsia is often confused with Cystopteris in herbarium collections, though the resemblance may be superficial. Woodsia is easily distinguished from Cystopteris by its persistent stipe bases, multilobed indusia, and obscure veins that end in hydathodes before reaching the blade margins. References Brown, D. F. M. 1964. A monographic study of the fern genus Woodsia. Nova Hedwigia 16: 1–154; Taylor, T. M. C. 1947. New species and combinations in Woodsia section Perrinia. Amer. Fern J. 37: 84–88; Windham, M. D. 1987. [Abstract] Chromosomal and electrophoretic studies of the genus Woodsia in North America. Amer. J. Bot. 74: 715.

Key to the Mexican Species of Woodsia 1. Blades linear-oblong, strongly tapered proximally; pinna pairs (12–)20–50. 2. Indusia sac-like, the lobes enclosing the sori, rarely open; segment margins undifferentiated. ............................................................. 5. W. mollis. 2. Indusia of separate plates; segment margins strongly differentiated, membranous. ........................................................................... 1. W. canescens. 1. Blades lanceolate to ovate, rarely linear-oblong, generally not strongly tapered proximally; pinna pairs 5–18. 3. Indusia composed of narrow, usually filamentous segments, these uniseriate for most of their length. 4. Translucent projections on pinnule margins mostly 1–2 celled, occasionally filamentous; spores averaging 48–57 ␮m; largest pinnae with 3–7 pairs of closely spaced pinnules. ........................................................................................................................ 6. W. neomexicana. 4. Translucent projections on pinnule margins mostly multicellular, often prolonged to form twisted filaments; spores averaging 33–48 ␮m; largest pinnae usually with 7–18 (rarely as few as 3) pairs of discrete, widely spaced pinnules. .............................. 7. W. phillipsii. 3. Indusia composed of relatively broad segments, these multiseriate for most of their length but often branched, divided, or fragmenting. 5. Blades glabrous; segment margins lustrous, with cells elongated and directed toward teeth tips, terminating in sharp conical translucent teeth. ............................................................................................................................................................................... 3. W. cystopteroides. 5. Blades sparsely to densely glandular; segment margins dull, or only rarely lustrous, cells not elongated and directed toward teeth tips. 6. Proximal portion of mature stipes reddish brown or dark purple; blades densely glandular, often somewhat viscid; vein tips not enlarged, barely visible on adaxial surfaces. .................................................................................................................. 8. W. plummerae. 6. Proximal portion of mature stipes light brown or stramineous (sometimes darker at very base); blades sparsely to moderately glandular, rarely viscid; vein tips usually enlarged to form whitish hydathodes on adaxial surfaces. 7. Blades sparsely to moderately glandular; segment margins thickened and lustrous; BCS, Son, Chih, Sin, Dgo. ....... 2. W. cochisensis. 7. Blades heavily glandular; segment margins dull; Coah, Qro, Zac, SLP, Me´x, Hgo, Ver. ................................................. 4. W. mexicana.

1. WOODSIA CANESCENS Woodsia canescens (Kunze) Mett., Ann. Sci. Nat. Bot, se´r. 5, 2: 249. 1864. Fig. 324L. Cheilanthes canescens Kunze, Linnaea 13: 143. 1839. Physematium canescens (Kunze) Trevis., Nuovo Giorn. Bot. Ital. 7: 155. 1875. Syntypes. Mexico. “Ad Polverillos,” Schiede s.n.; “ad San Pedro y San Pablo,” Ehrenberg s.n. (both presumably LZ, destroyed; isosyntypes of both at B).

Similar to Woodsia mollis in frond form and dissection, but distinct in having differentiated pinna margins resembling false indusia, the sori open and lacking sac-like indusia.

Distribution. Dry grassy oak slopes with Agave and Opuntia; 2100–2900 m. Mexico.

688

WOODSIA

Specimens Examined. DF (Lyonnet 235, NY; Schaffner 79, in 1875, NY, P). Hgo (Medina C. 944d, NY; Ventura A. 355, NY, 1613, NY). Me´x (Lo´pez H. 9, NY; Lott & Wendt P-95, NY; Rzedowski 29259, ARIZ). Pue (Purpus 4021, NY; Rzedowski 24905, MICH). Oax (Mickel 6808, NY). SLP (Schaffner 67, P).

Woodsia canescens is part of the W. mollis complex but seems sufficiently distinct to be recognized at the species rank. See W. mollis for further discussion. Although we have not seen type material of W. canescens (both isosyntypes at B on loan), the protologue states “margine reflexo crenato-sinuatis” and “indusiis marginalibus, hyalinis,” suggesting that this name applies to this species with strongly differentiated, recurved segment margins. The drawing in Kunze’s Farrnkra¨uter (1: t. 35, 1840), along with a lengthy description, confirms this conclusion. In making the combination in Woodsia, Mettenius mistakenly applied the name to Colombian material.

2. WOODSIA COCHISENSIS Woodsia cochisensis Windham, Contr. Univ. Michigan Herb. 19: 54. 1993. Type. U.S.A. Arizona: Cochise Co., SE wall of Huachuca Canyon in the Huachuca Mts, ca. 2.85 km SE of Blacktail Spring, Windham 781 (UT!; isotypes ARIZ!, ASU!, MICH!, UC!, US!). Figs. 325L–O, 326J, K. Rhizomes horizontal to ascending, compact, 2–3 mm diam.; rhizome scales linear-lanceolate, bicolorous orange-tan with black central streak, entire; fronds 7–24 cm, clumped; stipes 1⁄5–2⁄5 the frond length, stramineous, sparsely scaly and sparsely to moderately glandular; blades narrowly lanceolate, proximal pinnae slightly reduced, pinnate-pinnatifid to bipinnate, 2.5–4.5 cm wide; pinnae 7–12 pairs, deltate to narrowly so, attenuate, obtusely lobed, teeth with thickened shiny margins; margins with cilia and 1–2-celled translucent projections; indusia of relatively broad segments, these multiseriate much of their length but fimbriate distally, only slightly surpassing the mature sporangia; spores 37–53 ␮m; 2n⫽152 (USA).

loch 1228, MSC; Pringle 834, F, NY p.p., PH p.p.). Dgo (Pennell 18405, PH). Sin (Correll & Correll 28869, LL). Son (Gentry 8265, MICH; Gentry et al. 19331, NY; White 4084 MICH).

Woodsia cochisensis is distinct by being sparsely to moderately glandular and has thickened lustrous blade margins with some short-crustose projections on the teeth. It differs from W. phillipsii and W. neomexicana by having indusial segments broad and being non-filamentous at bases. Woodsia cochisensis is less glandular than W. mexicana and crosses with W. phillipsii to give intermediate triploids in Arizona, Sonora, Coahuila, San Luis Potosı´, and perhaps Chihuahua.

3. WOODSIA CYSTOPTEROIDES Woodsia cystopteroides Windham & Mickel, sp. nov. Type. Mexico. Sinaloa: Ocurahui, Sierra Surotato, Gentry 6432 (holotype NY!; isotypes ARIZ!, GH!, MICH!, PH!). Figs. 325F–H, J, K. Inter species Woodsiae laminis glabris marginibus nitidis cum dentibus conicis translucentibus distincta. (Alluding to its close resemblance to plants of Cystopteris.)

Rhizomes horizontal to ascending, compact, 3–4 mm diam.; rhizome scales linear-lanceolate, bicolorous orange-tan with black central streak, entire; fronds 12–22(–30) cm, clumped; stipes 1⁄4– 1 ⁄3 the frond length, stramineous, with scattered linear-lanceolate scales at bases; blades bipinnate to bipinnate-pinnatifid, 3–5.5 (–8) cm wide, lanceolate, proximal pinnae reduced; rachises glabrous except for occasional scales 1 mm long; pinnae 6–14(–17) pairs, narrowly deltate, pinnules obtuse; margins coarsely toothed, the teeth firm, lustrous, and slightly inrolled; adaxial blade surfaces with lustrous teeth having elongate cells directed toward teeth tips; abaxial blade surfaces glabrous; indusial segments plate-like half their length, divided into filaments distally; spores 37–48 ␮m.

Distribution. Among rocks or on grassy slopes in pine or oak forests; 900–3100 m. USA (Ariz); Mexico.

Distribution. Rocky slopes in pine-oak forests; 1450–2200 m. Mexico.

Selected Specimens Examined. BCS (Brandegee s.n., 18 Oct 1890, NY; Breedlove 43316, CAS). Chih (Correll & Gentry 22797, MSC; Knob-

Specimens Examined. Dgo (Reeder & Reeder 250, MEXU). Sin (Breedlove 16905, 18465, NY). Son (Van Devender 96-598, MEXU).

WOODSIA MOLLIS

Fronds of Woodsia cystopteroides have stipe bases stramineous (though may be dark at attachment), are 2-pinnate, nearly glabrous, with lustrous margins having elongate cells directed toward teeth tips, smooth, lustrous segment margins terminating in sharp conical translucent teeth, and indusial segments united proximally over half their length, divided into filaments distally.

4. WOODSIA MEXICANA Woodsia mexicana Fe´e, Me´m. Foug. 7: 66. 1857. Figs. 326E–H. Woodsia pusilla E. Fourn. var. mexicana (Fe´e) T. M. C. Taylor, Amer. Fern J. 37: 86. 1947. Type. Mexico. Distrito Federal: prope San Angel, Schaffner 306, in 1855 (RB). Woodsia pusilla E. Fourn., Bull. Soc. Bot. France 27: 329. 1880. Type. Mexico. San Luis Potosı´: San Miguelito, Schaffner 68, in 1877 (P!).

Rhizomes horizontal to ascending, compact, 2–3 mm diam.; rhizome scales bicolorous orange-tan with black central stripe, linear-lanceolate, entire; fronds 6–18 cm, clumped; stipes 1⁄6–1⁄3 the frond length, stramineous, with glandular hairs; blades pinnate-pinnatifid to bipinnate, 2.5–5.5 cm wide, lanceolate to narrowly deltate, basal pinna pair slightly reduced; pinnae 5–10 pairs, oblong, obtuse, lobes coarsely dentate, the teeth with hair extensions, indument of glandular hairs; indusial lobes as platelike at bases, filamentous at tips, conspicuously surpassing the sporangia; spores 42–53 ␮m.

689

Ver (Seaton 280 F, GH, cited by Palacios-Rios, 1992, but specimens are W. mollis vel aff.).

Woodsia mexicana is thought to be derived from past hybridization between W. phillipsii and W. plummerae. The plants are relatively small but have large segments/lobes, pale stipes, and are heavily glandular. Some teeth have crustose projections, and the indusial segments are united proximally, filamentous distally. The species has a distinct range and little overlap with its presumed parents.

5. WOODSIA MOLLIS Woodsia mollis (Kaulf.) J. Sm., J. Bot. (Hooker) 4: 191. 1841. Figs. 324A–H, J, K, M. Physematium molle Kaulf., Flora 12: 341. 1829. Type. Mexico. s. coll. s.n. (LE?). Woodsia guatemalensis Hook., Sp. Fil. 1: 60, t. 21A. 1844. Type. Guatemala. Skinner s.n. (K!). Woodsia fragilis Liebm., Mexic. Bregn. 280 (reprint 128). 1849. Syntypes. Mexico. Veracruz: Pico de Orizaba, above Santa Marı´a Alpatlahua, Liebmann s.n. [Pl. Mex. 2606, Fl. Mex. 502, 503] (C!; isosyntypes GH!, K!).

Rhizomes horizontal to ascending, compact; rhizome scales bicolorous, light brown with black central stripe, lanceolate, ciliate; fronds (12–)20–45(–68) cm, clumped; stipes 2–8 cm long, 1⁄8– 1 ⁄4 the frond length, stramineous, with scattered scales and catenate hairs; blades 9–20(–60) ⫻ 2–7 cm, linear-elliptic, pinnate-pinnatifid, proximal pinnae greatly reduced; rachises stramineous with scattered to dense, catenate hairs, 0.5–1 mm long; pinnae (12–)20–50 pairs, narrowly deltate, the lobes obtuse, margins entire to ciliate, with catenate, lax, white hairs on both surfaces 1–1.5 mm long, and short, spreading, golden, glandular hairs 0.5 mm long present or absent; sori submarginal, 2–3 pairs per ultimate segment; indusia (0.8–)1 mm diam., cup-like, globose or dissected into several plates with lacerate margins, glabrous to glandular; sporangia 32-spored; spores 51–70 ␮m.

Distribution. Limestone (less commonly igneous) rocks in pineoak forests; 1250–2650 m. Mexico. Selected Specimens Examined. Coah (Chiang et al. 10089A, LL; Johnston et al. 11505B, TEX). Hgo (Copeland herb. 36, GH, MEXU, MICH; Sa´nchez Mejorada & Cha´vez Avies 720, MEXU, UAMIZ). Me´x (Pringle 5273, MEXU). Qro (Zamudio & Pe´rez 7463, IEB). SLP (Orcutt 1650, MEX, TEX; Parry & Palmer 1000, GH, NY, PH p.p.; Whiting 1077, ARIZ, GH, MICH). Zac (Johnston et al. 10479C, LL). Unverified, Doubtful, or Mistaken Reports. Ags (Siqueiros 4344, Gonza´lez-Adame 594, both HUAA, cited by Siqueiros-Delgado & Gonza´lez-Adame, 2004, but not verified and doubtful). Chih (reported by Knobloch & Correll, 1962, but figures and specimens are W. phillipsii). Gto (Rzedowski 42021, IEB, cited by Dı´az-Barriga & Palacios-Rios, 1992, but not verified and very doubtful). Jal (Villa 005, CHAPA, NY, cited by Mickel, 1992, but the plant is W. phillipsii).

Distribution. Terrestrial on mesic, lightly wooded rocky slopes; 800–3100 m. Mexico; Guat, Salv, Nic. Selected Specimens Examined. Ags (Siqueiros D. 4198, IEB). Chih (Knobloch 1882, MSC). Chis (Matuda 2447, NY). Col (McVaugh 16109, MICH). DF (Orcutt 3621, NY). Dgo (Gentry & Argu¨elles 18165, NY).

690

WOODSIA

Gro (Keil & Canne 9110, NY). Gto (Duges s.n., in 1880, NY). Hgo (Ventura A. 1613, NY). Jal (McVaugh 13484, NY). Me´x (Hinton 1256, NY). Mich (Leavenworth 714, NY). Mor (Rose & Painter 6874, NY). Nay (Feddema 260, NY). NL (Hinton et al. 22151, NY, TEX). Oax (Mickel 6015, NY). Pue (Purpus 6445, NY). Qro (Ferna´ndez N. 4156, NY). Sin (Gentry 6499, NY). Tlax (Weber 729, MICH). Ver (Ventura A. 64, ARIZ, MICH). Zac (Rzedowski & McVaugh 948A, MICH).

3077, NY; Fisher 35409, NY; Hinton et al. 13356, NY; Mickel 675, NY; Purpus 1607, NY). Oax (Mickel 7435, NY).

Unverified, Doubtful, or Mistaken Reports. SLP (Schaffner 957, GH, cited by Brown, 1964, but not verified).

Woodsia neomexicana Windham, Contr. Univ. Michigan Herb. 19: 52. 1993. Type. U.S.A. New Mexico: Socorro Co., along small tributary of Water Canyon in the Magdalena Mts. ca. 5.84 km SE of the summit of North Baldy, Windham 90-365 (UT!; isotypes ARIZ!, ASU!, BRY!, COLO!, GH!, MICH!, MO!, NMC!, NY!, TEX!, UC!, UNM!, US!). Figs. 326L, M.

6. WOODSIA NEOMEXICANA

Woodsia mollis is the most widespread member of the genus in Mexico; other species are mostly limited to the northern and western states. The Mexican woodsias are in great need of biosystematic study, especially the W. mollis complex involving W. montevidensis (Spreng.) Hieron. of Hispaniola, South America, and South Africa. Mexican material of Woodsia mollis is fairly consistent in general aspect and rhizome scales but varies greatly in indusial dissection, and quantity and quality of indument. The nature of the indusium is one of the major characters used in defining species in Woodsia, but Mexican material of W. mollis has indusia varying from commonly globose and sac-like (most commonly) to deeply divided into plates, as well as numerous intermediates. Small glands may be present among the catenate hairs, or hairs may be totally absent on the pinnae (fig. 324H). The glandular morphotype does not appear to be correlated with geography. The abundance of catenate hairs varies widely, and a very woolly condition is found at high elevations in the state of Me´xico, Distrito Federal, and rarely Oaxaca. Segment margins also vary from slightly to strongly recurved and distinctly differentiated, the latter a variant considered here as distinct (see W. canescens). There may be a series of species and hybrids (fertile and sterile) involved here, or these may all be expressions of a single, highly variable species, the oldest name being W. montevidensis. Material of W. montevidensis in South America has relatively few laminar hairs (mostly on segment midveins) and open sori, only rarely showing sac-like indusia. A few specimens of W. mollis from scattered localities in Mexico, Guatemala, and El Salvador have mostly open sori and few hairs. The figure of W. guatemalensis (Hooker, Sp. Fil. 1: t. 21A. 1846) shows this condition. Most specimens have large spores, 51–70 ␮m diam. One specimen with virtually no hairs and open indusia (Mu¨ller 6, NY; Fig. 324M) has spores more in the range of a diploid (33–51 ␮m; avg. 40.5 ␮m). This may represent W. montevidensis, but other specimens with open indusia are disjunct in the country. More work is needed on this difficult complex.

Specimens Examined. Coah (Marsh 599 bis, TEX; Villarreal et al. 3452, ENCB). NL (Hinton 20081, MEXU, TEX; Rodriguez L. 151, MEXU, UAMIZ, UNL, 376, UNL, 387, UAMIZ). Zac (Palmer 257, NY).

Variant Specimens Examined Orange glands. Col (McVaugh 16109, MEXU). Gro (Lorea 852, MEXU). NL (Favela Lara s.n., 12 Nov 1981, UNL, s.n., 27 Jun 1982). Sin (Breedlove 18540, NY). Ver (Seaton 280, GH). Open indusia. Mor (Riba 1921, MEXU; Valdes et al. 87, UAMIZ). Qro (Zamudio & Pe´rez 7463, MEXU). Ver (Mu¨ller 6, NY; Riba & Pe´rez 1005, UAM-1). Woolly. DF (Pringle 15020, PH). Me´x (Dziekanowski et al.

Woodsia neomexicana is thought to be derived from hybridization between W. oregana and W. phillipsii and has intermediate blade dissection (Windham, 1993). Plants are only slightly glandular, the blade margins are somewhat crustose, always with some projections on teeth, and the indusia are filamentous— less obviously than in W. phillipsii—with filaments not far exceeding the sporangia. Woodsia neomexicana apparently backcrosses to W. phillipsii. Woodsia neomexicana has a range similar to W. mexicana but

Rhizomes horizontal to ascending, compact, 2–3 mm diam.; rhizome scales bicolorous orange-tan with black central stripe, linear-lanceolate, entire; fronds 4–30 cm, clumped; stipes 1⁄5–1⁄3 the frond length, stramineous to light brown, with scales at bases, sparsely glandular throughout; blades pinnate-pinnatifid to bipinnate, 1.5–4.5 cm wide, narrowly oblong, widest at or just above middle of blades; pinnae 6–11 pairs, oblong, mostly obtuse, margins with sparse short-glandular hairs; abaxial blade surfaces nearly glabrous; indusia segments plate-like at bases with filamentous tips equalling or somewhat exceeding the sporangia, with widely scattered glands and 1–2-celled translucent projections on most teeth; spores 48–57 ␮m; 2n⫽152 (Ariz).

Distribution. Epipetric (rarely terrestrial), pine-oak-fir forests, oak forests; 2100–2600 m. Sw USA; Mexico.

WOODSIA PLUMMERAE

is distinct in its completely filamentous indusial segments and fewer glands.

7. WOODSIA PHILLIPSII Woodsia phillipsii Windham, Contr. Univ. Michigan Herb. 19: 50. 1993. Type. U.S.A. Arizona: Cochise Co., Rucker Canyon, Chiricahua Mts, Phillips 2854 (GH!; isotypes ARIZ!, ASC!, UBC!, US!). Figs. 326A–D. Rhizomes horizontal to ascending, compact, 2–3 mm diam.; rhizome scales bicolorous orange-tan with black central stripe, linear-lanceolate, entire; fronds 6–30 cm, clumped; stipes (1⁄10– )1⁄5–1⁄3 the frond length, stramineous, often dark at attachment, with sparse small scales and glands; blades pinnate-pinnatifid to bipinnate, 2–7 cm wide, linear to narrowly oblong, proximal pinnae somewhat reduced; pinnae 10–18 pairs, lobes obtuse; margins ciliate; vein endings impressed; segment margins somewhat thickened, often lustrous adaxially, sparsely glandular, appearing ciliate due to the presence of multicellular translucent projections on teeth that are often prolonged to form twisted filaments; indusial lobes filamentous most of their length, far exceeding the mature sporangia, making abaxial surface and margin appear long-ciliate; spores 33–48(–53) ␮m; 2n⫽76 (USA).

Distribution. Among rocks and on ledges, usually granitic or volcanic substrates (occasionally limestone) in open pine-oak forests; (1000–)1800–3125 m. USA (Ariz, NM, Tex); Mexico. Selected Specimens Examined. BCN (Thorne 61149, MEXU, UC). BCS (Leo´n 665, ENCB). Chih (Correll & Gentry 23006, F, LL, MO, US; Palmer 448, NY). Coah (Henrickson 15180, LL). Dgo (Palmer 357, NY). Jal (Villa Castillo 005, CHAPA, NY). Me´x (Tejero-Dı´ez 2484, IZTA). Mich (Dı´az Barriga 7423, NY). Son (Phillips 675, MICH).

Woodsia phillipsii is distinct in having stramineous stipe bases (though may be dark at attachment), very narrow blades (like W. mollis but with smaller segments and fewer pinnae), crustose margins on segment teeth (sometimes flattened and ending in trichome-like filaments), and indusial lobes filamentous most of their length and far exceeding the sporangia. Glands are variable (few to many), and the segment margins are shiny.

691

Some specimens with larger spores (40–53 vs. 33–48 ␮m) may be allotetraploid hybrids.

8. WOODSIA PLUMMERAE Woodsia plummerae Lemmon, Bot. Gaz. 7: 6. 1882. Figs. 325A–E. Woodsia obtusa (Spreng.) Torr. var. glandulosa D. C. Eaton & Faxon, Bull. Torrey Bot. Club 9: 50. 1882 (renaming of W. plummerae). Woodsia obtusa var. plummerae (Lemmon) Maxon, Proc. U. S. Natl. Mus. 23: 644. 1901. Woodsia pusilla E. Fourn. var. glandulosa (D. C. Eaton & M. Faxon) T. M. C. Taylor, Amer. Fern J. 37: 86. 1947. Type. U.S.A. Arizona: Chiricahua Mts, 24 Sep 1881, Lemmon & Lemmon s.n. (UC!; isotypes F, GH, K, US).

Rhizomes horizontal to ascending, compact, 2–3 mm diam.; rhizome scales bicolorous orange-tan with black central stripe, linear-lanceolate, entire; fronds 10–25(–35) cm, clumped; stipes 1 ⁄5–2⁄5 the frond length, stramineous, dark at bases, extremely glandular throughout; blades pinnate-pinnatifid to bipinnate, 1.5–6 cm wide, lanceolate to narrowly oblong, basal pinna pair somewhat reduced; pinnae 6–8(–12) pairs, narrowly deltate or oblong, lobes acute; segment margins toothed, ciliate or glandular-ciliate; indusial lobes plate-like, only slightly fimbriate at tips; spores 35–55 ␮m; 2n⫽152 (USA).

Distribution. Damp ledges of igneous and limestone rocks in open pine-oak or fir forests; 1500–2200(–2800) m. Sw USA; Mexico. Selected Specimens Examined. BCN (Orcutt 306, MICH, NY, PH; Wiggins 16651, ENCB). BCS (Moran 20432, MSC). Chih (Knobloch 857, MSC; Pringle 455, F p.p., GH, MICH, NY, 834 GH, MEXU, MSC, NY, PH; Wendt et al. 9972, LL, NY). Jal (McVaugh 20534, CAS, MEXU, MICH, NY). Son (Lloyd 497, GH; Wiggins 7401, GH, MICH, UC).

Woodsia plummerae is distinct in being extremely glandular, the glands often expanded at tip and yellowish. The adaxial surfaces are dull and have no projections. The stipe bases are dark for about 1 cm or more, and indusial segments few, wide, entire, and with few projections. The diploid form is rare, most plants being tetraploids. Whether they are allo- or autotetraploids is

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unknown. Woodsia plummerae crosses with W. phillipsii to form sterile intermediates.

1 2 4. W O O D W A R DI A Woodwardia Sm., Me´m. Acad. Roy. Sci. (Turin) 5: 411. 1793. Lectotype (first chosen by J. Sm., Hist. Fil. 310. 1875): Woodwardia radicans (L.) Sm. [⬅ Blechnum radicans L.] Anchistea C. Presl, Epimel. Bot. 71. 1849 [1851]. Type: Anchistea virginica (L.) C. Presl [⬅ Blechnum virginicum L.] ⬅ Woodardia virginica (L.) sm.

Terrestrial; rhizomes commonly short, stout, compact, ascending to suberect (long-creeping in some species of swampy areas), often with abundant, large, entire, nearly concolorous scales; fronds medium-sized to large, firm, evergreen, chartaceous to subcoriaceous, monomorphic; stipes stramineous to tan, scaly at least at base and sometimes distally; blades pinnate-pinnatifid to bipinnate, 1-pinnate-bipinnatifid, or bipinnatifid, margins entire to spinulose; vegetative buds on blades present or absent (ours); rachises and costae scaly to glabrescent abaxially; veins netted in both fertile and sterile blades, without included free veinlets, forming a regular series of areoles along costae and costules, otherwise free or netted toward the margin; sori appearing costal (or costular) on each side of the costae (and/or costules), in chain-like rows or sometimes appearing confluent and longlinear, borne on the outer arc of costal areoles, protected by

indusia attached just outside of the sorus (opening toward the costae); spores bilateral, with perine irregularly folded; x⫽34 (35 in the segregate Anchistea). Woodwardia is a largely north-temperate and amphioceanic genus of 13 species, roughly divided between the Old World and New World, mostly in eastern Asia and United States/Mexico, plus one on Atlantic Ocean islands. The species in neotropical areas occur at middle to high elevations, usually above 1500 m. Woodwardia has blechnoid affinities and is basal in the family Blechnaceae (Cranfill, 2001), differing from Blechnum in the interrupted costal sori and more strongly areolate venation. The generic description above does not include the dimorphic segregate Lorinseria areolata (L.) C. Presl (Woodwardia areolata (L.) T. Moore) of the eastern United States, but does include Woodwardia virginica (L.) Sm., often segregated as Anchistea virginica (L.) C. Presl; the latter is a rather isolated element in the genus and is confined to the eastern United States and southeastern Canada, primarily in the coastal plain. References Cranfill, R. B. 2001. Monograph of Anchistea, Lorinseria, and Woodwardia (Blechnaceae). Chapter 2, Ph.D. dissertation, Phylogenetic studies in the Polypodiales (Pteridophyta) with an emphasis on the family Blechnaceae, University of California, Berkeley; Maxon, W. R. 1919. Notes on American ferns. XIV. Amer. Fern J. 9: 67–73; Weatherby, C. A. 1949. Two Mexican ferns. Amer. Fern J. 39: 88–91.

Key to the Mexican Species of Woodwardia 1. Free pinnae 5(–6) pairs per frond, or fewer; sori borne along the costae and rachises greatly elongate and often confluent at maturity, superficial or slightly impressed, running nearly the entire length of the costae; rhizome scales 5–10 mm long. .......... 2. W. martinezii. 1. Free pinnae 6 or more pairs per frond; sori absent from along costae or, when present, usually remaining discrete with age, never running nearly the entire length of the costae; rhizome scales 10–30 mm long. 2. Sori borne along distal 1⁄3–1⁄2(–4⁄5) of costae, as well as along costules; sori impressed to superficial (adaxial surface slight embossed), indusia thick to thin; distal pinnae often subcordate at their bases; rhizome and stipe base scales 10–20 mm long. ................................................................................................................................................................................................................... 3. W. semicordata.

2. Sori borne only along the costules of the ultimate segments, 1–5 mm long, absent from along costae; sori deeply impressed (the adaxial surface embossed), indusia thick; distal pinnae cordate to short- or long-decurrent at their bases; rhizome and stipe base scales 15–30 mm long. 3. Costae, costules, and veins abaxially glabrous or glabrescent, or with a few scattered and stramineous to whitish narrowly lanceolate scales, also often with yellowish resinous glands and sometimes short-stipitate glands; pinna segments often strongly ascending and falcate; fronds held mostly erect to stiffly ascending; proximal pinnae reduced and distant from pair above. ...................................................................................................................................................................................................... 1. W. fimbriata. 3. Costae, costules, and veins abaxially with scattered to often numerous scales, these reddish brown, those of the costae ovatelanceolate, glands lacking; pinna segments spreading to slightly ascending, not or somewhat falcate; fronds not held erect, mostly lax, often drooping, with proximal pinnae not markedly reduced, approximate to pair above. ................................. 4. W. spinulosa.

1. WOODWARDIA FIMBRIATA

Woodwardia chamissoi Brack., U.S. Expl. Exped., Filic. 16: 138. 1854. Lectotype (chosen here, informally selected by Cranfill, 2001: 218). U.S.A. California: Sacramento River, Brackenridge s.n. (US 62650, photo UC!).

Woodwardia fimbriata Sm. in Rees, Cycl. 38. 1818. Type. “West coast of North America” (protologue), “New Georgia” [probably U.S.A. Washington: Puget Sound area, in litt. from Eric Groves] at BM, Menzies s.n. (LINN 1628.8-Smith Herb., photo in Philipp. J. Sci. 50: 175–183. 1933; isotypes BM, E-herb. Menzies, LIV). Figs. 328C, D.

Rhizomes stout, short-creeping to suberect; fronds mostly erect to ascending; stipes (18–)30–60(–72) cm long, abundantly scaly at base, the scales brownish to orange-brown; blades pinnatepinnatifid, 25–100 cm long, broadest at or just below the middle, distal pinnae subcordate at the base; rachises sparsely scaly to usually glabrescent; pinnae (8–)10–24 pairs, the proximal pair distant from the rest, the lower and middle ones mostly 15–25

WOODWARDIA MARTINEZII ⫻ 3.5–9 cm, pinnatifid, asymmetric at base by the reduction of the basalmost basiscopic lobe(s); pinna lobes broadly attached at base, narrowed gradually distally or with margins parallel for some distance to the acute to obtuse apex, frequently with margins of proximal pinnae crenate to deeply incised, the lobes spinulose along the margins; indument abaxially of sparse fibrillose scales less than 0.5 mm long on costules, veins, and leaf tissue, also of sparse, stramineous to whitish, linear-lanceolate scales to ca. 2 mm along costae, these often deciduous, laminae abaxially often with yellowish resinous glands and short-stipitate glands; sori usually confined to costular areoles, short-linear, (1–)2–5 mm long, mostly 2–4 times as long as broad, straight or curved, deeply sunken into the blade tissue; indusia cartilaginous and vaulted, persisting and retaining form after sporangial dehiscence; spores 64 per sporangium; 2n⫽68 (Calif).

693

Rhizomes short to long-creeping, with pale brown scales 5–10 1–1.5 mm; fronds mostly erect to ascending; stipes 25–60 cm long, scaly at base, the scales orangish brown; blades pinnatepinnatifid, 20–42 cm long, deltate to broadly ovate; rachises with scattered, often deciduous, fibrillose scales; pinnae 1–5 pairs (usually 2–4 pairs completely free), the lower and longest ones 13– 23 ⫻ 5–11 cm, free pinnae cuneate at base, pinnatifid, sharply asymmetric at base by the reduction of the basalmost basiscopic lobe(s); pinna lobes broadly attached at base, attenuate to rounded at apex, or frequently with margins of proximal pinnae crenate to deeply incised, spinulose along the margins; indument abaxially absent or of sparse fibrillose scales 0.1–0.2 mm long on costules, veins, and leaf tissue, costae lacking scales, laminar glands absent; sori along costules, entire length of costae, and along rachises of the pinnatifid apex, varying greatly in length, linear, 3–40⫹ mm long, becoming confluent upon dehiscence of sporangia, superficial on the blade tissue, the lamina not or only faintly embossed; indusia thin, erose, spreading and hidden by dehiscing sporangia, retaining configuration after dehiscence of sporangia; spores 32 per sporangium?

⫻

Distribution. Terrestrial, along streambanks and springs; 400– 2100 m. Canada (British Columbia, especially Vancouver Is), USA (Ariz, Calif, Nev, Ore, Wash); Mexico. Selected Specimens Examined. BCN (Brandegee s.n., 4 May 1893, UC; Moran 25113, NY, 25576, UC; Wiggins 10027, 20976, DS). BCS (La Chuparosa, Brandegee s.n., 17 Oct 1893, UC). Son (Gentry 2111, UC; Martin et al. s.n., UC; Phillips 721, US). Unverified, Doubtful, or Mistaken Reports. NL (Rodrı´guez 8, UANL, ENCB, cited by Rodrı´guez-Lo´pez & Arreguı´n-Sa´nchez, 1992, but the specimen is likely W. spinulosa).

This species is difficult to separate from W. spinulosa, and some of the specimens cited are somewhat intermediate between W. spinulosa, as it occurs in southern and Central Mexico, and W. fimbriata, from the western United States. Characters listed by Maxon (1919) do not appear to be diagnostic. See discussion under W. spinulosa for differences with that species.

2. WOODWARDIA MARTINEZII Woodwardia martinezii Maxon ex Weath., Amer. Fern J. 39: 88. 1949. Type. Mexico. Hidalgo: near Zacualtipa´n, Martı´nez 27 (US!, photo UC!; isotypes B, BR, ENCB, LL, UC!). Figs. 328A, B.

Distribution. Terrestrial, in mesic woods, often in ravines or along streambanks, oak forests, montane cloud forests; 1200– 2000 m. Mexico. Selected Specimens Examined. Hgo (Acosta & Barrios 305, MEXU; Martı´nez 14, US; Moore 3202, UC; Puig 4873, NY). Pue (Knobloch 2173, US; Riba 446, NY; Rudd 1078, US; Ventura A. 354, NY). Ver (Ramı´rez 562, XAL).

This species is narrowly endemic and probably relatively basal in the genus (Cranfill, 2001); it may also be relictual, having once had a much wider distribution. The dissection and architecture of the blades, soral configuration, and rhizome structure all suggest relationship with some Asian species, e.g., W. japonica (L.f.) Sm., but probably not with W. kempii Numerous, as suggested by Tryon and Tryon (1982). Weatherby (1949) observed that this species combined features of Woodwardia s.str. (such as 1–2 rows of areoles outside the costal ones, spinulose-serrate margins, granular spores) with features of the segregate genus Anchistea (such as the strong development of costal sori which are superficial and membranous) and that this “emphasizes the untenability of Anchistea as a genus.” Hybrids are postulated be-

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WOODWARDIA

tween W. martinezii and W. spinulosa, and these are interpreted as resulting in the next species, W. semicordata (which see).

3. WOODWARDIA SEMICORDATA Woodwardia semicordata Mickel & Beitel, Mem. New York Bot. Gard. 46: 403. 1988, pro hybr. Type. Mexico. Veracruz: Orizaba, 1855, Mu¨ller 1485 bis (NY!). Figs. 328E–G. Rhizomes stout, short to suberect, with orange-brown scales 10–20 ⫻ 1.5–2.5 mm; fronds mostly erect to ascending; stipes 12– 45 cm long, scaly at base, the scales orangish brown; blades pinnate-pinnatifid, 25–94 cm long, ovate; rachises sparsely scaly and often glabrescent; pinnae 8–12 pairs, the longest ones 10–32 ⫻ 4–15 cm, pinnatifid, asymmetric at base by the reduction of two or more basalmost basiscopic and acroscopic lobe(s); pinna lobes broadly attached at base, attenuate at apex, or frequently with margins of proximal pinnae crenate to deeply incised, serrulate along the margins; indument abaxially of sparse to often numerous, red-brown fibrillose scales 0.3–1 mm long on costules, veins, and leaf tissue, also of sparse to occasionally numerous broader ovate scales to 3 ⫻ 1.5 mm along costae, scales often deciduous, laminae lacking glands; sori along costules, often also along distal 1⁄3–1⁄2(–4⁄5) of costae of all pinnae, sometimes along rachises in distal half of pinnatifid blade apex, variable in length and sometimes spacing, linear, 3–30⫹ mm long, sometimes confluent upon dehiscence of sporangia, those along costules deeply sunken in the blade tissue; indusia somewhat cartilaginous, vaulted to merely conduplicate; spores 32 per sporangium?

Woodwardia semicordata is intermediate between W. spinulosa and W. martinezii, and most likely arose through hybridization between these two species. However, it appears that in some cases, this hybrid is stabilized, perhaps through apogamy or chromosome doubling, and hence we recognize it as a species, without the hybrid symbol. In addition, the range of W. semicordata is considerably wider than one of the putative parents, W. martinezii, indicating that it is capable of reproducing on its own. Woodwardia semicordata resembles W. spinulosa in its robust habit, more gradually tapered blades, and tendency of its sori to remain discrete even after sporangial dehiscence; however, it approaches W. martinezii in the marked asymmetry of its proximal pinnae. In features such as development and configuration of the indusia, length of sori, and general blade dissection, W. semicordata is intermediate between the suspected parental species. Because of the reduced spore output of some populations, there is the supposition that W. semicordata reproduces by virtue of an apogamous life cycle, but this needs confirmation. Some specimens are reported to have malformed spores (Mickel & Beitel, 1988), but this is apparently not always true, and raises the possibility that different populations have different origins. Unfortunately, the type is insufficient to be able to ascertain its mode of reproduction, i.e., it lacks mature spores; however, the sporangia appear collapsed and the contents necrotic and black. Several collections of this species, as well as material in a number of botanical gardens, were originally identified as W. martinezii. Figure 103.1 (labelled W. martinezii) in Tryon and Tryon (1982: 664) is most likely W. semicordata.

4. WOODWARDIA SPINULOSA Woodwardia spinulosa M. Martens & Galeotti, Me´m. Foug. Mexique 64. 1842. Type. Mexico. Veracruz: Galeotti 6255 (BR, not found). The only Galeotti specimens of this species at BR are five sheets of 6573 from Cerro San Martı´n, three from Veracruz, and one from Oaxaca (supposedly at 2000’ although we have not seen it below 6000’). The BR specimen of Galeotti 6255 is Cochlidium serrulatum, Veracruz, 8000’. Most likely the published collection number was in error and 6573 is the type number). Figs. 327A–E.

Distribution. Terrestrial in montane cloud forests, pine-oakLiquidambar forests, in woods or ravines; 1000–2400 m. Mexico; Salv, CR (Grantham & Parsons 0230–90, UC). Selected Specimens Examined. Chis (Rovirosa 1068, PH). Hgo (Luna et al. 870, FCME). Me´x (Dura´n R. 14, IZTA). Oax (Gaddis s.n. ⫽ Univ. Calif. Bot. Gard. 92.1020, UC; Mickel 903, ISC). Pue (Riba 445, NY). Ver (Halsted 9, NY; Nee 23471, 23542, NY). Unverified, Doubtful, or Mistaken Reports. Gro (Vela´zquez 321, FCME, cited by Lorea-Herna´ndez and Vela´zquez M., 1998, as this species, but not seen). NL (Meyer & Rogers 2482, BM, cited by Cranfill, 2001, but not verified). Tam (Sharp 52216, BM, cited by Cranfill, 2001, but not verified).

Rhizomes suberect to erect, with scales concolorous, brownish, 15–30 ⫻ 3–5 mm; fronds arching and often drooping to suberect; stipes stramineous, (14–)20–50(–100) cm long, 1⁄2 the frond length, with scales similar to those of rhizome; blades pinnatepinnatifid, (18–)25–55(–100) cm long, 4–5 pairs of free pinnae below the pinnatifid apex, these with usually short-decurrent bases; rachises sparsely scaly and often glabrescent; pinnae 8–20 pairs, the largest (11–)20–35 ⫻ (3.5–)5–10 cm; pinna lobes 2–4.5 cm long, 1–1.5 cm wide, sometimes crenately lobed especially on the acroscopic side, margins spinulose; indument abaxially of sparse to often numerous, red-brown fibrillose or dissected scales 0.3–1 mm long on costules, veins, and leaf tissue, also of sparse to occasionally numerous, broadly ovate scales 1.5–3 ⫻ 0.8–1.5 mm, especially along costae, these often deciduous,

WOODWARDIA SPINULOSA

glands absent; sori along only the costular areoles, deeply impressed, upper surface of lamina embossed; indusia thick, vaulted, entire to suberose; spores 64 per sporangium.

695

Me´x (Purpus 1583, UC). Mich (Dı´az-Luna 7756, UC). Mor (Lyonnet 2864, DS). Nay (Dı´az-Luna 9579, UC). NL (Rodrı´guez 134, MO). Oax (Mickel 5320, UC). Pue (Arse`ne 1937, UC). SLP (Schaffner s.n., Oct 1876, NY). Sin (Sanders 22590, UC). Ver (Balls B4375, UC). Unverified, Doubtful, or Mistaken Reports. Qro (Zamudio 108, IEB, cited by Dı´az-Barriga & Palacios-Rios, 1992, but not verified; Diaz-Barriga & Carranza 7281, Rubio 1489, and Sevı´n 535, all QMEX, cited by Arreguı´n et al., 2001, but not verified). Tam (reported by Rodrı´guez-Lo´pez and Arreguı´n-Sa´nchez, 1992, as W. fimbriata, but not verified; also reported by Sharp, 1954, from Rancho del Cielo; specimens so identified are probably W. semicordata, which see).

Distribution. Terrestrial, usually on steep banks or ravines near streams in montane rain forests, pine forests, pine-oak forests, montane cloud forests dominated by alder; 1300–3100 m. Mexico; Guat, Hond, Salv, Nic. Selected Specimens Examined. Chih (Knobloch 8010, US). Chis (Alava 1236, UC). DF (Schaffner s.n., in 1875, B; Schmitz s.n., US). Dgo (Sanders 22700, UC). Gro (Lorea 2298, UC). Gto (Rzedowski 41573, ENCB). Hgo (Moore 5381, UC, US). Jal (Mexia 1515, NY, UC, US).

Woodwardia spinulosa is variable in blade and pinna size and shape, as well as width and lobing of pinnules and density of indument abaxially on the blades. It is distinguished from W. fimbriata by having fibrillose and often dissected scales on the abaxial costules, veins, and lamina, by the absence of yellowish resinous glands abaxially, and by the distal pinnae with shortdecurrent pinna bases (as opposed to cordate bases in W. fimbriata). However, in the northwestern part of the range of W. spinulosa, near the small area of overlap with W. fimbriata, these differences are not clearcut. Some specimens of Woodwardia, particularly from Sonora (see citations under W. fimbriata) and Durango (e.g., Palmer 92, NY, UC), are difficult to place with confidence, suggesting that further study is warranted. See W. semicordata, for discussion of hybrids between W. martinezii and W. spinulosa.

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ridophytes from the state of Chiapas, Mexico. Amer. Fern J. 77: 69–71. ———, ———, A. Valdes & Y. Sandoval. 1996. Pteridoflora del estado de Morelos, Me´xico. Lista de familias, generos, y especies. Acta Bot. Mex. 37: 45–65. ——— & M. A. Pe´rez-Farrera. 2000. New records for the pteridoflora of the state of Chiapas, Mexico. Amer. Fern J. 90: 104–105. ——— & B. Pe´rez-Garcı´a. 1979. Estudio bota´nico y ecolo´gico de la regio´n del Rı´o Uxpanapa, Veracruz. No. 9. Pterido´fitas. Biotica 4: 135–139. ——— & ———. 1997. Pterido´fitas. Pp. 175–181. In: E. Gonza´lez S., R. Dirzo & R. C. Vogt (eds.), Historia natural de Los Tuxtlas. Universidad Nacional Auto´noma de Me´xico, Me´xico. ——— & ———. 1999. Pterido´fitas. Familia Dryopteridaceae. Flora de Me´xico. 6(4): 1–48. Consejo Nacional de la Flora de Me´xico, Me´xico, D.F. ———, ——— & A. Orozco S. 1993. Las pterido´fitas en la historia de las plantas de la Nueva Espan˜a de Francisco Herna´ndez, protomedico Espan˜ol. Acta Bot. Mex. 25: 27– 48. Rodrı´guez P., A. 1953. Estudio de los helechos en los alrededores de Guadalajara. 68 pp. + 60 pp. + 2 pp. bibliography. Tesis, Maestro en Ciencias Biolo´gicas, Guadalajara, Jalisco. Rodrı´guez-Lo´pez, R. & M. L. Arreguı´n-Sa´nchez. 1992. Adiciones a la pteridoflora del estado de Nuevo Leo´n, Me´xico. II. Anales Esc. Nac. Ci. Biol. 37: 27–34. Rovirosa, J. N. 1889. Vida y trabajos de naturalista Belga Augusto B. (Bonifacio) Ghiesbreght, explorador de Me´xico. Naturaleza (Mexico City), Ser. 2, 1: 211–217. ———. 1909. Pteridografı´a del sur de Me´xico. iv + 298 pp., 70 pls. Ignacio Escalante, Me´xico, D. F. Rzedowski, J. 1959. Las collecciones bota´nicas de Wilhelm (Jose´ Guillermo) Schaffner en San Luis Potosı´. Acta Cientifica Potosina 3: 99–121. ———. 1973. Geographical relationships of the flora of Mexican dry regions. Pp. 61–72. In: A. Graham (ed.), Vegetation and vegetational history of northern Latin America. Elsevier Scientific Co., Amsterdam. ———. 1978. Vegetacio´n de Me´xico. 432 pp. Editorial Limusa, Me´xico, D.F. ———. 1991a. Diversidad y origenes de la flora faneroga´mica de Me´xico. Acta Bot. Mex. 14: 3–21. ———. 1991b. El endemismo en la flora faneroga´mica mexicana: una apreciacio´n analitica apreliminar. Acta Bot. Mex. 15: 47–64. ——— & R. McVaugh. 1966. La vegetacio´n de Nueva Galicia. Contr. Univ. Michigan Herb. 9: 1–123. Sano, R., M. Takamiya, M. Ito, S. Kurita & M. Hasebe. 2000. Phylogeny of the lady fern group, tribe Physematieae (Dryopteridaceae), based on chloroplast rbcL gene sequences. Molec. Phylogen. Evol. 15: 404–413. Savage, S. 1945. A catalogue of the Linnaean Herbarium. xv + 225 pp. Linnaean Society of London, London. Schelpe, E. A. C. L. E. & N. C. Anthony. 1986. Pteridophyta.

LITERATURE

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Steyermark, J. A., P. E. Berry & B. K. Holst (eds.). 1995. Flora of the Venezuelan Guayana. Vol. 2. Pteridophytes, Spermatophytes: Acanthaceae—Araceae. xiii + 706 pp. Timber Press, Portland. Stolze, R. G. 1976. Ferns and fern allies of Guatemala. Part. 1. Ophioglossaceae through Cyatheaceae. Fieldiana, Bot. 39: 1– 130. ———. 1981. Ferns and fern allies of Guatemala. Part. 2. Polypodiaceae. Fieldiana, Bot., n.s., 6: 1–522. ———. 1983. Ferns and fern allies of Guatemala. Part. 3. Marsileaceae, Salviniaceae, and the fern allies. Fieldiana, Bot., n.s., 12: 1–91. Tejero-Dı´ez, D. & M. de la L. Arreguı´n-Sa´nchez. 2004. Lista con anotaciones de los pterido´fitos del Estado de Me´xico. Acta Botanica Mexicana 69: 1–82. ——— & J. T. Mickel. 2004. Pterido´fitos de Oaxaca. In: A. Garcı´a Mendoza, M. A. Briones & M. J. Ordo´n˜ez (eds.), Biodiversidad del estado de Oaxaca, Mexico. Instituto de Biologı´a, Univ. Nacional de Me´xico, D. F. pp. 121–139. ——— & L. Pacheco. 2004. Notas taxono´micas y de distribucio´n en Polypodium plesiosorum Kunze y P. rhodopleuron Kunze (Polypodiaceae). Anales del Instituto de Biologı´a UNAM, serie Bota´nica 75(1): 11–37. Tryon, R. M. 1964b. The ferns of Peru: Polypodiaceae (Dennstaedtiaceae to Oleandraceae). Contr. Gray Herb. 194: 1–253. ———. 1970. The classification of the Cyatheaceae. Contr. Gray Herb. 200: 3–53. Tryon, R. M. & R. G. Stolze. 1989a. Pteridophyta of Peru. Part. I. 1. Ophioglossaceae- 12. Cyatheaceae. Fieldiana, Bot., n.s., 20: 1–145. ——— & ———. 1989b. Pteridophyta of Peru. Part. II. 13. Pteridaceae- 15. Dennstaedtiaceae. Fieldiana, Bot., n.s., 22: 1– 128. ——— & ———. 1991. Pteridophyta of Peru. Part. IV. 17. Dryopteridaceae. Fieldiana, Bot., n.s., 27: 1–176. ——— & ———. 1992. Pteridophyta of Peru. Part. III. 16. Thelypteridaceae. Fieldiana, Bot., n.s., 29: 1–80. ——— & ———. 1993. Pteridophyta of Peru. Part. V. 18. Aspleniaceae- 21. Polypodiaceae. Fieldiana, Bot., n.s., 32: 1– 190. ——— & ———. 1994. Pteridophyta of Peru. Part. VI. 22. Marsileaceae- 28. Isoe¨taceae. Fieldiana, Bot., n.s., 34: 1–123. ——— & A. F. Tryon. 1982. Ferns and allied plants, with special reference to tropical America. xii + 857 pp. SpringerVerlag, New York. Valde´s, J. & H. Flores. 1983. La pterido´fitas en la flora halofila y gipsofila de Me´xico. Anales Inst. Biol. Univ. Nac. Me´x. 54: 173–188. Vareschi, V. 1969. Flora de Venezuela: Helechos. 1032 pp + 207 plates. Instituto Botanico, Caracas, Venezuela. Va´zquez G., J. A., R. Cuevas G., T. S. Cochrane, H. H. Iltis, F. J. Santana M. & L. Guzma´n H. 1995. Flora de Manantla´n. Sida, Bot. Misc. 13: 1–312. Vega A., R., G. A. Bojo´rquez B. & F. Herna´ndez A. 1989. Flora de Sinaloa. 49 pp. Universidad Auto´noma de Sinaloa, Coordinacio´n General de Investigacio´n y Postgrado, Culiaca´n, Sinaloa.

702

LITERATURE

Walker, T. G. 1966. A cytotaxonomic suvey of the pteridophytes of Jamaica. Trans. Roy. Soc. Edinburgh 66: 169–237. ———. 1973. Additional cytotaxonomic notes on the pteridophytes of Jamaica. Trans. Roy. Soc. Edinburgh 69: 109– 135. ———. 1985. Cytotaxonomic studies of the ferns of Trinidad 2. The cytology and taxonomic implications. Bull. Brit. Mus. (Nat. Hist.), Bot. 13: 149–249. Whitehead, J. 1937. Some Arizona ferns collected in Sonora, Mexico. Amer. Fern J. 27: 43–51. Wiggins, I. L. 1973. Notes on three ferns from Baja California, Mexico. Amer. Fern J. 63: 152–157. ———. 1980. Pteridophytes: 52–71. Flora of Baja California. 1025 pp. Stanford University Press. Stanford. Wikstrom, N., P. Kenrick & J. C. Vogel. 2002. Schizaeaceae: a phylogenetic approach. Rev. Palaeobot. and Palynol. 119: 35–50. Windham, M. D. 1993. New taxa and nomenclature changes in the North American fern flora. Contr. Univ. Michigan Herb. 19: 31–61. ——— & G. Yatskievych. 2003. Chromosome studies of cheilanthoid ferns. I. Counts from the Americas provide new

insights on generic and subgeneric stability of base numbers. Amer. J. Bot. 90: 1788-1800. Windisch, P. G. 1982. Specimens from Fe´e’s pteridological collection at the Botanical Garden of Rio de Janeiro. Amer. Fern J. 72: 56–60. Wolf, P. G. 1995 [1996]. Phylogenetic analyses of rbcL and nuclear ribosomal RNA gene sequences in Dennstaedtiaceae. Amer. Fern J. 85: 306–327. ———, K. M. Pryer, A. R. Smith & M. Hasebe. 1998. Phylogenetic studies of extant pteridophytes. Pp. 541–556. In: D. E. Soltis et al. (eds.), Molecular systematics of plants II. DNA sequencing. Kluwer Academic Publishers, Boston. ———, S. D. Sipes, M. R. White, M. L. Martines, K. M. Pryer, A. R. Smith & K. Veda. 1999. Phylogenetic relationships of the enigmatic fern families Hymenophyllopsidaceae and Lophosoriaceae: evidence from rbcL nucleotide sequences. Pl. Syst. Evol. 219: 263–270. ———, P. S. Soltis & D. E. Soltis. 1994. Phylogenetic relationships of dennstaedtioid ferns: evidence from rbcL sequences. Molec. Phylogen. Evol. 3: 383–392. Yatabe, Y, M. Takamiya & N. Murakami. 1998. Variation in the rbcL sequence of Stegnogramma pozoi subsp. mollissima (Thelypteridaceae) in Japan. J. Pl. Res. 111: 557–564.

ACROSTICHUM

703

Fig. 1. A–C: Acrostichum danaeifolium (Rzedowski 24586a, NY). A. Fertile pinnae. B. Sterile pinna detail, abaxial surface. C. Paraphyses. D–F. Acrostichum aureum. D. (Matuda 2672, NY), stipe with ‘spines’. E, F. (Cowan 3193, NY). E. Sterile pinna detail, abaxial surface. F. Paraphyses.

704

ACTINOSTACHYS

Fig. 2. Actinostachys pennula (Martínez S. 14891, NY). A. Habit. B. Fertile pinnae.

ADENODERRIS

Fig. 3. Adenoderris sororia (Breedlove 56215, NY). A. Habit. B. Rhizome scale. C. Pinna.

705

706

ADIANTOPSIS

Fig. 4. A, B. Adiantopsis seemannii (Mickel 5214, NY). A. Habit. B. Pinnule. C, D. Adiantopsis radiata (Mickel 6970, NY). C. Frond. D. Pinnule.

ADIANTUM

707

Fig. 5. A–C. Adiantum braunii (Mickel 1650, NY). A. Pinna. B. Segment. C. Rachis. D, E. Adiantum deflectens (Mickel 5974, NY). D. Habit. E. Segment. F–H. Adiantum raddianum (Mickel 6026, NY). F. Pinna. G. Segment. H. Segment margin. J. Adiantum lunulatum (Mickel 6878, NY), habit. K, L. Adiantum concinnum (Camp 2450, NY). K. Pinna. L. Segments. M–O. Adiantum capillus-veneris (Mickel 6637, NY). M. Pinnule. N. Segment. O. Segment margin. P, Q. Adiantum raddianum (Mickel 6026, NY). P. Pinna. Q. Segment.

708

ADIANTUM

Fig. 6. A, B. Adiantum feei (Mickel 1608, NY). A. Frond. B. Segment. C, D. Adiantum tricholepis (Mickel 4215, NY). C. Pinna. D. Segment. E, F. Adiantum patens (Mickel 3957, NY). E. Blade. F. Segment. G, H. Adiantum galeottianum (Mickel 3992, NY). G. Habit. H. Bipinnate form, base of blade.

ADIANTUM

709

Fig. 7. A–C. Adiantum amplum (Mickel 5187, NY). A. Pinna. B. Rhizome and stipe base. C. Segment. D–F. Adiantum tenerum (Mickel 5818, NY). D. Pinnule. E. Rhizome and stipe bases. F. Segment. G, H, J. Adiantum andicola (Hallberg 1742, NY). G. Pinnule. H. Rhizome and stipe bases. J. Segment. K–N. Adiantum trapeziforme (Hallberg 1626, NY). K. Pinnule. L. Pinnule. M. Rhizome and stipe bases. N. Segment. O–W. Adiantum andicola. O–Q. (Mickel 804, NY). O. Pinnule. P. Rhizome and stipe bases. Q. Segment. R–T. (Mickel 1125, NY). R. Pinnule. S. Rhizome. T. Segment. U–W. (Mickel 6149, NY). U. Pinnule. V. Rhizome and stipe bases. W. Segment.

710

ADIANTUM

Fig. 8. A–C. Adiantum wilesianum (Mickel 5788, NY). A. Pinna. B. Segments. C. Rachilla. D. Adiantum latifolium (Mickel 5857, NY), segments. E. Adiantum tetraphyllum (Hallberg 1642, NY), segments. F. Adiantum pulverulentum × villosum (Mickel 5777, NY), pinna. G. Adiantum pulverulentum (Hallberg 1623, NY), pinna. H. Adiantum petiolatum (Hallberg 1621, NY), pinnae (see also Fig. 10D). J. Adiantum macrophyllum (Hallberg 1650, NY), pinnae. K–M. Adiantum amblyopteridium (Mickel 5163, NY). K. Pinna. L. Rachis. M. Segment base.

ADIANTUM

Fig. 9. A–D. Adiantum mcvaughii (McVaugh 19018, MICH). A. Habit. B. Sterile segment. C. Fertile segment. D. Stipe scale.

711

712

ADIANTUM

Fig. 10. A–C. Adiantum obliquum A. (Hernández G. 1828, NY), habit. B, C. (Hernández G. 249, NY). B. Pinna detail. C. Stipe scale. D. Adiantum petiolatum (Hallberg 1621, NY), pinna (see also Fig. 8H). E, F. Adiantum poiretii (Mickel 3918, NY). E. Pinnule. F. Segment. G, H, J. Adiantum aleuticum (Pinkava 11677, NY, from Arizona). G. Proximal pinna. H. Segment detail. J. Sterile segment margin. K–M. Adiantum jordanii (Wiggins 7541, NY). K. Fertile pinna. L. Fertile segment. M. Sterile segment.

ADIANTUM

713

Fig. 11. A–C. Adiantum shepherdii (Hinton 15434, NY). A. Habit. B. Pinna. C. Rhizome scale. D–G. Adiantum caryotideum (Breedlove 34205, DS). D. Pinna. E. Rachis. F. Rachis scale. G. Segment detail showing idioblasts. H, J, K. Adiantum oaxacanum (Mickel 5804, NY). H. Pinna. J. Segment detail. K. Rachis scale. L, M. Adiantum villosum (Hallberg 1661, NY). L. Proximal pinna. M. Rachis scale.

714

ADIANTUM

Fig. 12. A–D. Adiantum wilsonii (Matuda 3616, LL). A. Habit. B. Rachis. C. Venation. D. Rhizome scale. E–G. Adiantum seemannii (Williams 885, NY, from Panama). E. Fertile pinna. F. Sterile pinna. G. Rhizome scale. H, J, K. Adiantum decoratum (Breedlove 33179, LL). H. Pinna. J. Fertile segment. K. Rachis scale. L, M. Adiantum terminatum (Schipp 351, NY, from Belize). L. Segment detail. M. Rachis hair. N–P. Adiantum trichochlaenum (Mickel 6975, NY). N. Pinna. O. Rachis. P. Segment detail.

ALSOPHIL A

715

Fig 13. A–E. Alsophila salvinii (Mickel 4318, NY). A. Pinnules. B. Stipe. C. Stipe scale. D. Segment detail. E. Costular scales. F–H, J–L. Alsophila firma (Mickel 1010, ISC). F. Pinnules. G. Stipe. H. Stipe scale. J. Segment detail. K, L. Costular scales. M–R. Alsophila tryoniana. M–P. (Sousa 3604, MEXU). M. Stipe. N. Stipe scale. O. Segment detail, adaxial surface. P. Segment detail, abaxial surface. Q–R. (Steyermark 30009, US, from Guatemala). Q. Segment detail, abaxial surface. R. Blade scales.

716

ANANTHACORUS

Fig. 14. A–D. Ananthacorus angustifolius (Hallberg 1277, NY). A. Habit. B. Blade detail. C. Venation. D. Rhizome scale.

ANEMIA

717

Fig. 15. A, B. Anemia muenchii (Mickel 6874, NY). A. Habit. B. Pinna base. C, D. Anemia adiantifolia (Mickel 6292, NY). C. Habit. D. Segment. E. Anemia adiantifolia × mexicana var. makrinii (Mickel 1356, NY), pinna. F. Anemia adiantifolia × mexicana var. mexicana (Mickel 616, NY), pinna.

718

ANEMIA

Fig. 16. A, B. Anemia karwinskyana (Mickel 3993, NY). A. Habit. B. Segment detail. C, D. Anemia tomentosa var. mexicana (Mickel 4209, NY). C. Habit. D. Segment detail.

ANEMIA

719

Fig. 17. A, B. Anemia mexicana var. makrinii (McVaugh 16056, NY). A. Habit. B. Pinna detail. C, D. Anemia cicutaria (Britton et al. 15315, NY, from Cuba). C. Habit. D. Segment. E. Anemia mexicana var. mexicana (Pringle 1976, NY), blade base. F. Anemia speciosa (Sharp s.n., 29 Oct 1960, NY), frond.

720

ANEMIA

Fig. 18. A. Anemia hirsuta (Mickel 4222, NY), habit. B. Anemia oblongifolia (Mickel 3998, NY), habit. C. Anemia familiaris (Mickel 5984, NY), habit. D. Anemia pastinacaria (Mickel 4206, NY), habit.

ANEMIA

721

Fig. 19. A–C. Anemia affinis (Reveal 4048, NY). A. Habit. B. Pinna base, adaxial surface. C. Pinna base, abaxial surface. D. Anemia recondita (Mickel 1703, NY), habit. E. Anemia multiplex (Mickel 1709, NY), habit. F. Anemia jaliscana var. jaliscana (McVaugh 16416, NY), habit.

722

ANEMIA

Fig. 20. A, B. Anemia ×paraphyllitidis (Mickel 1118, NY). A. Habit. B. Pinna detail. C, D. Anemia phyllitidis (Mickel 6178, NY). C. Habit. D. Pinna detail. E, F. Anemia semihirsuta (Mickel 4758, NY). E. Habit. F. Pinna detail.

ANEMIA

723

Fig. 21. A–C. Anemia brandegeei (Brandegee s.n., in 1904, UC). A. Habit. B. Pinna, adaxial surface. C. Pinna, abaxial surface. D. Anemia intermedia (Jones 23472, UC), habit. E. Anemia affinis (González Tamayo 368, MICH), habit. F. Anemia jaliscana var. integrifolia (Hinton 1935, NY), habit. G. Anemia semihirsuta (Matuda 4802, MEXU), habit.

724

ANEMIA

Fig. 22. A–H. Anemia colimensis. A–C. (Calzada 18255, NY). A. Habit. B. Sterile pinnule. C. Fertile pinnule. D–G. (Calzada 18195, NY). D. Fertile frond. E. Fertile pinnule, adaxial surface. F. Sterile pinnule, abaxial surface. G. Sterile pinnule, adaxial surface. H. (Mickel 1347, NY), frond.

ANETIUM

Fig. 23. A–D. Anetium citrifolium (Türckheim 8625, from Guatemala, NY). A. Habit. B. Rhizome. C. Venation. D. Portion of fertile blade.

725

726

ANOGRAMMA

Fig. 24. A–C. Anogramma leptophylla. A. (Mickel 4710a, NY), habit. B. (McVaugh 22051, MICH), fertile pinnule. C. (Mickel 5032, NY), habit. D, E. Anogramma novogaliciana (González & Palafox 45, NY). D. Habit. E. Fertile segment. F–H, J. Anogramma chaerophylla (Mickel 4710b, NY). F. Habit. G. Fertile segment. H. Rhizome. J. Rhizome scale.

ARACHNIODES

Fig. 25. A, B. Arachniodes denticulata. A. (Hallberg 1465, NY), rhizome, basal pinna, blade apex. B. (Mickel 4677, NY), segment.

727

728

ARGYROCHOSMA

Fig. 26. A–C. Argyrochosma formosa (Mickel 6307, NY). A. Blade. B. Segment. C. Rhizome scale. D, E. Argyrochosma jonesii (Burgess 5759, ARIZ). D. Habit. E. Segment. F, G. Argyrochosma lumholtzii (Hartman 298, US). F. Habit. G. Segment. H, J, K. Argyrochosma microphylla (Pray 1678, NY, from New Mexico). H. Habit. J. Pinnule. K. Rhizome scale.

ARGYROCHOSMA

729

Fig. 27. A–C. Argyrochosma incana (Mickel 4427, NY). A. Habit. B. Pinnule. C. Rhizome scale. D–G. Argyrochosma delicatula. D. (Windham 482, NY), frond. E–G. (Pringle 2581, CAS). E. Habit. F. Segment, adaxial surface. G. Rhizome scale. H, J. Argyrochosma incana (Dorr 2509, TEX). H. Frond. J. Rhizome scale. K, L. Argyrochosma fendleri (Brandegee s.n., in 1878, NY, from Colorado). K. Habit. L. Rhizome scale.

730

ARGYROCHOSMA

Fig. 28. A–C. Argyrochosma limitanea subsp. limitanea A. (Ripley & Barneby 2816, NY, from Arizona), habit. B, C. (Lemmon s.n., May 1881, NY, from Arizona). B. Habit. C. Pinnule. D. Argyrochosma limitanea subsp. mexicana (Hinton et al. 20965, TEX), habit. E–H. Argyrochosma incana (Stanford et al. 684, MEXU). E. Pinna. F. Old pinna with abscised segments. G. Segment, adaxial surface. H. Segment, abaxial surface.

ARGYROCHOSMA

731

Fig. 29. A–C. Argyrochosma peninsularis (Brandegee 650, NY). A. Habit. B. Segments. C. Rhizome scale. D–F. Argyrochosma pilifera (Lyonnett 2570, US). D. Habit. E. Segment. F. Rhizome scale. G, H, J. Argyrochosma palmeri (Parry & Palmer 991, NY). G. Habit. H. Segments. J. Rhizome scale. K–M. Argyrochosma pallens (Pringle 10149, NY). K. Habit. L. Segments. M. Rhizome scale.

732

ASPIDOTIS

Fig. 30. A–F. Aspidotis meifolia (Fryxell & Kirkpatrick 2447, NY). A. Habit. B. Rhizome scale. C. Stipe and rachis, adaxial surface. D. Stipe and rachis, abaxial surface. E. Segment, abaxial surface. F. Segment, adaxial surface. G, H. Aspidotis californica (Montgomery & Huttleston 83-85, NY). G. Segment, abaxial surface. H. Segment, adaxial surface.

ASPLENIUM

733

Fig. 31. A, B. Asplenium pumilum (Hallberg 1635, NY). A. Habit. B. Segment. C, D. Asplenium minimum (González Quintero 1826, NY). C. Habit. D. Rachis and pinna base. E. Asplenium praemorsum (Mickel 3750, NY), pinna and rachis. F. Asplenium serra (Mickel 4109, NY), pinna. G. Asplenium dissectum (Mickel 7095, NY), pinna. H, J. Asplenium serratum (Hallberg 1640, NY). H. Frond. J. Blade margin.

734

ASPLENIUM

Fig. 32. A, B. Asplenium alatum (Ventura 342, NY). A. Habit. B. Pinnae. C, D. Asplenium miradorense (Hallberg 1551, NY). C. Habit. D. Pinnae. E–G. Asplenium pseuderectum (Hallberg 1296, NY). E. Habit. F. Pinnae. G. Rachis.

ASPLENIUM

735

Fig. 33. A, B. Asplenium monanthes (Mickel 5025, NY). A. Habit. B. Pinnae. C. Asplenium resiliens (Mickel 3721, NY), pinnae. D. Asplenium monanthes (Mickel 4529, NY), pinnae. E, F. Asplenium polyphyllum (Mickel 4640, NY). E. Habit. F. Pinnae. G. Asplenium palmeri (Mickel 6655, NY), pinnae. H. Asplenium oligosorum (Hallberg 1444, NY,), pinnae. J. Asplenium formosum (Mickel 3968, NY), pinnae. K. Asplenium castaneum (Mickel 4582, NY), pinnae. L, M. Asplenium hallbergii (Mickel 4579, NY). L. Pinnae. M. Rachis.

736

ASPLENIUM

Fig. 34. A. Asplenium laetum (Hallberg 1469, NY), pinnae. B. Asplenium hoffmannii (Mickel 7411, NY), pinnae. C. Asplenium tuerckheimii (Mickel 7300, NY), pinna. D. Asplenium abscissum (Mickel 6382, NY), pinnae. E. Asplenium yelagagense (Mickel 1058, NY), pinnae. F. Asplenium pteropus (Mickel 6451a, NY), pinnae. G. Asplenium pulchellum (Mickel 7394, NY), pinnae. H. Asplenium riparium (Mickel 1087, NY), pinnae. J. Asplenium harpeodes (Mickel 928, NY), pinnae.

ASPLENIUM

737

Fig. 35. A, B. Asplenium scolopendrium var. americanum (Ghiesbreght 307, NY). A. Frond. B. Blade detail. C, D. Asplenium auritum (Wendt 2690, NY). C. Portion of fertile blade. D. Fertile pinna. E. Asplenium sessilifolium var. sessilifolium (Mickel 3809, NY), fertile pinnae. F–H. Asplenium potosinum (Mickel 3973, NY). F. Frond. G. Proliferous blade tip. H. Sterile pinna, rachis cross section. J, K. Asplenium muenchii (Mickel 3818a, NY). J. Pinna, rachis cross section. K. Proliferous blade tip. L. Asplenium feei (Williams 904, NY, from Panama), fertile pinna.

738

ASPLENIUM

Fig. 36. A, B. Asplenium monodon. A. (Mickel 5101, NY), pinna. B. (Mickel 5840, NY), pinna. C. Asplenium sphaerosporum (Mickel 4775, NY), pinna. D–F. Asplenium cuspidatum D. (Hallberg 1310, NY), pinna. E. (Hallberg 1408, NY), pinna. F. (Mickel 3787a, NY), pinna. G. Asplenium fragrans (Mickel 4378, NY), pinna. H. Asplenium auriculatum (Hallberg 1370, NY) pinna. J. Asplenium salicifolium (Hallberg 1639b, NY), pinna. K, L. Asplenium auriculatum. K. (Hallberg 1369b, NY), pinna. L. (Hallberg 1445, NY), pinna.

ASPLENIUM

739

Fig. 37. A. Asplenium insolitum (Mickel 3810a, NY), pinnae. B, C. Asplenium blepharophorum (Mickel 7069, NY). B. Pinnae. C. Segment. D. Asplenium achilleifolium (Mickel 6188), pinnae. E, F. Asplenium blepharophorum (Mickel 4692, NY). E. Pinnae. F. Segment. G. Asplenium lamprocaulon (Mickel 6051, NY), pinnae. H. Asplenium exiguum (Mickel 6488, NY), habit. J. Asplenium tenerrimum (Mickel 6900, NY), habit. K. Asplenium stolonipes (Mickel 5376, NY), habit. L. Asplenium soleirolioides (Mickel 5386, NY), habit.

740

ASPLENIUM

Fig. 38. A. Asplenium rutaceum (Mickel 6549, NY), pinna. B, C. Asplenium cirrhatum (Mickel 5673, NY). B. Habit. C. Pinna. D. Asplenium myriophyllum (Mickel 3818b, NY), pinna. E. Asplenium uniseriale (Mickel 5704, NY), pinna. F. Asplenium radicans (Mickel 945, NY), pinna. G. Asplenium cristatum (Mickel 7301a, NY), pinna.

ASPLENIUM

741

Fig. 39. A, B. Asplenium nesioticum (Purpus 4880, UC). A. Habit. B. Pinnae. C, D. Asplenium sanchezii (Sánchez 749, US). C. Habit. D. Pinnae. E. Asplenium heterochroum (Schipp 211, UC, from Belize), pinnae. F. Asplenium vespertinum (Moran 30526, NY), pinnae. G, H. Asplenium fibrillosum (Schaffner 109, NY). G. Habit. H. Pinnae. J. Asplenium blepharodes (Brandegee s.n., 24 Jan 1890, NY), pinnae. K. Asplenium pringlei (Pringle 1837, UC), pinnae.

742

ASPLENIUM

Fig. 40. A. Asplenium venturae (Ventura A. 11176, IEB), pinnae. B. Asplenium breedlovei (Breedlove 30041, DS), proximal pinnae. C, D. Asplenium solmsii (Matuda 4173, NY). C. Pinnae. D. Pinnule. E, F. Asplenium eatonii (Cochrane et al. 12060, UC). E. Base of blade. F. Segment detail. G, H. Asplenium delicatulum (Breedlove 50838, CAS). G. Habit. H. Pinna. J, K. Asplenium myriophyllum (Mickel 5369, UC). J. Habit. K. Pinna.

ASPLENIUM

743

Fig. 41. A. Asplenium chihuahuense (Pringle 1444, NY), habit. B. Asplenium arcanum (Ferris 5360, DS), habit. C, D. Asplenium ultimum (Hinton 20142, TEX). C. Frond. D. Pinnae. E, F. Asplenium modestum (Palmer 162, in 1885, US). E. Habit. F. Pinna. G, H. Asplenium septentrionale (Brandegee s.n., 18 May 1893, NY). G. Fronds. H. Blade tips. J, K. Asplenium dalhousiae (Breedlove 18370, CAS). J. Habit. K. Pinnae.

744

ASPLENIUM

Fig. 42. A, B. Asplenium olivaceum (Mickel 7059c, NY). A. Habit. B. Pinnae. C, D. Asplenium trichomanes (Phillips 2348, UC, from Arizona). C. Habit. D. Pinnae. E. Asplenium gentryi (Correll & Gentry 23032, NY), pinnae. F, G. Asplenium dentatum (Vázquez Torres 2548, NY). F. Habit. G. Pinna. H, J. Asplenium harrisii (Breedlove 31750, DS). H. Habit. J. Pinnae. K–M. Asplenium tryonii (Correll & Gentry 23000, US). K. Habit. L. Pinnae. M. Blade apex with bud.

ASPLENIUM

745

Fig. 43. A, B. Asplenium delitescens (Mickel 5784, NY). A. Pinna. B. Pinna detail. C, D. Asplenium virillae (Breedlove 21798, DS). C. Habit. D. Pinna. E. Asplenium cruegeri (Breedlove 39059, NY), habit. F, G. Asplenium barbaense (Mickel 6753, UC). F. Pinnae. G. Pinna. H, J. Asplenium seileri (Matuda 186bis, US). H. Habit. J. Pinnae.

746

ASPLENIUM

Fig. 44. A. Asplenium sessilifolium var. occidentale (Mexia 1557, UC), pinnae. B. Asplenium semipinnatum (Pringle 3367, UC), pinnae. C. Asplenium juglandifolium (Purpus 7249, NY), proximal pinnae and blade apex. D. Asplenium obesum (Mexia 9048, UC), blade apex and proximal pinnae. E, F. Asplenium dianae (Webster & Webster 221, TEX). E. Habit. F. Pinna. G. Asplenium monanthes (Mickel 1615, NY), bud and plantlet at proximal pinna. H. Asplenium flabellulatum (Breedlove 31313, NY), fertile pinna.

ASTROLEPIS

747

Fig. 45. A–G. Astrolepis sinuata (Mickel 4992, NY). A. Habit. B. Sterile pinnae. C. Pinna detail to show scales, scurf, and glands. D. Scurf scales. E. Abaxial scales. F. Adaxial pinna surface. G. Adaxial scales. H, J–L. Astrolepis crassifolia (Mickel 4506, NY). H. Habit. J. Fertile pinnae, abaxial surface. K. Pinna detail, adaxial surface. L. Adaxial scale. M–Q. Astrolepis laevis (Rzedowski 21389, IEB). M. Fertile pinna, abaxial surface. N. Pinna detail, adaxial surface. O. Adaxial scale. P. Pinna detail, abaxial surface. Q. Abaxial scales.

748

ASTROLEPIS

Fig. 46. A–G. Astrolepis integerrima. A–D. (Mickel 3910, NY). A. Habit. B. Fertile pinnae, abaxial surface. C. Pinna detail, adaxial surface. D. Adaxial scale. E. (Carranza 3291, NY), pinna, adaxial surface. F. (Mayfield 1434, MEXU), pinna, adaxial surface. G. (Rzedowski 52494, NY), pinna, adaxial surface. H, J, K. Astrolepis cochisensis (Stevenson 646, NY). H. Habit. J. Pinna, adaxial surface. K. Adaxial scale. L–O. Astrolepis integerrima (Hinton 16559, MEXU). L. Habit. M. Pinna, abaxial surface. N. Pinna detail, adaxial surface. O. Adaxial scales.

ATHYRIUM

749

Fig. 47. A–D. Athyrium skinneri. A. (King & Soderstrom 9926, MICH), habit. B–D. (Mickel 6992a, NY). B. Pinna. C. Segment. D. Sorus. E–G. Athyrium palmense (Mickel 5082, NY). E. Pinna. F. Segment. G. Sorus. H, J, K. Athyrium tripinnatum (Tejero Díez 2919, NY). H. Pinna. J. Segment. K. Sorus.

750

ATHYRIUM

Fig. 48. A. Athyrium arcuatum (Mickel 4676, NY), frond (see also Figs. 49 A, B, E–G). B. Athyrium bourgeaui (Ventura A. 6030, NY), frond (see also Figs. 49 C, D, H, J–L).

ATHYRIUM

751

Fig. 49. A, B. Athyrium arcuatum (Ventura A. 3693, NY) (see also Fig. 48A). A. Pinna. B. Pinnule. C, D. Athyrium bourgeaui (Mickel 4055, NY) (see also Fig. 48B). C. Pinna. D. Pinnule. E–G. Athyrium arcuatum. E. (Mickel 657, ISC), pinnule. F, G. (Rzedowski 24175, NY). F. Pinna. G. Pinnule. H, J–L. Athyrium bourgeaui. H, J. (Correll & Gentry 23079, NY). H. Pinna. J. Pinnule. K, L. (Correll & Gentry 23095, LL). K. Pinnae. L. Segment.

752

AZOLL A

Fig. 50. A–C. Azolla microphylla (Mickel s.n., NY, from NYBG living collection). A. Habit. B. Leafy branch, upper surface. C. Leafy branch, lower surface. D–G. Azolla filiculoides (Smith s.n., UC 1730194). D. Habit. E. Lower surface of branch with sporocarps. F. Microsporangia with glochidia. G. Glochidium. H, J. Redrawn from Eames, Morphology of Vascular Plants, 1936: 251, said to be A. caroliniana, which we treat as a synonym of A. filiculoides. H. Megasporocarp (its walls transparent) containing the large basal spore covered by the pitted perispore, and the apical massulae (“floats”) which surround a central columella. J. Megaspore with an apical cap of sporocarp tissue, a cushion-like columella, four “floats,” and numerous whip-like appendages.

BLECHNUM

753

Fig. 51. A, B. Blechnum wardiae (Hallberg 1542, NY) (see also Fig. 54A). A. Sterile pinnae. B. Fertile pinna. C. Blechnum stoloniferum (Mickel 5248, NY), habit (see also Figs. 54J–L). D. Blechnum lehmannii (Hallberg 1538, NY), habit.

754

BLECHNUM

Fig. 52. A. Blechnum fragile (Ventura 390, NY), habit (sterile frond). B. Blechnum ensiforme (Mickel 991, NY), habit (sterile frond) (see also Fig. 54C). C–E. Blechnum schiedeanum (Mickel 1561, NY). C. Sterile pinna. D. Venation of sterile pinna. E. Apex of sterile pinna. F–H. Blechnum falciforme (Mickel 1238, NY). F. Sterile pinna. G. Venation and margin of sterile pinna. H. Fertile pinnae.

BLECHNUM

755

Fig. 53. A. Blechnum appendiculatum (Mickel 850, ISC), habit (see also Fig. 54D). B, C. Blechnum occidentale (Mickel 6988, NY) (see also Fig. 54E). B. Habit. C. Section of fertile pinna. D. Blechnum polypodioides (Hallberg 1307, NY), habit. E–G. Blechnum serrulatum (Rzedowski 1226, NY). E. Pinnae. F. Pinna apex. G. Pinna margin. H. Blechnum gracile (Mickel 6085, NY), habit.

756

BLECHNUM

Fig. 54. A. Blechnum wardiae (Mickel 5578, NY), sterile frond (see also Figs. 51A, B). B. Blechnum divergens (Mickel 963, ISC), base of sterile blade. C. Blechnum ensiforme (Mickel 5651, NY), base of sterile blade. D. Blechnum appendiculatum (Mickel 850, ISC), stipe detail (see also Fig. 53A). E. Blechnum occidentale (Hellwig 463, NY), stipe detail. F–H. Blechnum penna-marina (Tate 119, NY, from Bolivia). F. Fertile pinnae. G. Sterile pinnae, adaxial surface. H. Cross section of sterile pinna. J–L Blechnum stoloniferum (Mickel 5248, NY) (see also Fig. 51C). J. Fertile pinnae. K. Sterile pinnae, adaxial surface. L. Cross section of sterile pinna.

BLECHNUM

757

Fig. 55. A. Blechnum meridense (Britton & Hazen 243, NY, from Trinidad), frond. B. Blechnum ×confluens (Blechnum occidentale × polypodioides) (Palmer 1941A, NY), frond. C. Blechnum × caudatum (B. gracile × occidentale) (Hernández G. 1523, NY), habit.

758

BOLBITIS

Fig. 56. A–C. Bolbitis portoricensis. A. (McVaugh 18983, MICH), habit. B, C. (Hallberg 1646, NY). B. Venation. C. Fertile pinna.

BOLBITIS

759

Fig. 57. A–C. Bolbitis aliena (Valeur 535, UC, from Dominican Republic). A. Habit, sterile frond. B. Fertile frond. C. Venation, sterile frond. D, E. Bolbitis serratifolia (Breedlove 32817, DS). D. Proximal pinnae, sterile frond. E. Blade, fertile frond. F–H. Bolbitis umbrosa (Breedlove 22483, NY). F. Base of sterile blade. G. Blade detail, sterile frond. H. Venation, fertile blade.

760

BOLBITIS

Fig. 58. A, B. Bolbitis pergamentacea (Hernández G. 1005, NY). A. Sterile blade. B. Fertile blade. C–E. Bolbitis hemiotis. C, D. (Hart 6924, NY, from Trinidad). C. Sterile pinna. D. Fertile pinnae. E. (Fendler 101, NY, from Trinidad), rachis bud. F, G. Bolbitis bernoullii (Hernández G. 850, NY). F. Sterile pinnae. G. Fertile pinnae. H, J, K. Bolbitis hastata (Hallberg 1597, NY). H. Sterile blade. J. Sterile pinna apex, with bud. K. Venation, sterile blade.

BOMMERIA

761

Fig. 59. A. Bommeria elegans (Mickel 6231, NY), habit. B, C. Bommeria ehrenbergiana (Fisher s.n., 14 Aug 1937, NY). B. Frond. C. Venation. D–F. Bommeria hispida (McVaugh 17756, NY). D. Habit. E. Rhizome detail. F. Blade detail.

762

BOMMERIA

Fig. 60. A–D. Bommeria pedata (Mickel 806, NY). A. Frond. B. Segment, abaxial surface. C. Segment, adaxial surface. D. Venation. E–G. Bommeria subpaleacea (Mickel 4556, NY). E. Frond. F. Blade detail, abaxial surface. G. Blade detail, adaxial surface.

BOTRYCHIUM

763

Fig. 61. A, B. Botrychium tolucaense (Pringle 5193, VT). A. Habit. B. Pinna detail. C–E. Botrychium minganense (Schneider 966, F). C. Habit. D. Sterile blade. E. Fertile segment detail. F–H. Botrychium decompositum. F. (Wagner 80805, US), frond. G. (Wagner 80806, MICH), sterile blade. H. (Guzman 214, MEXU), proximal pinnae.

764

BOTRYCHIUM

Fig. 62. A. Botrychium decompositum (Roe et al. 1190A, WIS), habit. B. Botrychium virginianum (Mickel 3942, NY), habit.

BOTRYCHIUM

765

Fig. 63. A–D. Botrychium decompositum. A. (Mickel 4696, NY), habit. B. (Pringle 8922, ENCB), segments. C. (Calzada 15956, FCME), proximal pinnae. D. (Wagner 80818, MICH), proximal pinnae.

766

BOTRYCHIUM

Fig. 64. A–D. Botrychium schaffneri (Pringle 8922, MEXU). A. Habit of smaller plant. B. Sterile segment. C. Fertile segment. D. Proximal pinnae of larger plant. E, F. Botrychium socorrense (Levin 2044, SD). E. Proximal pinna. F. Segment.

CAMPYLONEURUM

767

Fig. 65. A, B. Campyloneurum angustifolium (Mickel 6959, NY) (see also Fig. 66F). A. Habit. B. Blade detail. C, D. Campyloneurum amphostenon (Mickel 4127, NY) (see also Fig. 66G). C. Habit. D. Blade detail. E–G. Campyloneurum tenuipes. E. (Hallberg 1453, NY), habit. F. (Mickel 6050, NY), blade detail. G. (Hallberg 1453, NY), blade detail. H. Campyloneurum xalapense (Mickel 964, NY), blade detail. J, K. Campyloneurum serpentinum (Breedlove 22151, NY). J. Habit. K. Blade detail. L–N. Campyloneurum phyllitidis (Mickel 1309, NY). L. Habit. M. (Mickel 5782, NY), blade detail. N. (Mickel 1309, NY), blade detail.

768

CAMPYLONEURUM

Fig. 66. A, B. Campyloneurum repens (Breedlove 34063, DS). A. Habit. B. Venation. C, D. Campyloneurum costatum (Proctor et al. 27063, NY, from Nicaragua). C. Habit. D. Venation. E. Camplyoneurum ensifolium (Lott 3049, NY), rhizome scale. F. Campyloneurum angustifolium (Nee 32158, NY), rhizome scale (see also Figs. 65A, B). G. Campyloneurum amphostenon (Mickel 4127, NY), rhizome scale (see also Figs. 65C, D).

CERADENIA

769

Fig. 67. A, B. Ceradenia pruinosa (Hallberg 1724, NY). A. Habit. B. Pinnae. C, D. Ceradenia jungermannioides (Hallberg 1524, NY). C. Habit. D. Blade detail. E–H. Ceradenia margaritata (Breedlove 31801, DS). E. Habit. F. Rhizome scale. G. Pinnae. H. Sorus. J, K. Ceradenia oidiophora (Mickel 3822, NY). J. Habit. K. Pinnae.

770

CERATOPTERIS

Fig. 68. A–D. Ceratopteris pteridoides. A. (Bruza 3121, NY, from Louisiana), habit. B–D. (Curtiss 3690, NY, from Florida). B. Fertile segment detail. C. Venation, sterile blade. D. Marginal plantlet. E–G. Ceratopteris thalictroides. E. Sterile frond (s. coll./s.n., from Brazil). F. Venation, sterile blade. G. Fertile frond (Maguire & Stahel 22719, NY).

CHEILANTHES

771

Fig. 69. A, B. Cheilanthes skinneri (Mickel 6902, NY). A. Habit. B. Rhizome scale. C–E. Cheilanthes bolborrhiza (Mickel 5207, NY). C. Pinna. D. Roots. E. Segment.

772

CHEILANTHES

Fig. 70. A, B. Cheilanthes farinosa (Mickel 5366, NY). A. Habit. B. Segment. C, D. Cheilanthes aurea var. aurea (Mickel 845, NY). C. Habit. D. Pinnule. E. Cheilanthes chaerophylla (Mickel 6254, NY), frond. F. Cheilanthes cuneata (Mickel 6625b, NY), blade (see also Figs. 71E, F). G–K. Cheilanthes pyramidalis (see also Figs. 86A, B). G. (Mickel 3857, NY), frond. H. (Mickel 1261, NY), pinnae. J, K. (Mickel 3857, NY). J. Pinnule. K. Segment.

CHEILANTHES

773

Fig. 71. A, B. Cheilanthes arizonica (Mickel 5252b, NY) (see also Figs. 77C–E). A. Frond. B. Pinnule. C, D. Cheilanthes membranacea (Mickel 7057, NY) (see also Figs. 75J–L). C. Pinna. D. Pinnule. E, F. Cheilanthes cuneata (Mickel 4444, NY) (see also Fig. 70F). E. Frond. F. Pinnule. G, H. Cheilanthes angustifolia (Mickel 4782, NY). G. Frond. H. Pinnule.

774

CHEILANTHES

Fig. 72. A, B. Cheilanthes allosuroides (Mickel 4952, NY). A. Pinnae. B. Segments. C, D. Cheilanthes notholaenoides (Hallberg 1463, NY) (see also Fig. 87N). C. Pinnae. D. Segments. E, F. Cheilanthes mickelii (Mickel 4210, NY). E. Pinnae. F. Segments. G, H. Cheilanthes cucullans (Mickel 784, NY). G. Pinnae. H. Segments. J, K. Cheilanthes aemula (Mickel 578, NY). J. Pinna. K. Pinnule. L–O. Cheilanthes fimbriata (Mickel 1483, NY). L. Frond. M. Pinnae. N. Segment, abaxial surface. O. Segment, adaxial surface.

CHEILANTHES

775

Fig. 73. A–C. Cheilanthes kaulfussii (Mickel 4949, NY). A. Frond. B, C. Pinnules. D, E. Cheilanthes longipila var. longipila (Mickel 1669, NY) (see also Figs. 87E–H). D. Pinnae. E. Segment. F–H. Cheilanthes lendigera (Mickel 3894, NY) (see also Fig. 83Q). F. Pinnae. G. Pinna, adaxial surface. H. Pinna, abaxial surface. J–P. Cheilanthes myriophylla (Mickel 773, NY) (see also Fig. 80A). J. Habit. K. Pinnae. L. Pinnule, adaxial surface. M. Pinnule, abaxial surface. N. Segments, abaxial surface. O, P. Blade scales.

776

CHEILANTHES

Fig. 74. A–E. Cheilanthes bonariensis (Mickel 6202, NY). A. Habit. B. Rhizome scale. C. Pinna detail, abaxial surface. D. Pinna detail, abaxial surface. E. Pinna detail, adaxial surface. F–K. Cheilanthes yatskievychiana (Burquez M. 96-302, MO). F. Habit. G. Rhizome scale. H. Stipe detail. J. Blade detail, abaxial surface. K. Blade detail, adaxial surface. L–Q. Cheilanthes brachypus (Mickel 6895, NY). L. Habit. M. Rhizome scale. N. Stipe scale. O. Fertile pinna. P. Segment detail, adaxial surface. Q. Segment detail, abaxial surface.

CHEILANTHES

777

Fig. 75. A–D. Cheilanthes aurantiaca (McVaugh 20111, NY). A. Habit. B. Segment, adaxial surface. C. Segment, abaxial surface. D. Rhizome scale. E–H. Cheilanthes aurea var. palmeri (Pringle 2584, NY). E. Habit. F. Pinnule, adaxial surface. G. Pinnule, abaxial surface. H. Rhizome scale. J–L. Cheilanthes membranacea (McVaugh 12869, MICH) (see also Figs. 71C, D). J. Pinna. K. Pinnule, abaxial surface. L. Rhizome scale. M–O. Cheilanthes brandegeei (Wiggins 7649, NY). M. Habit. N. Segment, abaxial surface. O. Rhizome scale.

778

CHEILANTHES

Fig. 76. A–C. Cheilanthes hintoniorum (Cowan 3753, UC). A. Habit. B. Segment, abaxial surface. C. Rhizome scale. D–F. Cheilanthes apiacea (Nesom 7716, NY). D. Habit. E. Segment. F. Rhizome scale. G–J. Cheilanthes pellaeopsis (Anderson & Laskowski 4394, NY). G. Pinna. H. Segment. J. Rhizome scale. K–N. Cheilanthes purpusii (Purpus 4881, UC). K. Habit. L. Pinna. M. Segment. N. Rhizome scale.

CHEILANTHES

779

Fig. 77. A, B. Cheilanthes decurrens (Molina et al. 30033, F, from Guatemala). A. Pinna. B. Pinna bases. C–E. Cheilanthes arizonica (Gentry & Arguelles 18163, NY) (see also Figs 71A, B). C. Pinna. D. Rhizome scale. E. Segment. F–H. Cheilanthes complanata var. complanata (Breedlove 41747, MEXU). F. Pinnae. G. Rhizome scale. H. Segment. J–L. Cheilanthes complanata var. interrupta (Espin M. 149, NY). J. Habit. K. Rhizome scale. L. Segment. M, N. Cheilanthes decomposita (Pringle 2588, NY). M. Basal pinna. N. Segment detail, abaxial surface. O, P. Cheilanthes decurrens (Maxon & Hay 3709, NY, from Guatemala). O. Frond. P. Pinna bases.

780

CHEILANTHES

Fig. 78. A–D. Cheilanthes horridula (Pringle 447, NY). A. Habit. B. Segment and rachis detail, abaxial surface. C. Segment and rachis detail, adaxial surface. D. Rhizome scale. E–H. Cheilanthes microphylla (Hernández G. 924, NY). E. Rhizome and partial frond. F. Segment and rachis detail, abaxial surface. G. Segment and rachis detail, adaxial surface. H. Rhizome scale. J–L. Cheilanthes alabamensis (Camp 2232, NY). J. Pinnae. K. Segments and rachis detail, abaxial surface. L. Segment and rachis detail, adaxial surface. M–P. Cheilanthes fendleri (McGill & Pinkava P12330, ASU). M. Pinnae. N. Segments and rachis detail, abaxial surface. O. Segments and rachis detail, adaxial surface. P. Rhizome scale.

CHEILANTHES

781

Fig. 79. A–D. Cheilanthes wrightii (Pringle 445, NY). A. Habit. B. Segments. C. Rachis and stipe detail, adaxial surface. D. Rhizome scale. E–H. Cheilanthes pringlei var. pringlei (Palmer 116, NY). E. Habit. F. Segments, adaxial surface. G. Segments, abaxial surface. H. Rhizome scale. J–M. Cheilanthes pringlei var. moncloviensis (Pray 3148, LAM). J. Habit. K. Segments. L. Rachis and stipe detail, adaxial surface. M. Rhizome scale. N–P. Cheilanthes peninsularis var. peninsularis (Jones 24559, NY). N. Habit. O. Segments. P. Rhizome scale. Q–S. Cheilanthes peninsularis var. insularis (Felger 15794, NY). Q. Habit. R. Segments and rachis detail. S. Rhizome scale.

782

CHEILANTHES

Fig. 80. A. Cheilanthes myriophylla (Mickel 756, NY), stipe (see also Figs. 73J–P). B–H. Cheilanthes villosa (Pringle 459, NY). B. Rhizome and partial frond. C. Stipe. D. Rhizome scale. E. Pinnule, adaxial surface. F. Pinnule, abaxial surface. G. Pinnule, abaxial surface, with scales removed. H. Blade scale. J–O. Cheilanthes clevelandii (Wiggins & Gillespie 4048, NY). J. Rhizome and partial frond. K. Rhizome scale. L. Pinnule, adaxial surface. M. Pinnule, abaxial surface. N. Pinnule, abaxial surface, with scales removed. O. Blade scale. P–V. Cheilanthes lindheimeri (McVaugh 16758, NY). P. Rhizome and partial frond. Q, R. Rhizome scales. S. Pinnule, adaxial surface. T. Pinnule, abaxial surface. U. Pinnule, abaxial surface, with scales removed. V. Blade scales. W–Bb. Cheilanthes yavapensis (Chambers 645, DS). W. Rhizome and partial frond. X. Rhizome scale. Y. Pinnule, adaxial surface. Z. Pinnule, abaxial surface. Aa. Pinnule, abaxial surface, with scales removed. Bb. Blade scales.

CHEILANTHES

783

Fig. 81. A–B. Cheilanthes lozanoi var. seemannii. A–B. (Mickel 6646, NY). A. Habit. B. Rhizome scale. C–E. Cheilanthes lozanoi var. lozanoi (Matuda 31543, US). C. Pinnae. D. Rhizome scale. E. (Rzedowski 22505, NY), pinnae. F. Cheilanthes lozanoi var. seemannii (Mickel 4951, NY), pinnae. G. Cheilanthes lozanoi var. lozanoi (Hinton et al. 4318, ARIZ), pinnae. H, J. Cheilanthes lozanoi var. seemannii (see also Figs. A, B, F). H. (Rose & Painter 6864, NY), pinnae. J. (Palmer 1437, NY), pinnae. K–P. Cheilanthes wootonii (Moran & Thomas 14371, CAS). K. Rhizome and partial frond. L. Rhizome scale. M. Pinnule, adaxial surface. N. Pinnule, abaxial surface. O. Pinnule, abaxial surface, with scales removed. P. Blade scale. Q–V. Cheilanthes covillei (Orcutt s.n., 8 Sep 1884, NY). Q. Rhizome and partial frond. R. Rhizome scale. S. Pinnule, adaxial surface. T. Pinnule, abaxial surface, with scales. U. Pinnule, abaxial surface, with scales removed. V. Blade scale.

784

CHEILANTHES

Fig. 82. A–E. Cheilanthes tomentosa (Knobloch 5946, NY). A. Pinnae. B. Pinnule and rachis detail, adaxial surface. C. Pinnule and rachis detail, abaxial surface. D. Rhizome scale. E. Stipe base scale. F–H. Cheilanthes chipinquensis (Pringle 2776, UC). F. Pinnae. G. Pinnule. H. Rhizome scale. J–N. Cheilanthes eatonii (Grimes 2284, NY). J. Rhizome and partial frond. K. Pinnule and rachis detail, abaxial surface. L–N. (Stanford et al. 341, NY). L. Pinnule and rachis detail, adaxial surface. M. Rhizome scale. N. Pinnule detail, abaxial surface. O–S. Cheilanthes jamaicensis (Pennell 17347, NY; P and S from Muller 3252, MICH). O. Rhizome and partial frond. P. Rachis detail. Q. Pinnule, abaxial surface. R. Pinnule, adaxial surface. S. Rhizome scale.

CHEILANTHES

785

Fig. 83. A–C. Cheilanthes leucopoda (Chase 7234, NY). A. Habit. B. Segments, adaxial surface. C. Segments, abaxial surface. D–G. Cheilanthes feei. D. (Worthington 12512, NY), habit. E–G. (Vasey s.n., in 1881, NY, from Texas). E. Pinnules, adaxial surface. F. Pinnule, abaxial surface. G. Rhizome scale. H, J–L. Cheilanthes mexicana (Parry & Palmer 988, in 1878, NY). H. Pinnae. J. Rachis detail. K. Pinnule. L. Segment detail, without hairs. M–P. Cheilanthes marsupianthes (Matuda 31197, MEXU). M. Rhizome and partial frond. N. Rachis detail. O. Pinnule detail. P. Rhizome scale. Q. Cheilanthes lendigera (Ventura A. 65, NY), segment (see also Figs. 73F–H).

786

CHEILANTHES

Fig. 84. A–D. Cheilanthes parryi (Felger 16896, ARIZ). A. Habit. B. Pinnule, adaxial surface. C. Pinnule, abaxial surface, with scales, and with scales removed. D. Rhizome scale. E–H. Cheilanthes newberryi (Franceschi s.n., winter 1892-93, NY). E. Habit. F. Pinnules, adaxial surface. G. Pinnules, abaxial surface. H. Rhizome scale. J–M. Cheilanthes viscida (Munz 7821, NY). J. Habit. K. Pinnule, adaxial surface. L. Pinnules, abaxial surface. M. Rhizome scale. N–Q. Cheilanthes longipila var. brevipila (Antonio 339, NY). N. Habit. O. Pinnules, adaxial surface. P. Pinnules, abaxial surface. Q. Rhizome scale.

CHEILANTHES

787

Fig. 85. A–C. Cheilanthes marginata (Brade 847, NY, from Costa Rica). A. Habit. B. Rhizome scale. C. Segment. D–G. Cheilanthes spiculata (Hinton 6837, LL). D. Habit. E. Rhizome scale. F. Pinnule, adaxial surface. G. Pinnule, abaxial surface. H, J, K. Cheilanthes marginata (Breedlove 22636, F). H. frond. J. Basal pinnae. K. Pinnule abaxial surface.

788

CHEILANTHES

Fig. 86. A–E. Cheilanthes pyramidalis (see also Figs. 70G–K). A, B. (Ventura A 2453, NY) A. Pinna. B. Segment. C–E. (Yatskievych 81-403, ARIZ). C. Frond. D. Segment, abaxial surface. E. Segment apex, adaxial surface. F, G. Cheilanthes marginata (see also Figs. 85A–C, H, J, K) (hort. berol., PH, isotype of C. hirsuta). F. Rachis and segment bases, abaxial surface. G. Segment apex, adaxial surface.

CHEILANTHES

789

Fig. 87. A–D. Cheilanthes maxonianum (Viereck 76, US). A. Frond. B. Rachis detail. C. Segment, adaxial surface. D. Segment, abaxial surface. E–H. Chelanthes longipila var. longipila (Pringle 1863, NY) (see also Figs. 73D, E). E. Pinnae. F. Rachis detail. G. Segment, adaxial surface. H. Segment, abaxial surface. J–M. Cheilanthes longipila var. brevipila (Calzada 16172, NY). J. Habit. K. Rachis detail. L. Segment, adaxial surface. M. Segment, abaxial surface. N. Cheilanthes notholaenoides (Díaz Barriga V. 6823, NY), rachis hairs (see also Figs. 72C, D).

790

CHEILOPLECTON

Fig. 88. A. Cheiloplecton rigidum var. lanceolatum (Mickel 4093, NY), frond. B–F. Cheiloplecton rigidum var. rigidum (Mickel 6313, NY). B. Habit. C. Segment. D. Recurved margin. E, F. Rhizome scales.

CIBOTIUM

Fig. 89. A–C. Cibotium schiedei (Ventura A. 235, NY). A. Pinna. B. Stipe base. C. Segments. D. Cibotium regale (Ghiesbreght s.n., NY), segments.

791

792

CNEMIDARIA

Fig. 90. A–C. Cnemidaria apiculata (Hallberg 1557, NY). A. Pinna. B. Stipe. C. Segment. D, E. Cnemidaria decurrens (Hallberg 1567, NY). D. Pinna. E. Pinna detail.

COCHLIDIUM

793

Fig. 91. A–C. Cochlidium linearifolium (Mickel 5953, NY). A. Habit. B. Blade detail, abaxial surface. C. Blade cross section, and adaxial surface. D, E. Cochlidium rostratum (Croat & Hannon 63757, UC, from Guatemala). D. Habit. E. Rhizome scale. F, G. Cochlidium serrulatum (Mickel 5954, NY). F. Habit. G. Blade detail.

794

CTENITIS

Fig. 92. A–C. Ctenitis submarginalis (Mickel 1091, ISC). A. Pinnae. B. Stipe base. C. Segment. D, E. Ctenitis nigrovenia (Mickel 7215, NY). D. Pinna. E. Segment. F–H. Ctenitis melanosticta (Hallberg 1338, NY). F. Pinna base. G. Segments. H. Sorus. J–M. Ctenitis excelsa. J–L. (Hallberg 1321, NY). J. Pinna base. K. Stipe base. L. Segment. M. (Mickel 6402, NY), rachis, adaxial surface.

CTENITIS

795

Fig. 93. A, B. Ctenitis refulgens (Breedlove 33376, DS). A. Proximal pinna, abaxial surface. B. Segment, abaxial surface. C, D. Ctenitis microchlaena (Breedlove 27456, NY). C. Proximal pinna, abaxial surface. D. Segment, abaxial surface. E, F. Ctenitis salvinii (Breedlove 21665, NY). E. Proximal pinna, abaxial surface. F. Segment, abaxial surface. G, H. Ctenitis lanceolata (Breedlove 38989a, DS). G. Habit. H. Base of pinna, abaxial surface. J, K. Ctenitis thelypteroides (Breedlove 33280, NY). J. Proximal pinna, abaxial surface. K. Segment, abaxial surface. L, M. Ctenitis baulensis (Breedlove 31328, DS). L. Proximal pinna, abaxial surface. M. Segment, abaxial surface.

796

CTENITIS

Fig. 94. A, B. Ctenitis erinacea (Croat 46382, UC). A. Base of proximal pinna. B. Segment, abaxial surface. C, D. Ctenitis bullata (Breedlove 25339, NY). C. Base of proximal pinnae. D. Segment, abaxial surface. E, F. Ctenitis grisebachii (Hernández M. 513, NY). E. Base of proximal pinna. F. Segment, abaxial surface. G, H, J. Ctenitis ursina (Breedlove 32274, NY). G. Base of proximal pinna. H. Segment, abaxial surface. J. Indusium. K, L. Ctenitis strigilosa (Pringle 6077, NY). K. Proximal pinnae. L. Segment, abaxial surface.

CTENITIS

797

Fig. 95. A, B. Ctenitis interjecta (Martínez S. et al. 24928, UC). A. Base of proximal pinna. B. Segments, abaxial surface. C, D. Ctenitis chiapasensis (Münch 117, US). C. Proximal pinna. D. Segment, abaxial surface. E–H. Ctenitis sloanei (Breedlove 33125, NY). E. Pinnules, abaxial surface. F. Stipe base. G. Segment, abaxial surface. H. Stipe scale. J, K. Ctenitis mexicana (Ventura A. 3308, NY). J. Base of proximal pinna. K. Segment, abaxial surface.

798

CTENITIS

Fig. 96. A, B. Ctenitis hemsleyana (Mickel 1001, NY). A. Rhizome and proximal pinnae. B. Segments, abaxial surface. C, D. Ctenitis equestris var. equestris (Breedlove 24029, NY). C. Rachis detail. D. Rachis scale. E, F. Ctenitis equestris var. erosa (Mickel 4164, NY). E. Rachis detail. F. Rachis scale.

CULCITA

Fig. 97. A–D. Culcita coniifolia (Mickel 917, NY). A. Pinnule. B. Stipe base. C. Segment. D. Segment detail.

799

800

CYATHEA

Fig. 98. A–F. Cyathea bicrenata (Mickel 1459, NY). A. Pinnules. B. Stipe. C. Stipe scale. D. Segment, adaxial surface. E. Segment, abaxial surface. F. Segment detail, abaxial surface. G, H, J, K. Cyathea myosuroides (Mickel 1495, NY). G. Pinnules. H. Stipe. J. Stipe scale. K. Segment. L–P. Cyathea schiedeana (Hallberg 1329, NY). L. Pinnules. M. Stipe. N. Stipe scale. O. Segment. P. Segment base detail.

CYATHEA

801

Fig. 99. A–E. Cyathea costaricensis (Mickel 1302, NY). A. Pinnules. B. Stipe. C. Stipe scale. D. Segment. E. Segment detail. F–H, J, K. Cyathea divergens var. tuerckheimii (Hallberg 1433, NY). F. Pinnules. G. Stipe. H. Stipe scale. J. Segment. K. Segment detail. L–P. Cyathea fulva (Mickel 4242, NY). L. Pinnules. M. Stipe. N. Stipe scale. O. Segment. P. Segment detail.

802

CYATHEA

Fig. 100. A–F. Cyathea godmanii (Breedlove 29657, NY). A. Stipe. B. Stipe scale. C. Sterile segment detail, abaxial surface. D. Fertile segment detail, abaxial surface. E. Segment, adaxial surface. F. Blade scale. G, H, J. Cyathea microdonta (Smith 2095, NY). G. Rachis and pinnules. H. Rachilla with spine. J. Segment.

CYCLOPELTIS

Fig. 101. A, B. Cyclopeltis semicordata A. (Ishiki et al. 2020, NY), rhizome and frond base. B. (Breedlove 20738, NY), pinna base, abaxial surface.

803

804

CYSTOPTERIS

Fig. 102. A–F. Cystopteris fragilis (see also Figs. 103A–E, H, J). A–D. (McVaugh 12873, MICH). A. Habit. B. Pinna. C. Segment. D. Sorus. E, F. (Mickel 6657, NY). E. Pinna. F. Segment. G, H, J. Cystopteris membranifolia (Mickel 4544, NY). G. Habit. H. Pinna. J. Segment.

CYSTOPTERIS

805

Fig. 103. A–E. Cystopteris fragilis (see also Figs. 102A–F). A, B. (Mickel 6490, NY). A. Pinna. B. Segment. C, D. (Mickel 6020, NY). C. Habit. D. Pinna. E. (Mickel 6657, NY), pinna. F, G. Cystopteris millefolia (Hinton 13499, ARIZ). F. Frond. G. Pinna. H, J. Cystopteris fragilis (Mickel 5362, NY) (see also Figs. 102A–F, 103A–E). H. Habit. J. Pinna.

806

DANAEA

Fig. 104. A, B. Danaea nodosa (Mickel 6457, NY). A. Sterile pinnae. B. Fertile pinna. C, D. Danaea cuspidata (Mickel 6456). C. Sterile pinnae and nodose stipe. D. Fertile pinnae. E–G. Danaea elliptica (Mickel 6511, NY). E. Fertile pinnae. F. Fertile pinna detail. G. Sterile pinnae. H. Danaea nodosa (Mickel 6408, NY), sterile pinnae.

DENNSTAEDTIA

807

Fig. 105. A–J. Dennstaedtia cornuta. A–F. (Mickel 6575, NY). A. Pinna bases. B. Bud in pinna axil. C. Fertile segment. D. Sterile segment. E. Sorus. F. Rachis hair. G–J. (Lorea 2921, NY). G. Pinnules. H. Segment. J. Sorus. K–Q. Dennstaedtia bipinnata. K–O. (Ventura A. 5047, NY). K. Pinnule. L. Adaxial axis groove. M. Segment. N. Sorus. O. Costal hair. P, Q. (Pringle 3407, NY). P. Sori. Q. Hairs. R–U. Dennstaedtia globulifera (Rose 11058, NY). R. Pinnule. S. Segment. T. Sorus. U. Costal hairs.

808

DENNSTAEDTIA

Fig. 106. A–E. Dennstaedtia distenta (Mickel 6003 and 1175, NY). A. Pinna bases. B. Axis grooves. C. Segment. D. Sorus. E. Axis hairs. F–H, J. Dennstaedtia cicutaria (Pringle 3824, NY). F. Pinna base. G. Segment. H. Sorus. J. Axis hairs.

DENNSTAEDTIA

809

Fig. 107. A–C. Dennstaedtia spinosa (Mickel 2730, US, from Costa Rica). A. Pinna base. B. Rachis. C. Segment. D, E. Dennstaedtia auriculata (Breedlove 33693, NY). D. Pinna. E. Segment. F, G. Dennstaedtia coronata (Münch 60, DS). F. Pinna base. G. Pinnule detail.

810

DICKSONIA

Fig. 108. A–C. Dicksonia sellowiana (Mickel 5348, NY). A. Pinna base. B. Stipe base. C. Segment.

DICRANOPTERIS

811

Fig. 109. A–D. Dicranopteris flexuosa. A. (Smith 1828, NY, from Costa Rica), portion of blade. B–D. (Nee 10151, NY, from Panama). B. Forking detail. C. Segments. D. Segment base.

812

DIDYMOCHLAENA

Fig. 110. A–E. Didymochlaena truncatula (Ventura 5095, NY). A. Blade apex. B. Pinna. C. Stipe base. D. Young sorus. E. Mature sorus.

DIPLAZIUM

Fig. 111. A–C. Diplazium lonchophyllum. A. (McVaugh 11775, MICH), habit. B. (Hinton 16258, MICH), pinna. C. (McVaugh 18980, MICH), pinna detail.

813

814

DIPLAZIUM

Fig. 112. A–C. Diplazium wendtii (Wendt et al. 5580, NY). A. Pinna. B. Segment. C. Stipe base scale. D, E. Diplazium puberulentum (Hallberg 1543, NY). D. Pinna base. E. Segment. F, G. Diplazium striatum (Mickel 7239, NY). F. Pinna base. G. Segments. H. Diplazium hians (Morton 7159, NY, from Honduras), segment. J. Diplazium drepanolobium (Ghiesbreght 361, NY), segments. K, L. Diplazium altissimum (Mickel 7100, NY). K. Fertile pinnules showing buds. L. Pinnule base. M, N. Diplazium cristatum (Mickel 6046a, NY). M. Pinna. N. Segment.

DIPLAZIUM

815

Fig. 113. A, B. Diplazium werckleanum (Hallberg 1413, NY). A. Pinna. B. Pinna detail. C, D. Diplazium donnell-smithii (Hellwig 457, NY). C. Pinnule. D. Segment. E, F. Diplazium franconis (Mickel 5354, NY). E. Pinna. F. Pinnule. G, H. Diplazium ternatum (Mickel 4264, NY). G. Frond. H. Pinna detail. J, K. Diplazium plantaginifolium (Hallberg 1643, NY). J. Blade. K. Blade detail.

816

DIPLAZIUM

Fig. 114. A, B. Diplazium expansum (Hallberg 1450, NY). A. Pinnule. B. Segment. C, D. Diplazium hellwigii (Hellwig 404, NY). C. Pinnule. D. Segment. E. Diplazium diplazioides (Mickel 5688, NY), segment. F. Diplazium urticifolium (Mickel 5654, NY), pinna. G, H. Diplazium striatastrum (Hallberg 1332, NY). G. Pinna. H. Segment. J, K. Diplazium lindbergii (Mickel 6049, NY). J. Pinna. K. Segment.

DIPLAZIUM

817

Fig. 115. A, B. Diplazium obscurum (Croat & Hannon 65231, UC). A. Proximal pinna. B. Blade apex, showing rachis buds. C, D. Diplazium neglectum (Matuda 3985, NY). C. Proximal pinna. D. Portion of frond apex, showing rachis buds. E, F. Diplazium gomezianum (Purpus 7112, UC). E. Base of proximal pinna. F. Segment. G, H. Diplazium vera-pax (Fink 143, UC). G. Habit. H. Base of proximal pinna with rachis bud.

818

DIPLAZIUM

Fig. 116. A, B. Diplazium errans (Campos 1531, NY). A. Blade. B. Pinna detail. C–E. Diplazium grandifolium (Mejia et al. 31399, from Dominican Republic). C. Pinnae and blade apex. D. Pinna detail. E. Sorus.

DIPLOPTERYGIUM

819

Fig. 117. A–F. Diplopterygium bancroftii. A. (Mickel 5680, NY), frond. B–F. Hallberg 1528, NY). B. Bud at first fork. C. Stipe section. D. Segments. E. Segment detail. F. Bud scale.

820

DORYOPTERIS

Fig. 118. A–C. Doryopteris palmata (Ventura A. 4985, NY). A. Habit. B. Blade base showing venation and buds. C. Blade detail. D, E. Doryopteris concolor (Mickel 5973, NY). D. Frond. E. Pinna showing venation.

DRYOPTERIS

Fig. 119. A–C. Dryopteris maxonii (Pringle 1839, MICH, rhizome from Reveles s.n., 28 Oct 1986, NY). A. Habit. B. Pinnules. C. Sori.

821

822

DRYOPTERIS

Fig. 120. A, B. Dryopteris muenchii (Breedlove 22027, NY). A. Proximal pinnae. B. Segment. C, D. Dryopteris arguta (Wiggins 4249, NY). C. Proximal pinna. D. Segment. E, F. Dryopteris futura (Fraser-Jenkins 13184, UC). E. Proximal pinna. F. Segment. G, H. Dryopteris filix-mas (Johnston et al. 11770, LL). G. Proximal and distal pinnae. H. Segment.

DRYOPTERIS

823

Fig. 121. A, B. Dryopteris wallichiana (Mickel 6777, NY). A. Pinnae. B. Segments. C, D. Dryopteris karwinskyana (Mickel 4205, NY). C. Pinna. D. Pinnule. E, F. Dryopteris pseudofilix-mas (Mickel 7054, NY). E. Pinnae. F. Segment. G, H. Dryopteris patula (Mickel 5071, NY) (see also Figs. 122H, J). G. Pinna. H. Segment base.

824

DRYOPTERIS

Fig. 122. A–G. Dryopteris cinnamomea (see also Figs. 123E, F). A–C. (Mickel 796, NY). A. Pinnae. B. Stipe base. C. Stipe scale. D. (Mickel 778, NY), pinnae. E–G. (Mickel 1260, NY). E. Pinna. F. Stipe base. G. Stipe scale. H, J. Dryopteris patula (Mickel 1109, NY) (see also Figs. 121G, H). H. Stipe base. J. Stipe scale. K. Dryopteris nubigena (Mickel 5285, NY), pinnae (see also 123G, H). L. Dryopteris cinnamomea (Mickel 4428a, NY), pinnae (see also Figs. 123E, F).

DRYOPTERIS

825

Fig. 123. A–D. Dryopteris rossii (McVaugh 16257, NY). A. Frond. B. Stipe scale. C. Segment detail. D. Indusium. E, F. Dryopteris cinnamomea (Breedlove 59129, NY) (see also Figs. 122A–G, L). E. Segment detail. F. Indusium. G, H. Dryopteris nubigena (Skutch 771, NY, from Guatemala) (see also 122K). G. Proximal pinna base. H. Segment.

826

ELAPHOGLOSSUM

Fig. 124. A–H. Elaphoglossum sartorii. A–C. (Mickel 999, NY) (see also Figs. 126F–H, J, 138L–P). A. Habit. B. Abaxial blade detail. C. Rhizome scale. D. (Mickel 4152, NY), frond. E–G. (Hellwig 218, NY). E. Abaxial blade detail. F, G. Stipe base scales. H. (Mickel 1620, NY), rhizome scale. J, K. Elaphoglossum lepidopodum (Mickel 6175, NY) (see also Figs. 138A–E). J. Rhizome and frond bases. K. Stipe scale. L–N. Elaphoglossum glaucum (Mickel 4336, NY). L. Habit. M. Abaxial blade detail. N. Rhizome scale.

ELAPHOGLOSSUM

827

Fig. 125. A–C. Elaphoglossum leebrowniae (Mickel 1230, NY). A. Habit. B. Rhizome scales. C. Abaxial blade detail. D, E. Elaphoglossum yourkeorum (Mickel 6678, NY). D. Habit. E. Abaxial blade detail. F, G. Elaphoglossum xanthopodum (Purpus 16580, US). F. Habit. G. Rhizome scales.

828

ELAPHOGLOSSUM

Fig. 126. A, B. Elaphoglossum seminudum (Mickel 911, NY). A. Habit. B. Stipe scale. C–E. Elaphoglossum lonchophyllum (Mickel 5706, NY). C. Habit. D. Rhizome scale. E. Venation. F–H, J. Elaphoglossum sartorii (see also Figs. 124A–H, 138L–P). F, G. (Mickel 3774, NY). F. Rhizome and stipe bases. G. Abaxial blade scales. H, J. (McVaugh 22375, NY). H. Rhizome scale. J. Stipe scale. K, L. Elaphoglossum glabellum (Mickel 6522, NY). K. Habit, fertile blade section. L. Rhizome scale. M. Elaphoglossum parduei (Mickel 1544, NY), habit (see also Figs. 131C, D).

ELAPHOGLOSSUM

829

Fig. 127. A–D. Elaphoglossum pallidum. A–C. (Mickel 5325, NY). A. Habit. B. Blade margin. C. Rhizome scale. D. (Mickel 4119a, NY), rhizome scale. E. Elaphoglossum erinaceum var. erinaceum (Mickel 5646, NY), rhizome scale (see also Figs. 139D–F). F–H, J–M. Elaphoglossum squamipes (Mickel 5266, NY). F. Habit. G, H. Rhizome scales. J. Stipe scale. K–M. Blade scales. N–P. Elaphoglossum tenuifolium (Pringle 4916, NY). N. Habit. O. Trichomidium. P. Rhizome scale. Q–S. Elaphoglossum affine (Camp 2683, NY). Q. Habit. R, S. Rhizome scales.

830

ELAPHOGLOSSUM

Fig. 128. A–F. Elaphoglossum muscosum (Mickel 1126, NY). A. Habit. B. Abaxial blade scales. C. Rhizome scale. D–F. Stipe scales. G, H, J–P. Elaphoglossum paleaceum. G–L. (Mickel 5367, NY). G. Habit. H. Abaxial blade scales. J. Blade scale. K. Rhizome scale. L. Stipe scale. M, N. (Mickel 930, NY). M. Blade base. N. Abaxial blade scales. O, P. (Mickel 6562, NY). O. Abaxial blade scales. P. Abaxial blade scales. Q. Elaphoglossum vestitum (Hellwig 319, NY), abaxial blade. R–W. Elaphoglossum pringlei (Anderson & Anderson 4778, NY). R. Habit. S. Abaxial blade. T. Rhizome scale. U, V. Stipe scales. W. Blade scales.

ELAPHOGLOSSUM

831

Fig. 129. A–D. Elaphoglossum tectum (Mickel 5096, NY). A. Habit. B. Rhizome scale. C. Adaxial blade scales. D. Abaxial blade scales. E–G. Elaphoglossum petiolatum (Mickel 3773, NY). E. Rhizome scale. F. Adaxial blade scales. G. Abaxial blade scales. H, J, K. Elaphoglossum ipshookense (Mickel 4748, NY). H. Rhizome scale. J. Adaxial blade scales. K. Abaxial blade scales. L–O. Elaphoglossum muelleri (Mickel 1164, NY). L. Habit. M. Rhizome scale. N. Adaxial blade scales. O. Abaxial blade scales.

832

ELAPHOGLOSSUM

Fig. 130. A–C. Elaphoglossum piloselloides (Mickel 5225, NY). A. Habit. B. Abaxial blade scales. C. Rhizome scale. D–F. Elaphohglossum lanceum (Mickel 4897, NY). D. Habit. E. Adaxial blade detail. F. Rhizome scale. G, H, J. Elaphoglossum setigerum (Mickel 3785, NY). G. Habit. H. Adaxial blade detail. J. Rhizome scale. K. Elaphoglossum peltatum (Mickel 4267, NY), habit. L–N. Elaphoglossum leonardii (Mickel 5296, NY). L. Habit. M. Adaxial blade detail. N. Rhizome scale.

ELAPHOGLOSSUM

833

Fig. 131. A. Elaphoglossum decursivum (Mickel 6417, NY), habit. B. Elaphoglossum guatemalense (Mickel 7287, NY), habit (see also Figs. 138F–K). C, D. Elaphoglossum parduei (Mickel 6615, NY) (see also Fig. 126M). C. Habit. D. Rhizome scale. E. Elaphoglossum squarrosum (Lorence 4042, NY), habit. F–H. Elaphoglossum lindenii (Rzedowski 23278, NY). F. Habit. G. Blade apex. H. Blade scale.

834

ELAPHOGLOSSUM

Fig. 132. A–C. Elaphoglossum gratum (Mickel 7437, NY). A. Habit. B. Blade scales. C. Rhizome scale. D, E. Elaphoglossum auricomum (Müller 386, NY). D. Blade. E. Stipe scales. F–H. Elaphoglossum dissitifrons (Mickel 4034, NY) (see also Figs. 140A–D). F. Habit. G. Stipe base scale. H. Rhizome scale. I–K. Elaphoglossum revolutum (Mickel 7326, NY). I. Habit. J. Blade detail. K. Rhizome scale.

ELAPHOGLOSSUM

835

Fig. 133. A–C. Elaphoglossum albomarginatum (Breedlove 31629, DS). A. Habit. B. Margin and veins. C. Rhizome scale. D–G. Elaphoglossum eximium (Geith 95MO17bis, NY). D. Habit. E. Adaxial blade detail. F, G. Rhizome scales. H, J. Elaphoglossum latum (Seidenschnur 109, NY, from Costa Rica, and Smith 2688a, NY, from Colombia). H. Habit. J. Adaxial blade margin. K. Elaphoglossum erinaceum var. occidentale (Rzedowski 25175, ASC), adaxial blade margin.

836

ELAPHOGLOSSUM

Fig. 134. A–E. Elaphoglossum pilosius (Breedlove 55848, CAS). A. Habit. B. Blade base, adaxial surface. C. Adaxial blade detail. D. Abaxial blade detail. E. Rhizome scale. F–H. Elaphoglossum setosum (Geith 95-MO16bis, NY). F. Habit. G. Fertile blade base. H. Sterile blade detail, adaxial surface. J–L. Elaphoglossum jaliscanum (Stevenson 570, NY). J. Habit. K. Abaxial blade detail. L. Adaxial blade detail. M, N. Elaphoglossum pusillum (Cedillo T. 1896, MEXU). M. Habit. N. Adaxial blade detail. O–Q. Elaphoglossum monicae (Rzedowski 26695, NY). O. Habit. P. Adaxial blade detail. Q. Abaxial blade detail.

ELAPHOGLOSSUM

837

Fig. 135. A–F. Elaphoglossum engelii (Rzedowski 23279, NY, and Molina R. et al. 16539, NY, from Guatemala). A. Habit. B. Fertile blade detail, abaxial surface. C. Abaxial blade scales. D. Adaxial blade scales. E. Stipe scale. F. Rhizome scale. G, H, J, K. Elaphoglossum hartwegii (Rzedowski 23469, MEXU). G. Habit. H. Adaxial blade scales. J. Stipe detail. K. Rhizome scale. L–P. Elaphoglossum rzedowskii (Rzedowski 26161, IEB). L. Habit. M. Abaxial blade scales. N. Adaxial blade scales. O. Blade scale. P. Rhizome scale. Q–S. Elaphoglossum huacsaro (Breedlove 22412, NY). Q. Habit. R. Abaxial blade scales. S. Rhizome scale.

838

ELAPHOGLOSSUM

Fig. 136. A–E. Elaphoglossum rufescens. A–D. (Hinton 2750, NY). A. Habit. B. Abaxial blade detail. C. Rhizome scale. D. Stipe scale. E. (Balls B4469, UC), habit. F–H, J–L. Elaphoglossum manantlanense (Wilbur & Wilbur 1971, US). F. Habit. G. Stipe detail. H. Rhizome scale. J. Fertile blade detail. K. Sterile blade detail, abaxial surface. L. Blade scale. M–P. Elaphoglossum obscurum (Ventura A. 309a, NY). M. Habit. N. Stipe detail. O. Rhizome scale. P. Abaxial blade detail.

ELAPHOGLOSSUM

839

Fig. 137. A–C. Elaphoglossum mcvaughii (McVaugh 9904, MICH). A. Habit. B. Stipe scale. C. Rhizome scale. D, E. Elaphoglossum herminieri (Jones & Facey 3292, NY, from Guatemala). D. Habit (fertile frond (Shafer 3068, NY, from Cuba)). E. Rhizome scale. F, G, H, J. Elaphoglossum rubescens (Johnson 541, NY, from Guatemala). F. Habit. G. Abaxial blade detail. H. Stipe detail. J. Rhizome scale.

840

ELAPHOGLOSSUM

Fig. 138. A–E. Elaphoglossum lepidopodum (see also Figs. 124J, K). A–D. (Rzedowski s.n., 22 Sep 1965, ENCB). A. Habit. B. Blade base detail, abaxial surface. C. Rhizome scale. D. Stipe scale. E. (Heath & Long 1045, MEXU), stipe detail. F–H, J, K. Elaphoglossum guatemalense (Riba et al. 304, MEXU) (see also Fig. 131B). F. Habit. G. Rhizome scale. H. Stipe detail. J. Abaxial blade detail. K. Stipe scale. L–P. Elaphoglossum sartorii (see also Figs. 124A–H, 126F–H, J). L, M. (Rose 2215, US). L. Habit. M. Rhizome scale. N–P. (Mickel 6677, NY). N. Habit. O. Abaxial blade base detail. P. Rhizome scale.

ELAPHOGLOSSUM

841

Fig. 139. A–C. Elaphoglossum potosianum (Parry & Palmer 1007, NY). A. Habit. B. Blade detail, adaxial surface. C. Blade detail, abaxial surface. D–F. Elaphoglossum erinaceum var. erinaceum (Mickel 5646, NY, and Breedlove 29815, NY) (see also Fig. 127E). D. Habit. E. Blade detail, abaxial surface. F. Rhizome scale.

842

ELAPHOGLOSSUM

Fig. 140. A–D. Elaphoglossum dissitifrons (Rzedowski & McVaugh 174, NY) (see also Figs. 132F–H). A. Habit. B. Stipe detail, scales. C. Rhizome scales. D. Abaxial blade detail. E–H, J. Elaphoglossum tenuiculum (Anderson 13212, NY). E. Habit. F. Stipe detail. G. Rhizome scales. H. Adaxial blade detail. J. Abaxial blade detail.

ELAPHOGLOSSUM

843

Fig. 141. A–E. Elaphoglossum viride (Mickel 1545, NY). A. Habit. B. Abaxial blade base. C. Rhizome. D. Rhizome scales. E. Rhizome scale detail. F–H, J, K. Elaphoglossum alansmithii (Mickel 3739, NY). F. Habit. G. Abaxial blade detail. H. Trichomidia. J. Rhizome scale. K. (Mickel 3735, NY), habit.

844

ENTEROSORA

Fig. 142. A, B. Enterosora ecostata (Breedlove 49972, CAS). A. Habit. B. Blade detail abaxial surface. C. Enterosora trichosora (Türckheim II1987, UC, from Guatemala), frond. D, E. Enterosora parietina (Mickel 5739, NY). D. Habit. E. Blade detail, abaxial surface.

EQUISETUM

845

Fig. 143. A–E. Equisetum hyemale var. affine. A–D. (Rzedowski 22193, NY) (see also Fig. 144 F, G, a). A. Habit. B. Node with persistent leaves. C. Node with deciduous leaves. D. Strobilus. E. (McVaugh 20021, MICH), rhizome with bases of aerial stems. F–H, J. Equisetum myriochaetum (Mickel 4225, NY) (see also Figs. 144H, J, b). F. Stem with fertile branch whorls. G. Ridge tubercles. H. Node with whorl of vegetative branches. J. Strobilus.

846

EQUISETUM

Fig. 144. A–E. Equisetum laevigatum (Clausen 4622, NY, from New Mexico) (see also Fig. 144c). A. Stem. B. Cone. C–E. Leaf sheaths. F, G. Equisetum hyemale var. affine (Pringle s.n., 23 May 1888, NY), leaf sheaths (see also Figs. 143A–E, 144a). F. Leaf sheath without teeth. G. Leaf sheath with teeth. H, J. Equisetum myriochaetum (Mickel 4225, NY) (see also Figs. 143F–H, J, 144b). H. Leaf sheath without teeth. J. Leaf sheath with teeth. a–f. Diagramatic stem cross sections of three species and three hybrids. (Figs. a–f reprinted from Hauke, 1963: t. 2, Figs. d, e, f, ed, af, ef, with permission of author.). a. Equisetum hyemale var. affine. b. Equisetum myriochaetum. c. Equisetum laevigatum. d. Equisetum ×schaffneri. e. Equisetum ×haukeanum. f. Equisetum ×ferrissii.

ERIOSORUS

847

Fig. 145. A, B. Eriosorus hirtus (Mickel 4239, NY). A. Frond. B. Segment. C, D. Eriosorus flexuosus var. flexuosus (Mickel 6744, NY). C. Pinna. D. Segment. E, F. Eriosorus flexuosus var. galeanus (Rzedowski & McVaugh 125, NY). E. Pinna. F. Segment.

848

GLEICHENELLA

Fig. 146. A–E. Gleichenella pectinata. A–C. (Hinton 7543, NY). A. Portion of blade. B. Ultimate fork. C. Penultimate fork. D, E. (Mickel 1444, NY). D. Segments. E. Sorus.

HECISTOPTERIS

Fig. 147. A–C. Hecistopteris pumila (Mickel 6440, NY). A. Habit. B. Blade tip. C. Sori.

849

850

HEMIDICTYUM

Fig. 148. A, B. Hemidictyum marginatum (Mickel 6381, NY). A. Pinna. B. Pinna detail.

HEMIONANTHES

Fig. 149. A–C. Hemionanthes gryphus (McVaugh 15908, MICH). A. Habit. B. Rhizome scale. C. Pinnule.

851

852

HEMIONITIS

Fig. 150. A. Hemionitis pinnatifida (Seiler 505, NY, from El Salvador), habit. B. Hemionitis levyi (Mickel 5976, NY), habit. C. Hemionitis palmata (Mickel 6877, NY), habit.

HEMIONITIS

Fig. 151. A, B. Hemionitis rufa (Matuda 3415, MICH). A. Habit. B. Pinna base. C, D. Hemionitis subcordata (Mickel 5151, NY). C. Frond. D. Pinna detail.

853

854

HISTIOPTERIS

Fig. 152. A–C. Histiopteris incisa (Mickel 1065, NY). A. Pinna. B. Rhizome. C. Segment.

HOLODICTYUM

Fig. 153. A–D. Holodictyum ghiesbreghtii (Palmer 336, NY). A. Habit. B. Blade detail, showing sori. C. Blade detail, showing hairs, veins. D. Rhizome scale.

855

856

HUPERZIA

Fig. 154. A. Huperzia hippuridea (Mickel 4898, NY), habit. B. Huperzia crassa (Sharp s.n., 26 Dec 1970, NY), habit. C, D. Huperzia pithyoides (Mickel 1095, NY). C. Habit. D. Fertile area. E, F. Huperzia reflexa (Mickel 4803, NY). E. Habit. F. Fertile area.

HUPERZIA

857

Fig. 155. A. Huperzia myrsinites (Mickel 4257, NY), habit. B. Huperzia pringlei (Mickel 7051, NY), habit (see also Figs. 158A–E). C. Huperzia cuernavacensis (Mickel 3945, NY), habit.

858

HUPERZIA

Fig. 156. A. Huperzia taxifolia (Hallberg 1484, NY), habit. B. Huperzia wilsonii (Hallberg 1416, NY), habit. C. Huperzia linifolia (Mickel 6683, NY), habit.

HUPERZIA

859

Fig. 157. A–C. Huperzia serrata (Ekman 14542, NY, from Hispaniola). A. Habit. B. Fertile area. C. Leaf. D–H. Huperzia beiteliana (Mickel 9650, NY). D. Habit. E. Fertile area. F. Leaf. G. Gemma, dorsal (left) and ventral views. H. Gemmaphore, dorsal (left) and lateral views. J–L. Huperzia orizabae. J. (Müller s.n., in June 1854, NY), habit. K. (Hallberg 1404a, NY), fertile area. L. (Müller s.n., in 1855, NY), branch.

860

HUPERZIA

Fig. 158. A–E. Huperzia pringlei (see also Fig. 155B). A–D (as H. tuerckheimii). A. (Molina R. 11936, NY, from Honduras), habit. B–D. (Türckheim 1864, NY, from Guatemala) B. Sterile area. C. Fertile area. D. Sporophyll. E. (Pringle 2974, NY, typical H. pringlei), sporophyll. F–H. Huperzia capillaris (Breedlove 29630, DS). F. Habit. G. Sterile area. H. Fertile area. J, K. Huperzia dichotoma (Pringle 3976, NY). J. Habit. K. Fertile area.

HYMENOPHYLLUM

861

Fig. 159. A, B. Hymenophyllum hirsutum (Mickel 1052, ISC). A. Habit. B. Pinna. C, D. Hymenophyllum crispum (Mickel 5288, NY). C. Habit. D. Pinna. E, F. Hymenophyllum microcarpum (Mickel 6617, NY). E. Habit. F. Pinnule. G, H. Hymenophyllum pulchellum (Mickel 1047, NY). G. Habit. H. Pinna.

862

HYMENOPHYLLUM

Fig. 160. A, B. Hymenophyllum trapezoidale (Mickel 1045, NY). A. Habit. B. Pinna. C, D. Hymenophyllum fragile (Mickel 4252a, NY). C. Habit. D. Frond. E, F. Hymenophyllum tegularis (Mickel 922, ISC). E. Habit. F. Pinna. G, H. Hymenophyllum lanatum (Hellwig 499, NY). G. Habit. H. Pinna.

HYMENOPHYLLUM

863

Fig. 161. A, B. Hymenophyllum asplenioides (Hallberg 1435, NY). A. Habit. B. Segment. C, D. Hymenophyllum myriocarpum (Mickel 918, NY) (see also Fig. 163M). C. Habit. D. Segment. E–G. Hymenophyllum fendlerianum (Mickel 1041, ISC). E. Habit. F. Segment. G. Sorus. H, J. Hymenophyllum brevistipes (Mickel 6532, NY) (see also Figs. 163C–E). H. Habit. J. Blade apex.

864

HYMENOPHYLLUM

Fig. 162. A, B. Hymenophyllum polyanthos (Mickel 5584, NY). A. Habit. B. Pinnule. C, D. Hymenophyllum tunbrigense (Mickel 5286, NY). C. Habit. D. Pinna. E, F. Hymenophyllum fucoides (Mickel 3781a, NY). E. Habit. F. Pinna. G. Hymenophyllum ectocarpon (Mickel 4146, NY), pinna detail. H, J. Hymenophyllum polyanthos (Mickel 1046, ISC). H. Habit. J. Pinna.

HYMENOPHYLLUM

865

Fig. 163. A, B. Hymenophyllum abruptum (Clute 54, NY, from Jamaica). A. Habit. B. Blade base. C–E. Hymenophyllum brevistipes (Breedlove 57305, CAS, NY) (see also Figs. 161H, J). C. Habit. D. Frond. E. Stipe. F–H. Hymenophyllum paucicarpum (Breedlove 36969, DS). F. Habit. G. Frond. H. Sorus. J–L. Hymenophyllum undulatum (Breedlove 58210, NY, and Mickel 1107, NY). J. Habit. K. Pinna. L. Sorus. M. Hymenophyllum myriocarpum (Mickel 4144, NY), sorus (see also Figs. 161C, D).

866

HYMENOPHYLLUM

Fig. 164. A–D. Hymenophyllum crassipetiolatum (Calzada et al. 8773, NY). A. Habit. B. Stipe detail. C. Blade detail. D. Sorus. E–H. Hymenophyllum maxonii (Ventura A. 205, IEB). E. Habit. F. Blade detail. G. Marginal hairs. H. Sorus. J–L. Hymenophyllum sieberi (Münch 11, US). J. Habit. K. Blade detail. L. Sorus.

HYPOLEPIS

867

Fig. 165. A–D. Hypolepis trichochlaena (Mickel 4322, NY). A. Rachis and pinnule. B. Rachis. C. Segment detail, abaxial surface. D. Indusium. E–G. Hypolepis nigrescens (Mickel 1029, NY). E. Rachis and pinnule. F. Rachis. G. Segment detail, abaxial surface. H, J–M. Hypolepis thysanochlaena. H, J–K (Mickel 1150, NY). H. Pinnule. J. Rachis. K. Segment detail, abaxial surface. L, M. (Mickel 5360, NY). L. Segment detail, abaxial surface. M. Indusium. N–P. Hypolepis repens (Mickel 5896, NY). N. Pinnule. O. Rachis. P. Segment detail, abaxial surface. Q–T. Hypolepis blepharochlaena (Mickel 4151, NY). Q. Pinnule. R. Rachis. S. Segment detail. T. Indusium.

868

HYPOLEPIS

Fig. 166. A–C. Hypolepis melanochlaena (Münch 35, DS). A. Pinna bases. B. Rachis. C. Segment. D–F. Hypolepis pulcherrima (Münch 111, DS). D. Pinna base. E. Segment. F. Indusium. G, H, J. Hypolepis stuebelii (Velázquez 281, FCME). G. Pinna base. H. Stipe. J. Segment base. K, L. Hypolepis muenchii. K. (Mickel 4632, NY), segment. L. (Breedlove 22777, DS), segment. M, N. Hypolepis microchlaena (Mickel 5657, NY). M. Segment. N. Segment, with sorus removed.

ISOËTES

869

Fig. 167. A–C. Isoëtes mexicana (McVaugh 13018, MICH). A. Habit. B. Leaf base, showing sporangium, megaspores, ligule, and velum. C. Megaspores. D. Isoëtes pringlei (Pringle 3333, NY), megaspores.

870

ISOËTES

Fig. 168. A–G. Isoëtes megaspores all at the same magnification. A. Isoëtes howellii (Brandegee s.n., Nov. 1902, UC), megaspores average 397 m diam. B. Isoëtes mexicana (Rzedowski & McVaugh 483, NY), megaspores average 359 m diam. C. Isoëtes montezumae (Taylor s.n., 9 Nov 1997, MIL), megaspores average 436 m diam. D. Isoëtes orcuttii (Howell 4175, UC), megaspores average 320 m diam. E. Isoëtes pallida (Kral 25320, MICH), megaspores average 413 m diam. F. Isoëtes pringlei (Pringle 3333, NY), megaspores average 503 m diam. G. Isoëtes mexicana × montezumae (Taylor s.n., 9 Nov 1997, MIL), megaspores range from less than 200 m to more than 800 m diam. H. Adaxial view of leaf bases showing velum coverage. Isoëtes orcuttii, left, with velum covering 100% of sporangium (a small portion of the velum peeled back near the leaf base to expose sporangium wall beneath) and I. howellii, right, with velum covering less than 25% of sporangium. Arrows indicate extent of velum.

JAMESONIA

Fig. 169. A–C. Jamesonia alstonii (Mickel 5513, NY). A. Habit. B. Pinnae. C. Pinna, adaxial surface.

871

872

LASTREOPSIS

Fig. 170. A–E. Lastreopsis effusa subsp. dilatata. A. (Hallberg 1288, NY), proximal pinna base. B, C. (Hallberg 1325, NY). B. Frond apex. C. Rachis bud. D, E. (Hallberg 1288, NY). D. Segment. E. Rachis section. F, G. Lastreopsis exculta subsp. guatemalensis (Purpus 6724, UC). F. Pinnae. G. Segment. H. Lastreopsis effusa subsp. divergens (Breedlove 38683, DS), segment.

LELLINGERIA

873

Fig. 171. A, B. Lellingeria hellwigii (Mickel 4324, NY). A. Habit. B. Fertile blade detail. C–E. Lellingeria limula (Breedlove 30808, NY). C. Habit. D. Fertile blade detail. E. Blade, adaxial surface. F, G. Lellingeria prionodes (Mickel 923a, NY). F. Habit. G. Fertile blade detail. H, J. Lellingeria delitescens (Hallberg 1729, NY). H. Habit. J. Fertile blade detail. K–M. Lellingeria mitchellae (Breedlove 48307, NY). K. Habit. L. Fertile blade detail. M. Sporangium. N, O. Lellingeria apiculata (Seiler 737, NY, from El Salvador). N. Fertile pinnae. O. Rhizome scale.

874

LINDSAEA

Fig. 172. A–C. Lindsaea stricta. A, B. (Téllez V. 6151, MEXU). A. Blade. B. Segment detail. C. (Martinez S. 15003, IEB), sorus. D–H. Lindsaea quadrangularis subsp. subalata. D. (Schipp 200, NY, from Belize), rachilla and segment bases. E–H. (Mickel 6517, NY). E. Pinna. F. Rachis. G. Segment. H. Sorus. J–M. Lindsaea lancea (Mickel 1048, NY). J. Pinna. K. Rachis. L. Segment. M. Sorus. N–P. Lindsaea arcuata (Mickel 1049, NY). N. Pinna. O. Segment. P. Sorus. Q–S. Lindsaea stricta (Mickel 7372, NY). Q. Habit. R. Pinna. S. Sorus.

LINDSAEA

875

Fig. 173. A–C. Lindsaea quadrangularis subsp. mexiae (Mexia 1837, MICH). A. Habit. B. Segment. C. Rhizome scale. D–F. Lindsaea klotzschiana (Nee & Calzada 22170, NY). D. Rachis and pinnae bases, abaxial surface. E. Rachis and pinna bases, adaxial surface. F. Segment detail, abaxial surface.

876

LLAVEA

Fig. 174. A–D. Llavea cordifolia (Mickel 3872, NY). A. Habit. B. Fertile segment detail. C, D. Rhizome scales.

LOMARIOPSIS

877

Fig. 175. A–C. Lomariopsis mexicana (Breedlove 33838, DS). A. Habit. B. Sterile pinna detail. C. Rhizome scale. D, E. Lomariopsis vestita. D. Sterile frond (Wendt 3822, NY). E. Fertile pinnae (Mickel 2789, NY, from Costa Rica). F. Lomariopsis recurvata (Mickel 7171, NY), sterile pinnae.

878

LONCHITIS

Fig. 176. A–C. Lonchitis hirsuta (Mickel 5233, NY). A. Pinna. B. Segment, abaxial surface. C. Segment, adaxial surface.

LOPHOSORIA

Fig. 177. A–C. Lophosoria quadripinnata (Mickel 1253, NY). A. Pinna base and stipe base. B. Segment. C. Segment detail.

879

880

LOXOGRAMME

Fig. 178. A–E. Loxogramme mexicana (Mickel 949b, NY). A. Habit. B. Rhizome. C. Rhizome scale. D. Blade detail. E. Cross section of blade.

LOXOSCAPHE

Fig. 179. A–C. Loxoscaphe theciferum (Ekman H6784, UC, from Dominican Republic). A. Habit. B. Segments, adaxial surface. C. Segments, abaxial surface.

881

882

LYCOPODIELLA

Fig. 180. A–D. Lycopodiella caroliniana. (Cowan 2233, NY). A. Habit. B. Rhizome. C. Erect stem detail. D. Strobilus detail. E–H. Lycopodiella alopecuroides (Breedlove 15165, DS). E. Habit. F. Rhizome. G. Erect stem detail. H. Strobilus detail. J–L. Lycopodiella cernua (Mickel 5870, NY). J. Habit. K. Stem detail. L. Strobilus.

LYCOPODIUM

883

Fig. 181. A–C. Lycopodium thyoides (Mickel 3940, NY). A. Habit. B. Upper surface of branch. C. Lower surface of branch. D, E. Lycopodium clavatum (Mickel 6766, NY). D. Habit. E. Leaves.

884

LYGODIUM

Fig. 182. A–D. Lygodium volubile. A. (Lot 1244, MEXU), pinnule. B–D. (Münch 98, MICH). B. Segment base. C. Fertile projection detail, abaxial surface. D. Fertile projection detail, adaxial surface. E. Lygodium heterodoxum (Hallberg 1302, NY), pinnule. F–H. Lygodium venustum. F–G. (Mickel 5961, NY). F. Pinna. G. Segment. H. (Martínez S. 16980, MEXU), pinna.

MACROTHELYPTERIS

Fig. 183. A–C. Macrothelypteris torresiana (Mickel 5921, NY). A. Pinna. B. Blade detail, abaxial surface. C. Stipe cross section.

885

886

MARATTIA

Fig. 184. A, B. Marattia excavata (Breedlove 22649, NY). A. Pinnule. B. Segment. C–F. Marattia weinmanniifolia. C. Rhizome and stipules (from slide). D–F. (Mickel 5264, NY) D. Pinnules. E. Segment. F. Synangium. G, H, J. Marattia laxa (Mickel 4117, NY). G. Pinnules. H. Pinnule detail. J. Synangium.

MARSILEA

887

Fig. 185. A–C. Marsilea polycarpa. A. Fertile leaves. B. Floating plant. C. Sporocarp. D–G. Marsilea deflexa. D. Sporocarp. E. Plant with floating leaves. F. Abaxial surface of floating leaf, showing hydropoten. G. Land leaf with sporocarps. H, J. Marsilea crotophora. H. Sporocarp. J. Habit. (Scale: short bar—A, B, E, F, G, J; long bar—C, D, H.) (Reprinted from Johnson (1986, Fig. 13) with permission.)

888

MARSILEA

Fig. 186. A–D. Marsilea macropoda. A. Habit. B. Sporocarps borne on branched peduncle. C. Sporocarps with indument removed to show variation in shape. D. Variation in attachment of sporocarps. E, F. Marsilea ancylopoda. E. Habit. F. Sporocarps to show variation in shape and in orientation of peduncle. (Scale: short bar—A, E; long bar—B, C, F.) (Reprinted from Johnson (1986, Fig. 15) with permission.)

MARSILEA

889

Fig. 187. A, B. Marsilea vestita. A. Habit of plant from California. B. Five sporocarps to show variation in size and in length of peduncle. C, D. Marsilea mollis. C. Two leaves. D. Two sporocarps from the same collection, the larger from a plant in cultivation. (Scale: short bar—A, C; long bar—B, D.) (Reprinted with alterations from Johnson (1986, Fig. 17) with permission.)

890

MEGALASTRUM

Fig. 188. A, B. Megalastrum lunense (Skutch 3113, NY, from Costa Rica). A. Proximal pinna. B. Costa detail. C–E. Megalastrum atrogriseum (Hernández G. 1414, NY). C. Proximal pinna. D. Rhizome scale. E. Segment detail. F, G, H, J. Megalastrum subincisum (Mickel 1445, NY). F. Pinna base. G. Pinnule base. H. Pinnule bases, abaxial surface. J. Costa scale. K–M. Megalastrum pulverulentum (Breedlove 31677, NY). K. Base of proximal pinna. L. Segment bases, abaxial surface. M. Costal scales.

MELPOMENE

891

Fig. 189. A–D. Melpomene zempoaltepetlensis (Mickel 4638, NY). A. Habit. B. Stipe detail. C. Fertile pinna, abaxial surface. D. Sterile pinnae, adaxial surface. E–G. Melpomene firma (Breedlove 33665, DS). E. Habit. F. Pinna, abaxial surface. G. Pinna, adaxial surface. H, J. Melpomene melanosticta (Breedlove 33014, DS). H. Habit. J. Fertile pinnae, abaxial surface. K, L. Melpomene xiphopteroides (Mickel 4811, NY). K. Habit. L. Fertile pinnae, abaxial surface.

892

MELPOMENE

Fig. 190. A, B. Melpomene leptostoma (Mickel 5737, NY). A. Habit. B. Pinnae. C, D. Melpomene deltata (Mickel 4637a, NY). C. Habit. D. Pinnae. E, F. Melpomene moniliformis (Mickel 4924, NY). E. Habit. F. Pinnae. G, H. Melpomene flabelliformis (Mickel 6786, NY). G. Habit. H. Pinnae. J, K. Melpomene pilosissima (Mickel 5252, NY). J. Habit. K. Pinnae.

METAXYA

Fig. 191. A, B. Metaxya rostrata (Breedlove 34100, DS). A. Pinna. B. Pinna detail.

893

894

MICROGRAMMA

Fig. 192. A, B. Microgramma piloselloides (Morton & Acuña 3412, UC, from Cuba). A. Habit. B. Blade detail. C, D. Microgramma reptans (Rovirosa 859, NY). C. Habit. D. Blade detail. E–G. Microgramma nitida (Barlow 14/2, NY). E. Habit. F. Rhizome scale. G. Blade detail. H, J, K. Microgramma tecta var. nana (Holm & Iltis 193, NY, from Costa Rica). H. Habit. J. Blade detail. K. Blade scale. L. Microgramma lycopodioides (Wendt et al. 2596, NY), habit. M–O. Microgramma percussa (Wendt et al. 2634, NY). M. Habit. N. Blade detail. O. Blade scales.

MICROPOLYPODIUM

895

Fig. 193. A–C. Micropolypodium trichomanoides (Breedlove 32225A, NY). A. Habit. B. Blade detail. C. Rhizome scale. D, E. Micropolypodium basiattenuatum (Mickel 3824, NY). D. Habit. E. Blade detail. F, G. Micropolypodium taenifolium (Mickel s.n., 24–31 July 1962, NY). F. Habit. G. Blade detail.

896

MILDELLA

Fig. 194. A–D. Mildella fallax (Mickel 4518, NY). A. Habit. B. Rhizome scale. C, D. (Mickel 4704, NY). C. Pinnae. D. Segment. E, F. Mildella intramarginalis (Mickel 4518, NY). E. Pinnae. F. Segment.

NEPHROLEPIS

897

Fig. 195. A–C. Nephrolepis undulata (Mickel 3981, NY). A. Habit. B. Pinna. C. Sorus. D–F. Nephrolepis hirsutula (Hellwig 354, NY) (see also Figs. 196N–R). D. Pinnae. E. Pinna base. F. Sorus. G, H, J–L. Nephrolepis pectinata (Mickel 1580, NY). G. Rhizome and frond bases. H. Pinnae. J. Pinna. K. Blade detail, adaxial surface. L. Sorus. M–O. Nephrolepis rivularis (Hallberg 1589, NY). M. Pinnae. N. Pinna base. O. Sorus.

898

NEPHROLEPIS

Fig. 196. A–C. Nephrolepis pendula (Breedlove 48362, NY). A. Pinnae. B. Pinna. C. Sorus. D–G. Nephrolepis cordifolia. D–F. (Rzedowski 29996, NY). D. Sterile pinnae. E. Rachis detail. F. Rachis scale. G. (Mexia 9274, NY), sorus. H, J–M. Nephrolepis biserrata (Hernández G. 1692, NY). H. Pinnae. J. Sorus. K. Stipe base. L. Rachis and pinna base, adaxial surface. M. Rachis and pinna base, abaxial surface. N–R. Nephrolepis hirsutula (Hellwig 455, NY) (see also Figs. 195D–F). N. Pinnae. O. Stipe base. P. Rachis and pinna base, adaxial surface. Q, R. Sori. S–W. Nephrolepis exaltata (Breedlove 23496, DS). S. Pinnae. T. Rachis, abaxial surface. U. Pinna base scale. V, W. Sori.

NEURODIUM

Fig. 197. A–D. Neurodium lanceolatum (Breedlove 49135, CAS). A. Habit. B. Rhizome. C. Rhizome scale. D. Blade detail.

899

900

NIPHIDIUM

Fig. 198. A–F. Niphidium crassifolium. A–E. (Mickel 5228, NY). A. Frond. B. Blade base. C. Blade detail. D, E. Rhizome scales. F. (Mickel 1013, ISC), frond.

NOTHOLAENA

901

Fig. 199. A–C. Notholaena meridionalis (Mickel 4214, NY). A. Habit. B. Rhizome scale. C. Pinna base. D–H, J. Notholaena brevistipes, (Mayfield et al. 786, UC). D. Habit. E. Rhizome scale. F. Pinna, abaxial surface. G. Costular hair. H. Pinna, adaxial surface. J. Blade hair. K–M. Notholaena rigida (Pringle 2599, MEXU). K. Habit. L. Rhizome scale. M. Pinnule, abaxial surface.

902

NOTHOLAENA

Fig. 200. A–D. Notholaena rosei (Mickel 6240, NY) (see also Figs. 206F–H). A. Habit. B. Rhizome scale. C. Stipe section, adaxial surface. D. Pinna. E–H. Notholaena leonina (Rodríguez L. 433, IEB). E. Habit. F. Rhizome scale. G. Pinna. H. Segment detail. J, K. Notholaena lemmonii var. lemmonii (Brandegee 659, NY). J. Blade. K. Pinna. L–P. Notholaena lemmonii var. australis (Mickel 844, NY). L. Habit. M. Stipe section. N. Rhizome scale. O. Pinnae. P. Segments.

NOTHOLAENA

903

Fig. 201. A–E. Notholaena galeottii (Mickel 4502, NY). A. Habit. B. Rhizome scale. C. Pinna. D. Pinna base. E. Laminar scales from abaxial surface. F–H, J. Notholaena schaffneri. F. (Freytag & Baxter 59, UC), pinna. G, H, J. (McVaugh 25806, NY). G. Pinnules, abaxial surface. H. Pinnules, adaxial surface. J. Rhizome scale. K–N. Notholaena aschenborniana (Pringle s.n., 22 Sep 1885, NY). K. Habit. L. Rhizome scale. M. Pinnules, abaxial surface. N. Pinnules adaxial surface.

904

NOTHOLAENA

Fig. 202. A–F. Notholaena bryopoda (Dorr 2612, NY). A. Habit. B. Segment adaxial surface. C. Segment, abaxial surface. D. Pinna. E. Frond. F. Rhizome scale. G, H, J. Notholaena greggii (Pringle 857, NY). G. Segments, abaxial surface. H. Segments, adaxial surface. J. Rhizome scale. K–N. Notholaena neglecta (Cowan 3804, UC). K. Habit. L. Segments, abaxial surface. M. Segments, adaxial surface. N. Rhizome scale. O–R. Notholaena californica subsp. californica (Orcutt 1461, NY). O. Habit. P. Segments, abaxial surface. Q. Segments, adaxial surface. R. Rhizome scale.

NOTHOLAENA

905

Fig. 203. A–D. Notholaena candida (Gentry et al. 19332, NY). A. Habit. B. Pinnules. C. Rhizome scale. D. Stipe base scale. E. Notholaena copelandii (Pringle 2038, NY), habit. F, G. Notholaena jacalensis (Pray 3095, UC). F. Habit. G. Segments.

906

NOTHOLAENA

Fig. 204. A–D. Notholaena standleyi. A–C. (Newberry s.n., Dec 1882, NY). A. Habit. B. Rhizome scale. C. Blade detail. D. (Bates et al. 1498, NY), habit. E–G. Notholaena sulphurea (Pray 3063, NY). E. Habit. F. Rhizome scale. G. Blade detail. H, J, K. Notholaena affinis (Pray 3104, NY). H. Habit. J. Pinnae. K. Rachis and pinna base. L–N. Notholaena brachycaulis (Clausen 7619, US). L. Habit. M. Rhizome scale. N. Blade detail.

NOTHOLAENA

907

Fig. 205. A–G. Notholaena aurantiaca (Pringle 1840, NY). A. Habit. B. Rhizome scale. C. Pinna, abaxial surface. D. Blade detail, abaxial surface. E. Blade scale. F. Pinna, adaxial surface. G. Blade detail, adaxial surface.

908

NOTHOLAENA

Fig. 206. A–E. Notholaena weatherbiana (Palmer 215, NY). A. Habit. B. Segments, adaxial surface. C. Segments, abaxial surface. D. Segment detail, abaxial surface, with scales removed. E. Rhizome scale. F–H. Notholaena rosei (Mickel 6896, NY) (see also Figs. 200A–D). F. Habit. G. Segments and marginal detail. H. Rhizome scale. J–M. Notholaena aliena (Bartlett 10999, MICH). J. Habit. K. Segments. L. Segment detail, with scales removed. M. Rhizome scale. N–Q. Notholaena grayi (Pringle 463, NY). N. Habit. O. Segments. P. Segment detail, with scales removed. Q. Rhizome scale.

ODONTOSORIA

909

Fig. 207. A–C. Odontosoria schlechtendalii (Mickel 1027, NY). A. Pinnule. B. Blade detail. C. Sorus. D–F. Odontosoria guatemalensis (Breedlove 31621, NY). D. Pinnule. E. Blade detail. F. Sorus.

910

OLEANDRA

Fig. 208. A–C. Oleandra articulata (Wendt et al. 3846, NY). A. Habit. B. Blade detail. C. Blade scale.

OLFERSIA

Fig. 209. A–C. Olfersia cervina (Mickel 3425, NY, from Costa Rica). A. Pinna. B. Venation. C. Base of fertile pinna.

911

912

ONOCLEOPSIS

Fig. 210. A–D. Onocleopsis hintonii. A, B. (McVaugh 26115, MICH). A. Frond. B. Blade detail. C, D. (Mickel 4162, NY). C. Fertile pinnae. D. Fertile segments.

OPHIOGLOSSUM

913

Fig. 211. A. Ophioglossum reticulatum (Hallberg 1780, NY), habit. B, C. Ophioglossum vulgatum (Pringle 8924, NY). B. Habit. C. Venation. D, E. Ophioglossum engelmannii (Pringle s.n., 15 Aug 1893, NY). D. Habit. E. Venation. F, G. Ophioglossum vulgatum (Mickel 9639, NY). F. Habit. G. Venation. H. Ophioglossum palmatum (Small s.n., 2 Apr 1916, NY, from Florida), habit.

914

OPHIOGLOSSUM

Fig. 212. A. Ophioglossum californicum (Jones 3750, NY), habit. B. Ophioglossum polyphyllum (Tryon & Tryon 58283, US), habit. C. Ophioglossum nudicaule (large form) (Romero-Rojas 1553, MEXU), habit. D. Ophioglossum nudicaule (small form) (Pringle 7086, NY), habit. E, F. Ophioglossum crotalophoroides (McVaugh 21682, NY). E. Habit. F. Venation.

OSMUNDA

915

Fig. 213. A–C. Osmunda regalis var. spectabilis (McVaugh 26416, MICH). A. Habit. B. Sterile pinnule. C. Fertile segment detail. D, E. Osmunda cinnamomea (Mickel 902, ISC). D. Sterile pinnae. E. Fertile pinnae.

916

PECLUMA

Fig. 214. A, B. Pecluma alfredii var. occidentalis (Moran 25502, UC). A. Habit. B. Blade detail. C, D. Pecluma alfredii var. cupreolepis (Mickel 5013, NY) (see also Fig. 215M). C. Pinnae. D. Blade detail. E, F. Pecluma plumula (Mickel 6115, NY). E. Pinnae. F. Blade detail. G, H. Pecluma ptilodon var. bourgeauana (Mickel 4819, NY). G. Pinnae. H. Blade detail. J, K. Pecluma atra (Mickel 6823, NY). J. Pinnae. K. Blade detail. L, M. Pecluma consimilis (Mickel 6518, NY) (see also Fig. 215L). L. Pinnae. M. Blade detail. N, O. Pecluma ferruginea (Mickel 6969, NY). N. Habit. O. Blade detail.

PECLUMA

917

Fig. 215. A–D. Pecluma hygrometrica (Breedlove 28505, NY). A. Habit. B. Pinna. C. Blade detail, abaxial surface. D. Sporangium. E, F. Pecluma sursumcurrens (Copeland herb. 128, UC). E. Pinnae. F. Blade detail, abaxial surface. G, H, J. Pecluma divaricata (Breedlove 27939, DS). G. Blade base. H. Rhizome. J. Rhizome scale and detail. K. Pecluma sursumcurrens (King 4354, NY), blade base. L. Pecluma consimilis (Mickel 6518, NY), blade base (see also Figs. 214L, M). M. Pecluma alfredii var. cupreolepis (Rzedowski & de la Sota 18336, NY), rachis scales (see also Figs. 214C, D). N, O. Pecluma dispersa (Breedlove 26041, NY). N. Blade base. O. Rachis scale.

918

PELLAEA

Fig. 216. A. Pellaea sagittata (Mickel 4438, NY), habit. B. Pellaea oaxacana (Mickel 6279, NY), pinnae. C. Pellaea ovata (Mickel 6475, NY), habit. D. Pellaea cordifolia (Mickel 4091, NY), pinna.

PELLAEA

919

Fig. 217. A–C. Pellaea notabilis (Windham 490, NY). A. Habit. B. Pinnae. C. Rhizome scale. D–F. Pellaea pringlei (Hinton 9475, NY). D. Habit. E. Pinnule. F. Rhizome scale. G, H, J, K. Pellaea andromedifolia (Franklin & Chandler 5648, NY). G. Habit. H. Segment, adaxial surface. J. Segment, abaxial surface. K. Rhizome scale. L–O. Pellaea intermedia (Pringle 930, NY). L. Pinna. M. Pinna axes, adaxial surface. N. Segment base, abaxial surface. O. Rhizome.

920

PELLAEA

Fig. 218. A–H, J. Pellaea atropurpurea. A–E. (Johnston & Muller 476, US). A. Habit. B. Pinna base. C. Rachis abaxial surface. D. Rachis, adaxial surface. E. Rhizome scale. F, G, H, J. (Windham 549, UT). F. Habit. G. Rachis, abaxial surface. H. Rachis, adaxial surface. J. Rhizome scale. K, L. Pellaea mucronata (Taylor & Taylor 15606, NY and Palmer 100, NY). K. Pinna. L. Rhizome scale. M, N. Pellaea truncata (Reeves 5716, ASU). M. Pinnae. N. Rhizome scale.

PELLAEA

921

Fig. 219. A–C. Pellaea ribae (Heil 5347, BRY). A. Habit. B. Rachis and pinna. C. Rhizome scale. D–F. Pellaea ternifolia subsp. ternifolia (Mickel 3849, NY). D. Habit. E. Rachis and pinna bases. F. Rhizome scale. G, H. Pellaea ternifolia subsp. arizonica (Knobloch 5042, MSC). G. Pinnae. H. Rachis and pinna bases. J, K. Pellaea villosa (Rzedowski 22894, NY). J. Habit. K. Rachis and pinna bases. L–N. Pellaea ternifolia subsp. brandegeei (Carter 2333, MEXU). L. Pinnae. M. Pinna base. N. Rhizome scale. O–Q. Pellaea wrightiana (Parry et al. s.n., NY). O. Pinnae. P. Segments. Q. Rhizome scale.

922

PENTAGRAMMA

Fig. 220. A–C. Pentagramma triangularis subsp. triangularis (Howe s.n., Apr 1898, NY, from California). A. Habit. B. Segment, abaxial surface. C. Segment, adaxial surface. D, E. Pentagramma triangularis subsp. maxonii (Montgomery & Huttleston 83-31, NY). D. Frond. E. Segment, adaxial surface. F, G. Pentagramma triangularis subsp. viscosa (Montgomery & Huttleston 83-41, NY). F. Frond. G. Segment, adaxial surface.

PHANEROPHLEBIA

923

Fig. 221. A–C. Phanerophlebia nobilis (Anderson & Laskowski 4363, MICH). A. Habit. B. Pinna detail. C. Sorus. D–F. Phanerophlebia auriculata (Pringle 831, NY). D. Pinna. E. Pinna detail, abaxial surface. F. Pinna. G, H, J, K. Phanerophlebia umbonata (Pringle 1982, NY). G. Pinna. H. Pinna detail, abaxial surface. J. Sorus. K. Indusium, section.

924

PHANEROPHLEBIA

Fig. 222. A–E. Phanerophlebia gastonyi (Mickel 6044, NY). A. Blade apex. B. Pinna detail. C, D. Pinna margin. E. Pinna. F–H, J–L. Phanerophlebia pumila. F, G. (Mickel 5384, NY). F. Habit. G. Pinna detail. H, J. (Mickel 5377, NY). H. Habit. J. Pinna detail. K, L. (Cultivated, NYBG 158/72). K. Pinna. L. Pinna detail. M–P. Phanerophlebia macrosora (Mickel 4156, NY). M. Pinna. N. Rhizome and stipe bases. O. Pinna detail. P. Stipe scales. Q, R. Phanerophlebia gastonyi (Mickel 3786b, NY). Q. Pinna. R. Pinna detail. S, T. Phanerophlebia remotispora (Mickel 7248, NY). S. Pinna. T. Pinna detail.

PHLEBODIUM

925

Fig. 223. A–D. Phlebodium areolatum A. Habit (King & Soderstrom 4757, MICH). B, C. (Mickel 4850, NY). B. Rhizome scale. C. Pinna detail, abaxial surface. D. (Mickel 1132, NY), pinna detail, abaxial surface. E, F. Phlebodium decumanum (Rovirosa 252, NY). E. Pinna. F. Pinna detail, abaxial surface.

926

PILULARIA

Fig. 224. A–C. Pilularia americana (Ahart s.n., 15 May 1978, NY, from California). A. Habit. B. Rhizome with sporocarps. C. Crozier.

PITYROGRAMMA

Fig. 225. A, B. Pityrogramma calomelanos. A. (Hinton 16295, MICH, frond; McVaugh 11936, MICH, base), habit. B. (McVaugh 11936, MICH), segment.

927

928

PITYROGRAMMA

Fig. 226. A, B. Pityrogramma dealbata (Pringle 6078, NY). A. Pinna, B. Segment detail. C–E. Pityrogramma ebenea. C, D. (Ventura A. 230, NY). C. Pinna. D. Segment. E. (Breedlove 23327, NY), segment (sori along raised veins). F–H, J, K. Pityrogramma trifoliata (Mickel 6082, NY). F, G. Vegetative pinnae. H. Sterile pinna detail. J. Fertile pinna. K. Fertile segment detail.

PLAGIOGYRIA

929

Fig. 227. A–E. Plagiogyria pectinata (Hellwig 247, NY), frond, rhizome (Lems s.n., 9 Jan 1964, NY, from Costa Rica), habit. B. Sterile pinna detail. C. Fertile frond. D. Fertile pinna detail, abaxial surface. E. (Mickel 3815, NY), base of sterile blade.

930

PLEOPELTIS

Fig. 228. A. Pleopeltis crassinervata (Hallberg 1389, NY), habit. B–F. Pleopeltis astrolepis (Mickel 5165, NY). B. Habit. C. Blade detail. D, E. Blade scales. F. Rhizome scale. G, H, J, K. Pleopeltis conzattii (Mickel 1390, NY). G. Habit. H. Blade detail. J. Blade scale. K. Rhizome scale. L. Pleopeltis × melanoneuron (Purpus 5798, US), habit. M. Polypodium × bartlettii (Bartlett 10286, US), habit. N. Pleopeltis × sordidula (Copeland herb. 149, US), habit. O–Q. Pleopeltis fallax (Mickel 6468, NY). O. Habit. P. Fertile blade detail. Q. Blade scale. R–V. Pleopeltis mexicana (Mickel 4357, NY). R. Habit. S. Blade scale. T–V. Rhizome scales.

PLEOPELTIS

931

Fig. 229. A–D. Pleopeltis polylepis var. interjecta (Mickel 1162, NY). A. Habit. B. Rhizome scale. C. Blade detail, abaxial surface. D. Blade scales. E–H. Pleopeltis polylepis var. erythrolepis (Pringle 825, NY). E. Habit. F. Rhizome scale. G. Blade detail, abaxial surface. H. Blade scale. J–M. Pleopeltis polylepis var. polylepis (Mickel 7065, NY). J. Habit. K. Rhizome scale. L. Blade detail, abaxial surface. M. Blade scales. N. Pleopeltis angusta var. angusta (Mickel 6342, NY), habit. O–R. Pleopeltis angusta var. stenoloma (Mickel 1470, NY). O. Habit. P. Rhizome scale. Q. Blade detail. R. Blade scale.

932

POLYBOTRYA

Fig. 230. A–C. Polybotrya polybotryoides (Hernández G. 964, NY). A. Apex of sterile blade. B. Venation. C. Fertile pinna. D–H. Polybotrya caudata. D. (Breedlove 34101, DS), base of proximal sterile pinna. E. Stipe scale. F. Scale detail. G. Segment base. H. (Jones & Facey 3246, NY, from Guatemala), base of proximal fertile pinna.

POLYPODIUM

933

Fig. 231. A–D. Polypodium bradeorum (Mickel 7299, NY). A. Habit. B. Blade detail. C. Blade scale. D. Rhizome scale. E–H. Polypodium muenchii (Mickel 5524, NY). E. Habit. F. Pinna detail. G. Blade scale. H. Rhizome scale. J–L. Polypodium polypodioides var. aciculare (Mickel 1019, NY). J. Habit. K. Pinna. L. Blade scale. M–R. Polypodium polypodioides var. knoblochianum (Hartman 341, UC). M. Habit. N. Rhizome scale. O. Pinna, abaxial surface. P. Pinna, adaxial surface. Q. Abaxial blade scale. R. Adaxial blade scale. S, T. Polypodium polypodioides var. polypodioides (Martínez S. 15425, NY). S. Pinna. T. Blade scale. U, V. Polypodium polypodioides var. aciculare (Magaña s.n., 27 Apr 1968, NY). U. Pinna. V. Blade scales. W, X. Polypodium polypodioides var. michauxianum (Palmer 295, NY). W. Pinna. X. Blade scales.

934

POLYPODIUM

Fig. 232. A–D. Polypodium bradeorum (Breedlove 22284, NY). A. Habit. B. Rhizome scale. C. Pinna detail. D. Blade scale. E, F. Polypodium dulce. E. (Mickel 3152, NY, from Costa Rica), habit. F. (Purpus 6871, NY), pinna detail.

POLYPODIUM

935

Fig. 233. A–D. Polypodium lepidotrichum (Ventura 2357, NY). A. Rhizome, stipe, and pinnae. B. Pinna base. C. Stipe scale. D. Rhizome scale. E–H. Polypodium pyrrholepis (Hallberg 1312, NY). E. Rhizome and pinnae. F. Pinna base. G. Stipe scale. H. Rhizome scale. J–S. Polypodium sanctae-rosae. J–M. (Mickel 1639, NY). J. Rhizome, stipe, and pinnae. K. Pinna base. L. Stipe scale. M. Rhizome scale. N–Q. (Mickel 4470, NY). N. Habit. O. Pinna base. P. Stipe scale. Q. Rhizome scale. R, S. (Mickel 4471, NY). R. Rhizome. S. Rhizome scale.

936

POLYPODIUM

Fig. 234. A–D. Polypodium furfuraceum (Mickel 3883, NY) (see also Figs. 241K–M). A. Habit. B. Pinna. C. Blade scale. D. Rhizome scale. E–H, J. Polypodium platylepis (Mickel 4540, NY). E. Habit. F. Sterile and fertile pinna detail. G. Adaxial pinna detail. H. Blade scale. J. Rhizome scale. K–N. Polypodium thyssanolepis (Mickel 4964, NY) (see also Figs. 242D, E). K. Habit. L. Pinna base. M. Blade scale. N. Rhizome scale. O–R. Polypodium cryptocarpon (Hellwig et al. 1297, NY). O. Habit. P. Pinnae. Q. Blade scale. R. Rhizome scale.

PLEOPODIUM

937

Fig. 235. A–C. ×Pleopodium tricholepis (Mickel 3873, NY; see comments under Pleopeltis mexicana). A. Habit. B. Pinna base. C. Blade scale. D–F. Polypodium collinsii (Breedlove 24924, NY). D. Pinna base. E. Rachis detail. F. Rachis scale. G. Polypodium subpetiolatum (Ugent & Flores 2767, NY), pinna detail (see also Figs. 237F, G). H, I. Polypodium hispidulum (Hallberg 1693, NY), pinna base. I. sporangial capsule, with acicular hair. J, K. Polypodium puberulum (Mickel 4982, NY) (see also Figs. 237H, J). J. Fertile pinnae. K. Blade detail. L, M. Polypodium diplotrichum (Mickel 6181, NY). L. Habit. M. Pinna base. N, O. Polypodium echinolepis (Hallberg 1362, NY). N. Pinna detail. O. Rhizome scale.

938

POLYPODIUM

Fig. 236. A–E. Polypodium madrense (Mickel 793, NY). A. Habit. B. Pinna. C. Rhizome. D. Blade scale. E. Rhizome scale. F–H. Polypodium plebeium (Mickel 1014, NY). F. Pinnae. G. Pinna base. H. Blade scale. J–N. Polypodium alansmithii (Mickel 4573, NY). J. Habit. K. Pinna detail. L. Rhizome. M. Blade scale. N. Rhizome scale. O–S. Polypodium rzedowskianum (Rzedowski 18605, NY). O. Habit. P. Pinna detail. Q. Rhizome. R. Blade scale. S. Rhizome scale.

POLYPODIUM

939

Fig. 237. A–C. Polypodium martensii (Mickel 1163, NY). A. Habit. B. Pinna detail. C. Rhizome scale. D, E. Polypodium hartwegianum (Mickel 1153, NY). D. Pinnae. E. Pinna detail. F, G. Polypodium subpetiolatum (Mickel 5036, NY) (see also Fig. 235G). F. Pinnae. G. Pinna detail. H, J. Polypodium puberulum (Mickel 6599, NY) (see also Figs. 235J, K). H. Pinnae. J. Pinna detail. K, L. Polypodium fraternum (Mickel 4747, NY). K. Pinnae. L. Pinna detail. M, N. Polypodium longepinnulatum (Mickel 6688, NY) (see also Figs. 244M, N). M. Pinnae. N. Pinna detail. O, P. Polypodium fraternum (Mickel 1112, NY). O. Pinnae. P. Pinna detail.

940

POLYPODIUM

Fig. 238. A–C. Polypodium rhodopleuron (Hallberg 1365, NY). A. Rhizome and pinnae. B. Pinna detail. C. Rhizome scale. D–F. Polypodium falcaria (Mickel 1527, NY). D. Pinnae. E. Pinna detail. F. Rhizome scale. G, H, J. Polypodium plesiosorum (Mickel 4473, NY) (see also Figs. 245F–H, J–L). G. Rhizome and pinnae. H. Pinna detail. J. Rhizome scale. K–M. Polypodium californicum (Mickel 7426, NY). K. Habit. L. Pinna detail. M. Rhizome scale. N–P. Polypodium arcanum var. arcanum (Mickel 4167a, NY). N. Rhizome and pinnae. O. Pinna detail. P. Rhizome scale. Q–S. Polypodium colpodes (Mickel 5045, NY). Q. Rhizome and pinnae. R. Pinna detail. S. Rhizome scale.

POLYPODIUM

941

Fig. 239. A–C. Polypodium pleurosorum (Mickel 1099, NY). A. Rhizome and pinnae. B. Pinna detail. C. Rhizome scale. D–F. Polypodium eperopeutes (Mickel 1595, NY). D. Rhizome and pinnae. E. Pinna detail. F. Rhizome scale. G, H, J. Polypodium triseriale (Hallberg 1284, NY). G. Rhizome and pinnae. H. Pinna detail. J. Rhizome scale. K–M. Polypodium dissimile (Hallberg 1281, NY). K. Rhizome and pinnae. L. Pinna detail. M. Rhizome scale. N–Q. Polypodium rhachipterygium. N, O, Q. (Hallberg 1605, NY). N. Rhizome and pinnae. O. Pinna detail. P. (Hallberg 1285, NY), pinna detail. Q. Rhizome scale.

942

POLYPODIUM

Fig. 240. A–D. Polypodium fraxinifolium (Türckheim 642, NY). A. Pinnae. B. Rhizome. C. Pinna detail. D. Rhizome scale. E–H. Polypodium microgrammoides (Carranza et al. 4320, UC). E. Habit. F. Blade detail, abaxial surface. G. Blade detail, adaxial surface margin. H. Rhizome scale. J–L. Polypodium praetermissum (Reeder & Reeder 2473, US). J. Habit. K. Pinna detail. L. Rhizome scale.

POLYPODIUM

943

Fig. 241. A, B. Polypodium bradeorum. A. (Breedlove 22284, F, from Chiapas), sori. B. (Mickel 3484, NY, from Costa Rica), sori. C, D. Polypodium lindenianum (Méndoz Ton 5335, NY). C. Pinnae. D. Segment. E–H, J. Polypodium villagranii (Rzedowski 29979, ENCB). E. Habit. F. Pinna, abaxial surface. G. Pinna, adaxial surface. H. Stipe scales. J. Rhizome scale. K–M. Polypodium furfuraceum (Rzedowski 22652, NY) (see also Figs. 234A–D). K. Fertile pinna detail, abaxial surface. L. Sterile pinna detail, abaxial surface. M. Pinna detail, adaxial surface.

944

POLYPODIUM

Fig. 242. A–C. Polypodium riograndense (Fishbein 1784, ARIZ). A. Habit. B. Pinna. C. Blade scale. D, E. Polypodium thyssanolepis (Reveal 2730, ARIZ) (see also Figs. 234K–N). D. Habit of small plant. E. Stipe detail. F–H, J. Polypodium fallacissimum (Palmer 426, NY). F. Habit. G. Pinna. H. Stipe detail. J. Blade scale. K–M. Polypodium rosei (Pringle 4535, NY). K. Pinnae and rhizome. L. Pinna, abaxial surface. M. Pinna detail, adaxial surface. N, O. Polypodium remotum (Brade 75, NY, from Costa Rica). N. Habit. O. Rhizome scale.

POLYPODIUM

945

Fig. 243. A, B. Polypodium hesperium (Correll & Gentry 23218, US). A. Habit. B. Pinna. C–G. Polypodium fissidens (Mickel 1241, NY). C. Pinnae and rhizome. D. Blade base. E. Rachis detail. F. Sporangium. G. Rhizome scale. H, J–M. Polypodium christensenii. H, J, K. (Breedlove 34803, DS). H. Fertile pinna. J. Pinna base, abaxial surface. K. Pinna base, adaxial surface. L. (Breedlove 41087, DS), rhizome scale. M. (Breedlove 32006, NY), sterile pinna bases. N, O. Polypodium alavae (Alava 1287, MICH). N. Pinnae. O. Sporangia.

946

POLYPODIUM

Fig. 244. A–D. Polypodium guttatum (Rzedowski 24040, NY). A. Habit. B. Pinna bases. C. Rhizome scale. D. Blade scale. E, F. Polypodium chiapense (Breedlove 27453, NY). E. Pinnae. F. Pinna base. G, H, J. Polypodium liebmannii (Ventura A. 201, ENCB). G. Rhizome and pinnae. H. Pinna base. J. Rhizome scale. K, L. Polypodium conterminans (Breedlove 28918, NY). K. Pinnae. L. Pinna detail. M, N. Polypodium longepinnulatum (see also Figs. 237M, N). M. (Gittins 4184, NY), pinna detail. N. (Vera Santos 3670, NY), pinna detail.

POLYPODIUM

947

Fig. 245. A, B. Polypodium scouleri (Moran 6422, MEXU). A. Habit. B. Rhizome scale. C–E. Polypodium eatonii (Ventura A. 16945, NY). C, D. Blade base. E. Pinna detail. F–H, J–L. Polypodium plesiosorum (Mickel 4473, NY) (see also Figs. 238G, H, J). F. Rhizome G. Rhizome scale. H. Pinna detail. J–L. (Diaz-Barriga 69, IEB). J. Pinnae. K. Pinna base detail, abaxial surface. L. Sporangium. M–P. Polypodium arcanum var. septentrionale (Pringle 13294, CAS). M. Rhizome and pinna. N. Pinna base detail, abaxial surface. O. Contorted scale. P. Rhizome scale. Q, R. Polypodium arcanum var. bakeri (Pringle 3352, NY). Q. Rhizome and pinnae. R. Rhizome scale.

948

POLYSTICHUM

Fig. 246. A–D. Polystichum rachichlaena (McVaugh 11612, MICH). A. Frond. B. Pinna. C. Pinnule. D. Stipe scale. E. Polystichum schizophyllum (Lorea 4574, UC), pinna base.

POLYSTICHUM

949

Fig. 247. A. Polystichum acrostichoides (Dorr 2596, UC, NY), habit. B. Polystichum munitum (Anthony 9, NY and US ), pinnae. C–E. Polystichum muenchii (Münch 14, DS). C. Habit. D. Pinna. E. Sorus. F–H. Polystichum speciosissimum (Mickel 1200, NY). F. Stipe base. G. Pinna. H. Pinnule and rachis detail. J–L. Polystichum turrialbae. J. (Mickel 7056, NY) Pinnae. K, L. (Schaffner 86, NY). K. Pinnule. L. Stipe base scale.

950

POLYSTICHUM

Fig. 248. A, B. Polystichum muricatum (Breedlove 28923, NY). A. Pinnae. B. Stipe base. C–F. Polystichum furfuraceum (Münch 113, DS). C. Pinna. D. Rachis. E, F. Rachis scales. G, H, J. Polystichum alfaroi (Purpus 7231, UC). G. Pinna. H. Pinnule. J. Sorus. K–N. Polystichum orbiculatum (Breedlove 24306, DS). K. Pinnae. L. Stipe base. M. Pinna, adaxial surface. N. Pinna apex, abaxial surface.

POLYSTICHUM

951

Fig. 249. A–C. Polystichum hartwegii (Mickel 4011, NY) (see also Figs. 250P–S). A. Pinna. B. Pinnule, abaxial surface. C. Pinnule margin, adaxial surface. D–G. Polystichum erythrosorum (Mickel 4915, NY). D. Pinna. E. Stipe base. F. Pinnule. G. Stipe scale. H, J–L. Polystichum mickelii (Mickel 4836, NY) (see also Figs. 250L–O). H. Pinna. J. Stipe. K. Pinnule. L. Stipe scale. M–P. Polystichum fournieri (Mickel 4170, NY). M. Pinnae. N. Stipe base. O. Segment. P. Stipe scale. Q–T. Polystichum ordinatum (Mickel 4122a, NY). Q. Pinna. R. Stipe. S. Pinnule. T. Stipe scale.

952

POLYSTICHUM

Fig. 250. A–F. Polystichum platyphyllum (Mickel 4004, NY). A. Pinna. B. Blade apex. C. Stipe base. D. Rachis bud. E. Pinnule. F. Stipe scale. G, H, J, K. Polystichum distans (Mickel 3932, NY). G. Pinnae. H. Stipe base. J. Segment. K. Stipe scale. L–O. Polystichum mickelii (Mickel 7189, NY) (see also Figs. 249H, J–L). L. Pinna. M. Stipe base. N. Pinnule. O. Stipe scale. P–S. Polystichum hartwegii (Mickel 4307, NY) (see also Figs. 249A–C). P. Pinna. Q. Stipe base. R. Pinnule. S. Stipe scale.

POLYTAENIUM

953

Fig. 251. A–C. Polytaenium feei (Mickel 6465, NY). A. Habit. B. Cross-section of blade. C. Rhizome scale. D–F. Polytaenium lineatum (Mickel 6190, NY). D. Habit. E. Cross-section of blade. F. Rhizome scale. G, H, J. Polytaenium chlorosporum (Hallberg 1546, NY). G. Habit. H. Cross-section of blade. J. Rhizome scale. K–M. Polytaenium cajenense (Mickel 6467, NY). K. Habit. L. Cross-section of blade. M. Rhizome scale.

954

PSILOTUM

Fig. 252. A–C. Psilotum nudum (Orcutt 6317, DS). A. Habit (rhizome from Palmer 688, in 1886, NY). B. Stem cross-sections. C. Fertile branch. D, E. Psilotum complanatum (Ventura 368, NY). D. Habit. E. Fertile branch.

PTERIDIUM

955

Fig. 253. A–C. Pteridium aquilinum var. feei (Mickel 1392, NY). A. Pinnules. B. Segments, abaxial surface. C. Hairs. D, E. Pteridium aquilinum var. pubescens (Gallegos s.n., in 1923, US). D. Segments, adaxial surface. E. Segments, abaxial surface. F. Pteridium aquilinum var. latiusculum (Hernández 7629, MEXU), segments, abaxial surface. G, H, J. Pteridium arachnoideum G. (Mickel 4241, NY), pinnules. H, J. (Rzedowski & McVaugh 69, NY). H. Rachilla and segment base. J. Segment base, abaxial surface. K–M. Pteridium caudatum. K, L. (Mickel 4542, NY). K. Base of pinna. L. Segment bases, abaxial surface. M. (Mickel 4176, NY), segment bases, abaxial surface.

956

PTERIS

Fig. 254. A. Pteris cretica (Mickel 3722, NY), frond. B. Pteris longifolia (Mickel 4012, NY), pinnae (see also Figs. 259D, E). C, D. Pteris muricata (Mickel 4546, NY). C. Pinna. D. Stipe. E, F. Pteris grandifolia (Mickel 1482, NY). E. Pinna. F. Pinna detail. G, H. Pteris pulchra (Hallberg 1630, NY). G. Pinna. H. Pinna detail. J, K. Pteris biaurita (Mickel 5166, NY). J. Pinna. K. Segment. L, M. Pteris quadriaurita (Mickel 6990, NY). L. Pinna. M. Segment.

PTERIS

957

Fig. 255. A, B. Pteris altissima (Mickel 1456, NY). A. Pinnule. B. Venation. C–E. Pteris erosa (Mickel 6142, NY). C. Pinna. D. Pinnule. E. Indusium (Boutin & Kimnach 3112, NY). F, G. Pteris orizabae (Mickel 4664, NY). F. Pinna. G. Segments. H, J. Pteris podophylla (Mickel 1524, NY). H. Pinnule. J. Segments.

958

PTERIS

Fig. 256. A–E. Pteris tripartita (Zanoni et al. 22569, NY, from Dominican Republic). A. Base of blade. B. Pinnules, adaxial surface. C. Rachis detail, adaxial surface. D. Pinnules, adaxial surface. E. Blade detail

PTERIS

959

Fig. 257. A, B. Pteris paucinervata (Ghiesbreght 380, NY). A. Pinna. B. Segment C, D. Pteris muricella (Breedlove 25068, NY). C. Basal pinna. D. Segment. E–G. Pteris pungens. E. (Mell 2033, NY), juvenile frond. F, G. (Hernández G. 1901, NY). F. Basal pinna. G. Segment.

960

PTERIS

Fig. 258. A–D. Pteris tremula (Vigener s.n., NY, from cultivation). A. Pinna. B. Pinna base, adaxial surface. C. Fertile segment. D. Sterile segment. E, F. Pteris propinqua (Stern et al. 684, NY, from Panama). E. Blade apex and pinna. F. Segment.

PTERIS

961

Fig. 259. A–C. Pteris vittata (Hinton 24120, NY). A. Frond. B. Sterile pinna detail. C. Fertile pinna base. D, E. Pteris longifolia (Koch & Fryxell 83225, NY) (see also Figs. 254B). D. Sterile pinna detail. E. Fertile pinna base. F–H, J. Pteris chiapensis (Breedlove 22492, DS). F. Pinna. G. Adaxial rachilla groove. H. Adaxial rachis groove. J. Fertile pinna detail.

962

RADIOVITTARIA

Fig. 260. Radiovittaria stipitata (Mickel 3082, NY, from Costa Rica). A. Habit. B. Rhizome scale. C. Blade detail.

SACCOLOMA

963

Fig. 261. A, B. Saccoloma elegans (Hernández G. 1555, NY). A. Pinna. B. Fertile margin. C–E. Saccoloma inaequale (Mickel 974, ISC). C. Pinnule. D. Segment. E. Segment detail.

964

SALVINIA

Fig. 262. A–D. Salvinia minima (Breedlove 56065, NY). A. Fertile plant. B. Sterile plant. C. Blade hairs. D. Blade hair. E–H, J, K. Salvinia auriculata (Breedlove 34932, NY). E. Habit. F. Leaf. G. Blade hairs. H. Blade hair. J. Habit. K. Blade hair.

SCHAFFNERIA

Fig. 263. A, B. Schaffneria nigripes (Mickel 7333, NY). A. Habit. B. Blade detail.

965

966

SCHIZAEA

Fig. 264. A, B. Schizaea elegans (Hallberg 1330, NY). A. Blade. B. Blade base. C–E. Schizaea poeppigiana (Tillett 7847, NY). C. (rhizome from Huber et al. 8046, NY, from Venezuela), habit. D. Fertile pinnae. E. Pinna detail, adaxial and abaxial surfaces.

SCOLIOSORUS

Fig. 265. A–E. Scoliosorus ensiformis (Mickel 6186, NY). A. Habit. B. Cross-section of blade. C. Rhizome scale. D. Sporangia with paraphyses. E. Paraphyses.

967

968

SELAGINELLA

Fig. 266. A–C. Selaginella lepidophylla (Mickel 6663, NY). A. Habit. B. Upper surface of branch. C. Lower surface of branch. D–G. Selaginella basipilosa (Mickel 4965, NY). D. Habit. E. Branch. F. Vegetative leaf. G. Sporophyll. H, J–L. Selaginella wrightii (Mickel 5005, NY). H. Habit. J. Branch. K. Vegetative leaf. L. Sporophyll. M–P. Selaginella rupincola (Mickel 4561, NY). M. Habit. N. Branch. O. Vegetative leaf. P. Sporophyll.

SELAGINELLA

969

Fig. 267. A–F. Selaginella pallescens (Mickel 7006, NY). A. Branch habit. B. Upper surface of branch. C. Lower surface of branch. D. Median leaf. E. Axillary leaf. F. Lateral leaf. G, H. Selaginella harrisii (Hallberg 1375, NY) (see also Figs. 282A–F). G. Upper surface of branch. H. Lower surface of branch. J–O. Selaginella hoffmannii (Mickel 5914, NY) (see also Figs. 269N–S). J. Habit. K. Upper surface of branch. L. Lower surface of branch. M. Median leaf. N. Axillary leaf. O. Lateral leaf. P–T. Selaginella cuneata (Mickel 1044, NY). P. Upper surface of branch. Q. Lower surface of branch. R. Median leaf. S. Axillary leaf. T. Lateral leaf. U–Z. Selaginella oaxacana (Mickel 6350, NY). U. Branch habit. V. Upper surface of branch. W. Lower surface of branch. X. Median leaf. Y. Axillary leaf. Z. Lateral leaf.

970

SELAGINELLA

Fig. 268. A–F. Selaginella delicatissima (Mickel 882, NY). A. Habit. B. Lower surface of branch. C. Upper surface of branch. D. Median leaves. E. Axillary leaf. F. Lateral leaf. G, H, J–M. Selaginella flexuosa (Mickel 946, NY). G. Habit. H. Lower surface of branch. J. Upper surface of branch. K. Median leaves. L. Axillary leaf. M. Lateral leaf. N–S. Selaginella corrugis (Mickel 1043, NY). N. Habit. O. Lower surface of branch. P. Upper surface of branch. Q. Median leaves. R. Axillary leaf. S. Lateral leaf. T–Y. Selaginella mickelii. T. (Mexia 9190, NY), habit. U–Y. (Mickel 5871, NY). U. Lower surface of branch. V. Upper surface of branch. W. Median leaves. X. Axillary leaf. Y. Lateral leaf. AA–EE. Selaginella mosorongensis (Martínez-C. 331, NY). AA. Lower surface of branch. BB. Upper surface of branch. CC. Median leaves. DD. Axillary leaf. EE. Lateral leaf.

SELAGINELLA

971

Fig. 269. A–G. Selaginella hirtifolia (Mickel 4790, NY) (see also Figs. 271V–BB). A. Habit. B. Lower surface of branch. C. Upper surface of branch. D. Median leaves. E. Axillary leaf. F. Lateral leaf, lower surface. G. Lateral leaf, upper surface. H, J–M. Selaginella chiapensis (Hallberg 1625b, NY). H. Lower surface of branch. J. Upper surface of branch. K. Median leaves. L. Axillary leaf. M. Lateral leaf. N–S. Selaginella hoffmannii (Mickel 6938, NY) (see also Figs. 267J–O). N. Habit. O. Lower surface of branch. P. Upper surface of branch. Q. Median leaves. R. Axillary leaf. S. Lateral leaf. T–Y. Selaginella flagellata (Mickel 5133a, NY). T. Habit. U. Lower surface of branch. V. Upper surface of branch. W. Median leaf. X. Axillary leaf. Y. Lateral leaf. Z, AA–FF. Selaginella porphyrospora (Mickel 3984, NY). Z. Habit. AA. Lower surface of branch. BB. Upper surface of branch. CC. Median leaves. DD. Axillary leaf. EE. Lateral leaf, lower surface. FF. Lateral leaf, upper surface.

972

SELAGINELLA

Fig. 270. A–E. Selaginella prolifera (Mickel 6880, NY). A. Lower surface of branch. B. Upper surface of branch. C. Median leaf. D. Axillary leaf. E. Lateral leaf. F–H, J, K. Selaginella martensii (Mickel 6152, NY). F. Lower surface of branch. G. Upper surface of branch. H. Median leaf. J. Axillary leaf. K. Lateral leaf. L–P. Selaginella silvestris (Mickel 3793, NY). L. Lower surface of branch. M. Upper surface of branch. N. Median leaf. O. Axillary leaf. P. Lateral leaf. Q–U. Selaginella stellata (Hallberg 1587, NY). Q. Lower surface of branch. R. Upper surface of branch. S. Median leaf. T. Axillary leaf. U. Lateral leaf.

SELAGINELLA

973

Fig. 271. A–G. Selaginella popayanensis (Mickel 4295, NY). A. Habit. B. Upper surface of branch, with strobilus. C. Lower surface of branch, with strobilus. D. Median leaves. E. Axillary leaf. F. Lateral leaf, lower surface. G. Lateral leaf, upper surface. H, J–O. Selaginella mixteca (Mickel 6300, NY). H. Habit. J. Upper surface of branch. K. Lower surface of branch. L. Median leaves. M. Axillary leaf. N. Lateral leaf, lower surface. O. Lateral leaf, upper surface. P–U. Selaginella minima (Mickel 4204, NY). P. Habit. Q. Upper surface of branch, with strobili. R. Lower surface of branch, with strobili. S. Median leaves. T. Axillary leaf. U. Lateral leaf. V–Z, AA–BB. Selaginella hirtifolia (Mickel 4761, NY) (see also Figs. 269A–G). V. Habit. W. Upper surface of branch. X. Lower surface of branch. Y. Median leaves. Z. Axillary leaf. AA. Lateral leaf, lower surface. BB. Lateral leaf, upper surface. CC–HH. Selaginella schiedeana (Mickel 5820, NY). CC. Habit. DD. Upper surface of branch. EE. Lower surface of branch. FF. Median leaves. GG. Axillary leaf. HH. Lateral leaf.

974

SELAGINELLA

Fig. 272. A–G. Selaginella illecebrosa (Wendt 4749, NY). A. Habit. B. Lower surface of branch. C. Upper surface of branch. D. Lateral leaf. E. Median leaves. F. Axillary leaf. G. Lower stem leaf. H–N. Selaginella tarda (Mickel 6927, NY). H. Habit. I. Upper surface of branch. J. Lower surface of branch. K. Median leaf. L. Lateral leaf. M. Axillary leaf. N. Node. O–T. Selaginella bernoullii (Mickel 5092, NY). O. Habit. P. Upper surface of branch. Q. Lower surface of branch. R. Median leaves. S. Axillary leaf. T. Lateral leaf.

SELAGINELLA

975

Fig. 273. A–E. Selaginella carnerosana (Pringle s.n., 20 Sep 1890, NY). A. Habit. B. Upper surface of branch and strobilus. C. Leaf from upper surface of stem. D. Leaf from lower surface of stem. E. Sporophyll. F–H, J, K. Selaginella peruviana (Bourgeau 248, NY). F. Habit. G. Upper surface of branch and strobilus. H. Leaf from upper surface of stem. J. Leaf from lower surface of stem. K. Sporophyll. L–O. Selaginella cinerascens (Broome et al. 1951, NY). L. Habit. M. Upper surface of branch, including strobilus. N. Leaf. O. Sporophyll. P–S. Selaginella asprella (Moran 15451, UC). P. Habit. Q. Upper surface of branch and strobilus. R. Leaf. S. Sporophyll.

976

SELAGINELLA

Fig. 274. A–F. Selaginella landii (McVaugh 25825, NY). A. Habit. B. Upper surface of branch, including strobili. C. Lower surface of branch. D. Leaf from upper surface of stem. E. Leaf from lower surface of stem. F. Sporophyll. G, H, J–M. Selaginella parishii (Palmer 306, NY). G. Habit. H. Upper surface of branch, including strobilus. J. Lower surface of branch. K. Leaf from upper surface of stem. L. Leaf from lower surface of stem. M. Sporophyll. N–T. Selaginella eremophila (Thorne & Charlton 60172, MEXU). N. Habit. O. Branch and strobilus. P. Upper surface of branch. Q. Lower surface of branch. R. Leaf from upper surface of stem near apex. S. Leaf from lower surface of stem. T. Sporophyll. U–Z. Selaginella arizonica. U, W, Y, Z. (Mearns 2701, NY). U. Habit. V. (Dorn 2857, NY), upper surface of branch, including strobilus. W. Lower surface of branch. X. (Dorn 2857, NY), sporophyll. Y. Leaf from lower surface of branch. Z. Leaf from from upper surface of branch.

SELAGINELLA

977

Fig. 275. A–D. Selaginella bigelovii (Franklin 5798, NY). A. Habit. B. Branch including strobilus. C. Leaf. D. Sporophyll. E–H. Selaginella viridissima (Correll 13627, NY, from Texas). E. Habit. F. Branch including strobilus. G. Leaf. H. Sporophyll. J–M. Selaginella steyermarkii (Gittins 4257, NY). J. Habit. K. Upper surface of branch, including strobilus. L. Leaf. M. Sporophyll. N–P. Selaginella sartorii (Mickel 4498, NY). N. Branch with strobilus. O. Leaf. P. Sporophyll. Q, R. Selaginella sellowii (Mickel 6654b, NY). Q. Branch. R. Leaf.

978

SELAGINELLA

Fig. 276. A–E. Selaginella extensa (Fryxell et al. 3562, NY). A. Habit. B. Upper surface of branch, including strobilus. C. Leaf. D. Sporophyll, adaxial surface. E. Sporophyll, abaxial surface. F–H. Selaginella underwoodii (Correll & Gentry 21778, NY). F. Habit. G. Upper surface of branch. H. Leaf. J–L. Selaginella macrathera (Johnston 8067, UC). J. Habit. K. Leaf. L. Sporophyll. M–O. Selaginella arsenei (Basile 2, NY). M. Habit. N. Leaf. O. Sporophyll.

SELAGINELLA

979

Fig. 277. A–H, J–L. Selaginella sertata. A–F. (Mickel 5125, NY). A. Habit. B. Upper surface of branch. C. Lower surface of branch. D. Axillary leaf. E. Median leaf. F. Lateral leaf. G, H. (Mickel 5136, NY), leaves. G. Axillary leaf. H. Lateral leaf. J, K. (Mickel 5992, NY). J. Axillary leaf. K. Lateral leaf. L. (Iltis et al. 31021, NY), median leaf. M–O. Selaginella schizobasis (Mickel 5783, NY). M. Axillary leaf. N. Median leaf. O. Lateral leaf. P–S. Selaginella eurynota (Breedlove 28620, NY). P. Axillary leaf. Q. Median leaf. R, S. Lateral leaves. T–X. Selaginella marginata (Anderson 12566, NY). T. Upper surface of branch, including strobilus. U. Lower surface of branch. V. Axillary leaf. W. Median leaf. X. Lateral leaf.

980

SELAGINELLA

Fig. 278. A–E. Selaginella ribae (Thomas 2865, NY). A. Habit. B. Strobilus. C. Strobilus detail with some sporophylls removed. D. Median leaf. E. Lateral leaf. F–H. Selaginella mutica (Henrickson 12873, LL). F. Habit. G. Leaf. H. Sporophyll. J–R. Selaginella schaffneri (McVaugh 25810, NY). J. Habit. K. Upper surface of branch. L. Lower surface of branch. M, N. Median leaves. O, P. Axillary leaves. Q, R. Lateral leaves.

SELAGINELLA

981

Fig. 279. A–F. Selaginella convoluta (Valdez 50, NY). A. Habit. B. Upper surface of branch. C. Lower surface of branch. D. Median leaves. E. Axillary leaf. F. Lateral leaf. G, H, J–L. Selaginella novoleonensis (Pringle 2489, NY). G. Upper surface of branch, including strobilus. H. Lower surface of branch. J. Median leaves. K. Axillary leaf. L. Lateral leaf. M–Q. Selaginella gypsophila (Dorr et al. 2553, NY). M. Upper surface of branch, including strobilus. N. Lower surface of branch. O. Median leaf. P. Axillary leaf. Q. Lateral leaf.

982

SELAGINELLA

Fig. 280. A–F. Selaginella pilifera (Pringle 886, NY). A. Habit. B. Upper surface of branch, including strobilus. C. Lower surface of branch. D. Median leaves. E. Axillary leaf. F. Lateral leaf. G, H, J–M. Selaginella rzedowskii (Contreras 609, NY). G. Habit. H. Upper surface of branch, including strobilus. J. Lower surface of branch. K. Median leaves. L. Axillary leaf. M. Lateral leaves. N–T. Selaginella nothohybrida (Koch et al. 7989, NY). N. Habit. O. Upper surface of branch. P. Lower surface of branch. Q. Strobilus. R. Median leaves. S. Axillary leaf. T. Lateral leaf.

SELAGINELLA

983

Fig. 281. A–G. Selaginella pulcherrima (García & Torres 1524, NY). A. Habit. B. Main stem. C. Upper surface of branch. D. Lower surface of branch. E. Median leaves. F. Axillary leaf. G. Lateral leaf.

984

SELAGINELLA

Fig. 282. A–F. Selaginella harrisii (Montgomery 87-04, NY) (see also Figs. 267G, H). A. Habit B. Upper surface of branch, including strobilus. C. Lower surface of branch. D. Median leaves. E. Axillary leaf. F. Lateral leaf. G, H, J–N. Selaginella polyptera (Mickel 599, NY). G. Habit. H. Upper surface of main stem. J. Upper surface of branch. K. Lower surface of branch. L. Median leaf. M. Axillary leaf. N. Lateral leaf.

SELAGINELLA

985

Fig. 283. A–E. Selaginella huehuetenangensis (Breedlove 33161, NY). A. Upper surface of branch, including strobilus. B. Lower surface of branch, including strobilus. C. Median leaves. D. Axillary leaf. E. Lateral leaf. F–H, J–L. Selaginella guatemalensis (Breedlove & Dressler 29870, NY). F. Branch habit. G. Upper surface of branch, including strobili. H. Lower surface of branch, including strobili. J. Median leaves. K. Axillary leaf. L. Lateral leaf. M, N. Selaginella barnebyana (Méndez 5731, CAS). M. Median leaves on main stem. N. Lateral leaf. O–T. Selaginella lindenii (Linden s.n., P). O. Habit. P. Upper surface of branch, including strobili. Q. Lower surface of branch, including strobilus. R. Median leaves. S. Axillary leaf. T. Lateral leaf. U–X. Selaginella subrugosa (Mickel 6686, NY). U. Habit. V. Upper surface of branch, including strobilus. W. Lateral and median leaves. X. Lateral and axillary leaves.

986

SELAGINELLA

Fig. 284. A–G. Selaginella acutifolia (Mickel 6905, NY). A. Habit. B. Strobilus. C. Upper surface of branch, including strobilus. D. Lower surface of branch. E. Median leaves. F. Axillary leaf. G. Lateral leaf. H, J–N. Selaginella reflexa (Pringle 3292, NY). H. Branch habit. J. Upper surface of branch with strobilus. K. Lower surface of branch. L. Median leaves. M. Axillary leaf. N. Lateral leaf. O–T. Selaginella carioi (Breedlove 37259, NY). O. Habit. P. Upper surface of branch, including strobilus. Q. Lower surface of branch. R. Median leaves. S. Axillary leaf. T. Lateral leaf. U–Z. Selaginella arsiclada (Díaz Barriga 4627, NY). U. Habit. V. Upper surface of branch, including strobilus. W. Lower surface of branch. X. Median leaves. Y. Axillary leaf. Z. Lateral leaf.

SELAGINELLA

987

Fig. 285. A–F. Selaginella apoda. A. (Breedlove 21287, NY), habit. B, C. (Breedlove 22010, NY). B. Upper surface of branch, including strobilus. C. Lower surface of branch. D–F. (Breedlove 21287, NY). D. Median leaves. E. Axillary leaf. F. Lateral leaf. G, H, J–M. Selaginella breedlovei (Breedlove 28528, DS). G. Habit. H. Upper surface of branch, including strobilus. J. Lower surface of branch. K. Median leaves. L. Axillary leaf. M. Lateral leaf. N–S. Selaginella orizabensis (Müller 268, NY). N. Habit. O. Upper surface of branch, including strobilus. P. Lower surface of branch. Q. Median leaves. R. Axillary leaf. S. Lateral leaf. T–Y. Selaginella stenophylla (Hellwig 1222, NY). T. Habit. U. Upper surface of branch, including strobilus. V. Lower surface of branch. W. Median leaves. X. Axillary leaf. Y. Lateral leaf.

988

SELAGINELLA

Fig. 286. A–F. Selaginella simplex (Mickel 3996a, NY). A. Habit. B. Upper surface of branch, including strobili. C. Lower surface of branch, including strobili. D. Median leaves. E. Axillary leaf. F. Lateral leaf. G, H, J–M. Selaginella lineolata (Mickel 4014, NY). G. Habit. H. Upper surface of branch, including strobilus. J. Lower surface of branch. K. Median leaves. L. Axillary leaf. M. Lateral leaf. N–S. Selaginella tuberosa (Mickel 6153, NY). N. Habit. O. Upper surface of branch, including strobili. P. Lower surface of branch. Q. Median leaves. R. Axillary leaf. S. Lateral leaf.

SELAGINELLA

989

Fig. 287. A–F. Selaginella finitima (Wendt 2543, NY). A. Habit. B. Upper surface of branch, including strobilus. C. Lower surface of branch. D. Median leaves. E. Axillary leaf. F. Lateral leaf. G, H, J–M. Selaginella moritziana (Breedlove 24378, DS). G. Habit. H. Upper surface of branch, including strobilus. J. Lower surface of branch. K. Median leaves. L. Axillary leaf. M. Lateral leaf. N–T. Selaginella tenella (Mexia 1276, MICH). N. Habit. O. Upper surface of branch, including strobilus. P. Lower surface of branch. Q. Median leaves. R. Axillary leaf. S. Lateral leaf. T. Lateral leaf with idioblasts.

990

SPHAEROPTERIS

Fig. 288. A–E. Sphaeropteris horrida (Mickel 1302, NY). A. Pinna base. B. Stipe. C. Stipe scale. D. Segment. E. Blade scale.

SPHENOMERIS

Fig. 289. A–D. Sphenomeris clavata (Breedlove 35362, NY). A. Habit. B. Sterile segment. C. Fertile segment. D. Rhizome hair.

991

992

STICHERUS

Fig. 290. A–E. Sticherus bifidus. A. (Mexia 1349, MICH), habit. B–E. (Mickel 1103, ISC). B. Ultimate fork. C. Bud scale. D. Segments. E. Segment detail.

STICHERUS

993

Fig. 291. A–E. Sticherus palmatus (Mickel 1103, ISC). A. Portion of blade. B. Segments. C. Segment detail. D. Ultimate fork. E. Bud scale. F–H, J. Sticherus underwoodianus (Mickel 4243, NY). F. Ultimate fork. G. Blade scale. H. Segments. J. Segment detail. K–M. Sticherus brevipubis (Mickel 6535, NY). K. Blade fork. L. Ultimate fork. M. Bud scale.

994

STIGMATOPTERIS

Fig. 292. A, B. Stigmatopteris longicaudata (Mickel 975, ISC). A. Pinna and blade apex. B. Pinna detail. C, D. Stigmatopteris sordida (Mickel 6403, NY). C. Pinna. D. Segments.

TECTARIA

995

Fig. 293. A–C. Tectaria fimbriata (Maxon 4430, UC, from Cuba). A. Habit. B. Blade detail. C. Indusium. D, E. Tectaria transiens (Hallberg 1566, NY). D. Proximal pinna. E. Pinna detail. F, G. Tectaria trichodes (Zamudio 193.5, ENCB, frag. NY). F. Frond. G. Blade detail. H, J. Tectaria incisa f. incisa (Breedlove 22522, NY). H. Proximal pinna. J. Blade detail.

996

TECTARIA

Fig. 294. A. Tectaria panamensis (Allen 1320, NY, from Panama), habit. B–D. Tectaria heracleifolia (Mickel 6832, NY). B. Blade. C. Blade detail. D. Sorus. E–G. Tectaria mexicana (Mickel 6158, NY). E. Pinna. F. Blade detail. G. Sorus. H, J–L. Tectaria incisa f. vivipara. H, J, K. (Hallberg 1610, NY). H. Proximal pinna. J. Blade detail. K. Sorus. L. (Hernández G. 2587, NY), bulblet in pinna axil.

TERPSICHORE

997

Fig. 295. A–D. Terpsichore semihirsuta. A, B. (Hellwig 435, NY). A. Habit. B. Pinna, abaxial surface, with fungus. C, D. (Mickel 6762, NY). C. Frond. D. Pinnae. E. Terpsichore mollissima (Hallberg 1592, NY), pinnae. F, G. Terpisichore delicatula (Mickel 5292, NY). F. Habit. G. Pinnae.

998

TERPSICHORE

Fig. 296. A–C. Terpsichore subtilis (Breedlove 67840, CAS). A. Habit. B. Pinnae, abaxial surface, with fungus. C. Pinnae, adaxial surface with lime dots. D, E. Terpsichore senilis (Breedlove 68357, UC). D. Fronds. E. Pinnae. F, G. Terpsichore anfractuosa (Mickel 4292, NY). F. Habit. G. Blade detail.

TERPSICHORE

999

Fig. 297. A, B. Terpsichore cultrata (Mickel 6737, NY). A. Habit. B. Pinnae. C–H. Terpsichore spathulata. C–E. (Pringle 4145b, NY). C. Habit. D, E. Pinnae. F–H. (Bourgeau s.n., NY). F. Frond showing branching rachis. G, H. Blade detail. J, K. Terpsichore asplenifolia (Mickel 4293, NY). J. Habit. K. Pinnae.

1000

THELYPTERIS

Fig. 298. A–C. Thelypteris (Amauropelta) cheilanthoides var cheilanthoides. A. Habit (frond, McVaugh 13589, MICH; rhizome, McVaugh & Koelz 1365, MICH). B, C. (Mickel 4223, NY). B. Segments. C. Sorus. D, E. Thelypteris cheilanthoides var. mucosa (Mickel 4317, NY). D. Segment base. E. Sorus.

THELYPTERIS

1001

Fig. 299. A–C. Thelypteris (Amauropelta) mortonii (Sánchez 38, UC). A. Rhizome. B. Segment, abaxial surface. C. Segment, adaxial surface. D, E. Thelypteris tablana (Münch 146, DS). D. Segment, abaxial surface. E. Segment, adaxial surface. F, G. Thelypteris sancta (Breedlove 47427, CAS). F. Habit. G. Pinna. H, J. Thelypteris struthiopteroides (Hallberg 1759, NY). H. Segments. J. Segment detail. K, L. Thelypteris thomsonii (Breedlove 23121, NY). K. Pinna. L. Segment. M, N. Thelypteris cinerea (Breedlove 31678, NY). M. Pinna. N. Segment margin.

1002

THELYPTERIS

Fig. 300. A–C. Thelypteris (Amauropelta) linkiana (Mickel 1017, NY). A. Segments, abaxial surface. B. Distal pinna base with bud in axil. C. Segments, adaxial surface. D–F. Thelypteris concinna (Mickel 6304, NY). D. Segments, abaxial surface. E. Segment, adaxial surface. F. Sporangium. G, H. Thelypteris oligocarpa (Mickel 1122, NY). G. Segments, abaxial surface. H. Segment, adaxial surface. J, K. Thelypteris deflexa (Mickel 985, NY). J. Segments, abaxial surface. K. Segment, adaxial surface. L. Thelypteris pilosohispida (Mickel 5720, NY), segments. M. Thelypteris rudis (Mickel 4688, NY), segments.

THELYPTERIS

1003

Fig. 301. A, B. Thelypteris (Amauropelta) nubigena (Mickel 5533, NY). A. Segments. B. Rachis with hairs. C. Thelypteris balbisii (Mickel 5183, NY), pinna base. D–G. Thelypteris resinifera var. resinifera (Hallberg 1283, NY). D. Pinna base. E. Segments. F. Laminar hairs. G. Sorus. H. Thelypteris oaxacana (Mickel 6556, NY), segments. J. Thelypteris pilosula (Mickel 1187, NY), segments. K–M. Thelypteris melanochlaena (Mickel 5110, NY). K. Segments. L. Sorus. M. Laminar hairs. N–P. Thelypteris scalaris (Mickel 1054, NY). N. Segments. O. Sorus. P. Pinna axil with bud.

1004

THELYPTERIS

Fig. 302. A–C. Thelypteris (Amauropelta) venturae (Ventura 8141, ENCB). A. Pinna. B. Segment, abaxial surface. C. Segment, adaxial surface. D–F. Thelypteris atrovirens (Breedlove 32669, NY). D. Pinnae. E. Segment, abaxial surface. F. Segment, adaxial surface. G. Thelypteris resinifera var. proxima (Pringle s.n., 4 Nov 1889, UC), base of blade. H, J–L. Thelypteris caucaensis (Holm-Nielsen et al. 3352, UC, from Ecuador). H. Rhizome and stipe bases. J. Proximal pinnae. K. Segments, abaxial surface. L. Costular scale.

THELYPTERIS

1005

Fig. 303 A–E. Thelypteris (Amauropelta) loreae (Lorea 4540, UC). A. Middle and proximal pinnae. B. Stipe detail. C. Stipe scale. D. Adaxial blade surface. E. Abaxial blade surface.

1006

THELYPTERIS

Fig. 304. A, B. Thelypteris (Cyclosorus) kunthii (Mickel 4769, NY). A. Segments. B. Sorus. C–E. Thelypteris ovata var. lindheimeri (Mickel 1604, NY). C. Pinnae. D. Segments. E. Rachis, adaxial surface. F. Thelypteris puberula var. puberula (Mickel 852, NY), segments. G. Thelypteris kunthii (Mickel 3947, NY), segment, adaxial surface. H. Thelypteris patens (Hallberg 1677, NY), segments. J. Thelypteris albicaulis (Mickel 6156, NY), segments. K. Thelypteris dentata (Hallberg 1322, NY), segments. L. Thelypteris hispidula (Mickel 6087, NY), segments.

THELYPTERIS

1007

Fig. 305. A, B. Thelypteris (Cyclosorus) augescens (Cabrera 4536, CAS). A. Apical portion of blade. B. Segments. C–E. Thelypteris blepharis (Breedlove 33605, DS). C. Segments. D. Stipe base. E. Scale. F. Thelypteris tuerckheimii (Breedlove 23535, NY), base of proximal pinnae. G, H. Thelypteris cretacea (Vázquez Torres 370, NY). G. Segments. H. Abaxial blade detail. J. Thelypteris interrupta (Cowan 2001, NY), segments. K, L. Thelypteris puberula var. sonorensis (Schaffner s.n., Oct 1879 & 81, NY). K. Segment, abaxial surface. L. Segment, adaxial surface. M–O. Thelypteris lanosa (Ventura A. 2717, NY). M. Base of proximal pinna. N. Segments. O. Sorus.

1008

THELYPTERIS

Fig. 306. A–C. Thelypteris (Goniopteris) blanda (Mickel 1424b, NY). A. Pinna. B. Rachis. C. Segments. D–F. Thelypteris tetragona (Mickel 5802, NY). D. Pinna. E. Segments. F. Sporangium. G, H, J. Thelypteris hatchii (Hallberg 1599, NY). G. Pinna. H. Segments. J. Laminar hairs. K. Thelypteris biolleyi (Mickel 1031, NY), laminar hairs. L–N. Thelypteris paucipinnata (Hallberg 1331, NY). L. Pinna. M. Segments. N. Sorus. O, P. Thelypteris toganetra (Mickel 5802, NY). O. Pinna. P. Segments. Q, R. Thelypteris imbricata (Mickel 4198, NY). Q. Pinna. R. Segments.

THELYPTERIS

1009

Fig. 307. A–C. Thelypteris (Goniopteris) guadalupensis (Proctor 39907, UC, from Puerto Rico). A. Habit. B. Segment. C. Rachis. D, E. Thelypteris minor (Breedlove 41979, DS). D. Base of proximal pinnae. E. Segment. F, G. Thelypteris nicaraguensis (Breedlove 22378, DS). F. Base of proximal pinnae. G. Segment. H, J, K. Thelypteris obliterata (Schipp 83, NY, from Belize). H. Pinnae. J. Segments. K. Sporangium. L, M. Thelypteris muenchii (Münch s.n., DS). L. Fertile pinnae. M. Sterile pinnae.

1010

THELYPTERIS

Fig. 308. A, B. Thelypteris (Goniopteris) stolzeana (Breedlove 22355, DS). A. Blade. B. Segments. C, D. Thelypteris reptans (Breedlove 32948, NY). C. Fertile blade. D. Pinna base. E–G. Thelypteris resiliens (Martínez S. 22710, UC, from Guatemala). E. proximal pinnae. F. Distal portion of blade with rachis bud. G. Segment. H, J, K. Thelypteris praetermissa (Croat 24548, UC, from Belize). H. Pinna bases. J. Segment. K. Costa detail. L, M. Thelypteris martinezii (Martínez S. 10123, MEXU). L. Pinna bases. M. Segment.

THELYPTERIS

1011

Fig. 309. A–C. Thelypteris (Goniopteris) schaffneri (Purpus 5944, UC). A. Base of proximal pinna. B. Segments. C. indusia. D, E. Thelypteris ghiesbreghtii (Hernández G. 1967, NY). D. Pinna detail. E. Sorus. F, G. Thelypteris hondurensis (Ordóñez 28, UC, from Honduras). F. Pinna detail. G. Sorus. H, J. Thelypteris rhachiflexuosa (Riba 1683, UC). H. Blade. J. Pinna detail. K, L. Thelypteris poiteana (Mickel 5799, NY). K. Pinna detail. L. Sporangium. M. Thelypteris meniscioides (Hallberg 1634, NY), pinna detail.

1012

THELYPTERIS

Fig. 310. A–D. Thelypteris (Goniopteris) leptocladia (Goll 1016, NY, from Puerto Rico). A. Habit. B. Segment. C. Sporangium. D. Blade hairs.

THELYPTERIS

1013

Fig. 311. A, B. Thelypteris (Meniscium) falcata (Grayum 2757, NY, from Costa Rica). A. Pinna. B. Pinna detail. C, D. Thelypteris angustifolia (Grayum et al. 4911, UC, from Costa Rica). C. Base of fertile proximal pinna. D. Base of sterile proximal pinna. E–G. Thelypteris reticulata (Mexia 9202, NY). E. Base of fertile pinna. F. Fertile pinna detail. G. Sterile pinna detail. H, J. Thelypteris serrata (Hallberg 1762, NY). H. Pinna apex. J. Pinna detail. K–N. Thelypteris standleyi (Standley 24126, US). K. Bases of proximal fertile pinnae. L. Fertile pinna detail. M. Bases of proximal sterile pinnae. N. Sterile pinna detail.

1014

THELYPTERIS

Fig. 312. A–D. Thelypteris (Stegnogramma) pilosa (see also Fig. 313A). A–C. Mickel 7076, NY. A. Habit. B. Segments. C. Sporangium. D. (Correll & Gentry 23091, UC), habit.

THELYPTERIS

Fig. 313. A. Thelypteris (Stegnogramma) pilosa (Pringle 2589, NY), habit (see also Figs. 312A–D).

1015

1016

THELYPTERIS

Fig. 314. A–C. Thelypteris (Steiropteris) glandulosa var. brachyodus (Breedlove 34189, DS). A. Apical portion of blade. B. Base of proximal pinna. C. Segments.

THELYPTERIS

1017

Fig. 315. A–C. Thelypteris (Thelypteris) palustris var. pubescens (Schaffner s.n., in 1876, NY). A. Habit. B. Sterile segment. C. Fertile segment (indusia hidden among sporangia).

1018

TRICHOMANES

Fig. 316. A–C. Trichomanes radicans (see also Figs. 319K–M). A. (Pringle 5535, NY), habit. B, C. (Mickel 5643, NY). B. Segment. C. Sorus and involucre. D–F. Trichomanes collariatum (Ventura A. 8117, NY). D. Pinna. E. Segment. F. Sorus.

TRICHOMANES

1019

Fig. 317. A, B. Trichomanes diversifrons (Wendt et al. 3845A, NY). A. Fertile and sterile fronds. B. Fertile frond detail. C, D. Trichomanes ovale (Mickel 6870, NY). C. Habit. D. Frond. E, F. Trichomanes galeottii, juvenile (Breedlove 21893, NY). E. Habit, winged stipe. F. Fertile pinna. G, H, J. Trichomanes membranaceum (Wendt et al. 3844A, NY). G. Habit. H. Sori. J. Marginal scales. K–M. Trichomanes angustatum (Breedlove 68356, CAS). K. Habit. L. Pinnae. M. Sorus. N, O. Trichomanes galeottii (Mexia 9282, NY). N. Habit. O. Pinna detail.

1020

TRICHOMANES

Fig. 318. A, B. Trichomanes angustifrons (Breedlove 32179, DS). A. Habit. B. Bladedetail. C, D. Trichomanes petersii (Leonard 2121, NY, from South Carolina). C. Habit. D. Blade detail. E, F. Trichomanes curtii (Proctor et al. 17172, NY, from Nicaragua). E. Habit. F. Sori. G, H. Trichomanes ekmanii (Hernández X. 631, LL). G. Habit. H. Blade margin detail.

TRICHOMANES

1021

Fig. 319. A, B. Trichomanes reptans (Breedlove 32612, MICH). A. Habit. B. Blade detail. C, D. Trichomanes holopterum (Harris 9284, NY, from Jamaica). C. Habit. D. Blade detail. E–H, J. Trichomanes tuerckheimii. E, F. (Burger & Gentry 8943, NY, from Costa Rica). E. Pinna. F. Blade base detail. G–J. (Burch 4429, NY, from Costa Rica). G. Segments. H. Fertile blade detail. J. Sorus. K–M. Trichomanes radicans (Correll & Gentry 23088, NY) (see also Figs. 316A–C). K. Habit. L. Blade detail. M. Sorus.

1022

TRICHOMANES

Fig. 320. A, B. Trichomanes pinnatum (Mexia 9133, NY). A. Habit. B. Venation. C, D. Trichomanes polypodioides (Hallberg 1282, NY). C. Habit. D. Segments. E, F. Trichomanes lucens (Maxon 5635, Panama, NY). E. Habit. F. Pinna. G, H. Trichomanes crispum (Hallberg 1588, NY). G. Habit. H. Pinna.

TRICHOMANES

1023

Fig. 321. A–C. Trichomanes hymenophylloides (Mickel 5564, NY). A. Habit. B. Stipe. C. Pinna. D–F. Trichomanes pyxidiferum (Mickel 5761, NY). D. Habit. E. Stipe. F. Pinna. G, H. Trichomanes rigidum (Mickel 1035, NY). G. Habit. H. Pinnule. J, K. Trichomanes capillaceum (Mickel 6585, NY). J. Habit. K. Pinna.

1024

TRICHOMANES

Fig. 322. A, B. Trichomanes krausii (Mickel 5770, NY). A. Habit. B. Blade detail. C–E. Trichomanes godmanii (Mickel 5848, NY). C. Habit. D. Blade detail. E. Sorus. F–H. Trichomanes bucinatum (Mickel 5105, NY). F. Habit. G, H. Blade detail. J, K. Trichomanes hymenoides (Mickel 6120, NY). J. Habit. K. Frond.

VITTARIA

1025

Fig. 323. A–C. Vittaria bradeorum (Mickel 6670, NY). A. Habit. B. Blade detail. C. Rhizome scale. D–F. Vittaria flavicosta (Mickel 5663, NY). D. Habit. E. Blade detail. F. Rhizome scale. G, H, J, K. Vittaria graminifolia (Mickel 5267, NY). G. Habit. H. Blade detail. J. Spore. K. Rhizome scale. L–O. Vittaria lineata (Nee & Taylor 29183, NY). L. Habit. M. Blade detail. N. Spore. O. Rhizome scale.

1026

WOODSIA

Fig. 324. A–H, J, K. Woodsia mollis. A–F. (Mickel 766, NY). A. Habit. B. Rachis. C. Pinna, abaxial surface. D. Segment, adaxial surface. E. Segment, abaxial surface. F. Rhizome scale. G, H, J. (Mickel 6259, NY). G. Pinna. H. Segment. J. Sorus. K. (Pringle 11274, NY), segment. L. Woodsia canescens (Mickel 4489, NY), segment. M. Woodsia mollis (or possibly W. montevidensis) (Müller 6, NY), segment.

WOODSIA

1027

Fig. 325. A–E. Woodsia plummerae, (Wiggins 7401, MICH). A. Habit. B. Rachis. C. Rhizome scale. D. Adaxial surface margin. E. Indusia. F–H, J, K. Woodsia cystopteroides. F–H, J. (Breedlove & Thorne 18425, NY). F. Habit. G. Rachis. H. Pinna, abaxial surface. J. Adaxial surface margin. K. (Gentry 6432, ARIZ), indusia. L–O. Woodsia cochisensis (Baker 10399, ARIZ) (see also Figs. 326J, K). L. Habit. M. Rachis. N. Adaxial surface margin. O. Indusia.

1028

WOODSIA

Fig. 326. A–D. Woodsia phillipsii (Knobloch 2073, MSC). A. Habit. B. Rachis. C. Segment, adaxial surface. D. Indusium. E–H. Woodsia mexicana (Orcutt 1650, TEX). E. Habit. F. Rachis. G. Adaxial pinna margin. H. Indusium. J, K. Woodsia cochisensis (Pringle 834, F) (see also Figs. 325L–O). J. Rachis. K. Segment, adaxial surface. L, M. Woodsia neomexicana (Hinton 20081, TEX). L. Rachis. M. Segment, adaxial surface margin.

WOODWARDIA

1029

Fig. 327. A–E. Woodwardia spinulosa. A–C. (McVaugh 21555, MICH). A. Blade. B. Stipe base. C. Stipe scales. D, E. (Mickel 3794, NY). D. Segments. E. Segment detail.

1030

WOODWARDIA

Fig. 328. A, B. Woodwardia martinezii (Riba 446, NY). A. Pinnae. B. Pinna detail. C, D. Woodwardia fimbriata (Moran 25113, NY). C. Pinna. D. Pinna detail. E–G. Woodwardia semicordata (Nee 23542, NY). E. Pinna. F. Segment detail. G. Blade apex.

INDEX TO SCIENTIFIC NAMES Adiantum (Continued ) mathewsianum 42 mcvaughii 21, 32, 33, 34, 41, 711* mendoncae 30 mexicanum 38 microphyllum 203 modestum 27 monotis 37 oaxacanum 21, 25, 34, 35, 45, 713* obliquum 25, 33, 34, 35, 36, 712* patens 24, 31, 35, 39, 708* subsp. oatesii 35 pedatum 24 subsp. aleuticum 24 var. aleuticum 24 pellucidum 36 petiolatum 33, 35, 36, 710*, 712* philippense 32 phyllitidis 44 platyphyllum 39 poeppigianum 44 poiretii 21, 36, 37, 38, 712* polyphyllum 44 princeps 25 pulverulentum 25, 37, 43, 710* pulverulentum ⫻ villosum 710* raddianum 37, 38, 707* radiatum 20 schaffneri 27 seemannii 28, 38, 39, 714* shepherdii 21, 31, 35, 39, 713* strictum 367 tenerum 25, 39, 40, 709* var. dissectum 25 terminatum 40, 714* tetraphyllum 29, 37, 40, 41, 45, 710* var. majus 44 thalictroides 36, 37 trapeziforme 41, 42, 709* var. oblongatum 41 var. plumieri 41 trapezoides 39, 40 trichochlaenum 25, 40, 41, 42, 43, 45, 714* tricholepis 21, 37, 42, 43, 708* villosum 37, 42, 43, 713* wilesianum 43, 44, 710* wilsonii 44, 714* Aleuritopteris 71, 176, 193, 402 affinis 413 candida 416 chihuahuaensis 192, 193 cretacea 425 farinosa 192 lutea 184 mexicana 192, 193 schaffneri 192, 423 Allantodia costalis 268 Allosorus alabamensis 181 angustifolius 182 var. minor 182 atropurpureus 445

Allosorus alabamensis (Continued ) chaerophyllus 187 ciliata var. minor 200 decompositus 191 flexuosus 448 formosus 66 greggii 419 hirsutus 200 karwinskii 368 mucronatus 446 ochracea 184 pulchellus 66 sagittatus 449 Alsophila 45 australis 45 bicrenata 235 blechnoides 395, 396 firma 45, 47, 715* fulva 237 godmanii 237 marginalis 239 mexicana 237 microdonta 238, 239 muenchii 46 munchii 46 myosuroides 238 pilosa 632 pruinata 371 quadripinnata 372 rostrata 396 salvinii 45, 46, 47, 715* scabriuscula 235 schiedeana 239 tryoniana 46, 47, 715* Amauropelta 622, 623 breutelii 623 cheilanthoides 626 concinna 627 deflexa 628 mucosa 627 oligocarpa 632 pilosula 633 resinifera 633 rudis 635 sancta 635 sanctiformis 627 scalaris 636 thomsonii 637 Amphidesmium 395 blechnoides 395, 396 rostratum 396 Anabaena azollae 139 Ananthacorus 47, 525, 544, 549 angustifolius 47, 716* Anapeltis 396 lycopodioides 396 Anarthropteris 372 Anaxeton 410 Anaxetum 410 crassifolium 410

Anchistea 692, 693 virginica 692 Anemia 18, 48, 377 subg. Anemia 48 subg. Anemiorrhiza 48, 49, 50 subg. Coptophyllum 48 adiantifolia 49, 50, 51, 55, 717* var. asplenifolia 49 var. pumila 49 var. subaurita 49 adiantifolia ⫻ mexicana var. makrinii 49, 717* adiantifolia ⫻ mexicana var. mexicana 49, 717* affinis 50, 52, 54, 59, 721*, 723* anthriscifolia 60 asplenifolia 49 bipinnata 51 brandegeei 50, 53, 723* carvifolia 49 cicutaria 3, 49, 50, 719* clinata 53 colimensis 49, 50, 51, 724* collina 58 cornea 57 ⫻didicusana 58, 59 distans 60 familiaris 51, 52, 720* flexuosa var. anthriscifolia 60 fulva var. mexicana 60 guatemalensis 54, 60 helveola 57 hirsuta 50, 52, 54, 58, 59, 720* var. achillaeifolia 52 hirsuta ⫻ jaliscana 52 humilis 56 intermedia 50, 52, 53, 723* jaliscana 50, 52, 53, 56, 59 var. integrifolia 53, 54, 723* var. jaliscana 53, 54, 721* karwinskyana 52, 54, 60, 718* longistipes 57 makrinii 55 mexicana 49, 54, 58 var. makrinii 51, 55, 719* var. mexicana 55, 60, 719* var. paucifolia 60 muenchii 55, 56, 58, 717* multiplex 56, 59, 721* munchii 55 oblongifolia 56, 720* var. humilis 56 obovata 52 opaca 52 ⫻paraphyllitidis 56, 58, 722* pastinacaria 50, 52, 56, 57, 59, 720* phyllitidis 48, 52, 56, 58, 59, 722* pilosa 57 var. brevistipes 57 var. longistipes 57 recondita 54, 58, 59, 721

INDEX TO SCIENTIFIC NAMES Anemia (Continued ) rosei 54 seemannii 57 semihirsuta 58, 59, 722*, 723* speciosa 49, 55, 60, 719* tomentosa 52, 60 var. mexicana 60, 718* villosa var. karwinskyana 54 Anemidictyon phyllitidis 58 Anetia 60 Anetium 60, 544 citrifolium 60, 61, 725* Anisosorus 371 hirsutus 371 Anogramma 61, 463 chaerophylla 61, 62, 63, 726* guatemalensis 61, 62, 63 leptophylla 61, 62, 63, 726* novogaliciana 63, 726* Antigramma nigripes 548 Antrophyum 525, 526, 549 sect. Loxogramme 372 subg. Polytaenium 525 cajenense 526 carnosum 550 chlorosporum 526 ensiforme 549, 550 falcatum 550 feei 527 galeottii 550 guayanense 527 lacantunense 526 lanceolatum 527 lineatum 528 stenophyllum 527 subsessile 526 Arachniodes 63, 271, 455 aristata 63 aspidioides 63 denticulata 63, 727* Argyrochosma 3, 64, 132, 176, 411, 443 delicatula 65, 67, 68, 729* fendleri 65, 66, 69, 729* formosa 66, 728* incana 65, 66, 67, 68, 70, 729*, 730* jonesii 67, 69, 728* limitanea 66, 67, 69 subsp. limitanea 68, 730* subsp. mexicana 68, 70, 730* lumholtzii 67, 68, 69, 728* microphylla 66, 69, 728* nivea 64 pallens 69, 70, 71, 731* palmeri 70, 71, 731* peninsularis 70, 71, 731* pilifera 71, 731* Arthropteris 427 Aspidium 272, 609 agatolepis 273 albicaule 640, 647 ameristoneuron 230

Aspidium (Continued ) apertum 278 argutum 272 articulatum 427 athyrioides 273 augescens 640 biolleyi 650 biserratum 404 bourgaei 223 bourgeaui 223 cheilanthoides 626 cheiloplotium 638 chiapasense 222 chontalense 360 chrysocarpon 280 conspersoides 648 crinitum 281 ctenitis 219 denticulatum 63 donianum 281 equestre 222 expansum 227 exsudans 634 var. feei 634 var. majus 634 extensum 227 fimbriatum 610 flaccidum 273 fragile var. fumarioides 241 geropogon 640 ghiesbreghtii 634 glandulosum 19 goggilodus 643 gongylodes 638, 643 hartwegii 517 heracleifolium 610 hispidulum 643 jucundum 63 juglandifolium var. macrosorum 457 karstenii 228 karwinskyanum 275 kunzei 626 latifolium 613 leptorhachis 281 lindenii 230 lunense 387 macrophyllum 612 martinicense 612 melanostictum 227 mexicanum var. acutilobum 281 var. obtusilobum 278 microcarpon 228 microchlaena 228 molle 642 muenchii 518 munchii 518 munitum 519 nobile 455, 458 nodosum 427 obtusilobum 233 orbiculatum 520

Aspidium (Continued ) var. orbiculatum 520 ordinatum 521 paleaceum 281 pallidum var. majus 645 parallelogrammum 281 patulum 278 paupertinum 278 pectinatum 407 pendulum 408 pilosulum 633 platyphyllum 521 plumieri var. brasiliense 615 producens 634 pseudofilix-mas 279 puberulum 646 pumilum 459 roseum 273 rostratum 396 rutaceum 118 scabriusculum 223 scalare 636 semicordatum 240 sprengelii 625 strigilosum 231 tablanum 637 trejoi 517 trifoliatum 609 truncatulum 251 uliginosum 379 undulatum 409 varians 661 wallichianum 281 Aspidotis 5, 71 californica 71, 72, 732* meifolia 72, 732* Aspleniaceae 73, 91 Asplenium 3, 5, 72, 328, 373, 548 sect. Hymenasplenium 73, 86, 91, 118 sect. Sphenopteris 93, 122 abscissum 79, 80, 89, 92, 105, 108, 115, 128, 131, 736* achilleifolium 80, 81, 85, 94, 125, 739* acutiserratum 88, 131 adiantoides 113 adiantum-nigrum 87 var. adiantum-nigrum 87 adiantum-nigrum ⫻ trichomanes 105 aethiopicum 73, 113 alatum 81, 734* alfredii 130 alleopteron 116 altissimum 253 amabile 129 anisophyllum 95 anthriscifolium 115 arboreum 254 arcanum 81, 82, 743* arcuatum 105 argentinum 104 aspidiiforme 257

INDEX TO SCIENTIFIC NAMES Asplenium (Continued ) auriculatum 82, 83, 119, 738* auritum 73, 83, 89, 90, 97, 106, 107, 125, 126, 737* var. bipinnatifidum 89, 126 var. bipinnatisectum 89, 126 var. dissectum 131 var. mexicanum 89 var. monodon 106 var. subsimplex 106 var. tripinnatum 97 bangii 104 barbaense 83, 84, 104, 112, 124, 745* barclayanum 93 bertolonii 111 bicrenatum 119 bisectum 93 blechnoides 150 blepharodes 84, 96, 113, 119, 120, 130, 741* blepharophorum 81, 85, 94, 739* boliviense 104 bradeorum 104 brasiliense 104 breedlovei 85, 86, 101, 742* castaneum 86, 87, 96, 98, 99, 106, 110, 111, 119, 120, 735* chihuahuense 73, 87, 105, 743* cicutarium 88 var. paleaceum 93 cirrhatum 87, 88, 116, 131, 740* cladolepton 93, 107 claussenii 89, 104 clutei 125 commutatum 85 conchatum 80 conquisitum 118 cordovense 266 coriaceum 131 coriifolium 97 cristatum 88, 89, 108, 118, 129, 130, 740* cruegeri 89, 104, 105, 745* cuneatum 113 cuneifolium 87 ⫻curtissii 108 curvatum 105 cuspidatum 73, 83, 89, 90, 91, 97, 126, 738* var. foeniculaceum 97 var. tenerrimum 126 var. triculum 91 cyrtopteron 116 dalhousiae 73, 90, 743* delicatulum 72, 90, 91, 127, 742* delitescens 80, 91, 92, 745* dentatum 92, 744* denticulosum 254 dianae 92, 93, 746* diplazioides 254 diplosceum 84, 104 dissectum 93, 123, 733* distans 257

Asplenium (Continued ) divaricatum 108 dolosum 105 donnell-smithii 99 dubiosum 87 eatonii 81, 85, 93, 742* erectum var. harpeodes 99 excelsum 125 exiguum 3, 73, 94, 128, 129, 739* expansum 256 falcinellum 101, 102 feei 94, 95, 128, 737* fibrillosum 84, 87, 95, 96, 113, 119, 120, 130, 741* finckii 328 flabellulatum 96, 101, 116, 130, 746* var. dentatum 96 var. flabellulatum 96 var. partitum 96 foeniculaceum 97 formosum 96, 97, 130, 735* fournieri 82, 126 fragile 126 fragrans 82, 83, 90, 97, 738* var. mexicanum 89 franconis 257, 263 var. gracile 257 furcatum 113 galeottii 105, 136 galipense 131 gentryi 97, 98, 117, 120, 744* ghiesbreghtii 328 glenniei 94 gracile 94 grande 80, 81 grandifolium 259 haenkeanum 93 hahnii 255 hallbergii 98, 99, 106, 117, 120, 735* harpeodes 99, 102, 104, 112, 114, 736* harrisii 99, 100, 125, 744* hastatum 83 hesperium 111, 112 heterochroum 100, 106, 108, 109, 110, 111, 117, 741* hoffmannii 91, 100, 101, 103, 115, 736* incisoserratum 101, 103 induratum 260 insigne 122 insolitum 86, 101, 739* juglandifolium 101, 102, 746* karstenianum 131 kellermanii 109 klotzschii 254 komarovii 120 lacerum 89, 126 laciniatum 81 laetum 89, 91, 92, 101, 102, 103, 105, 736* lamprocaulon 103, 109, 739* leptophyllum 105 leucothrix 115

Asplenium (Continued ) levyi 83 limbatum 323 lindbergii 260 lividum 131 longissimum 152 lugubre 102 lunulatum var. latius 111 macilentum 83, 107 macraei 93 macrocarpon 85 macrurum 131 marginatum 323 marinum 72 martensii 136 melanorhachis 103 membranifolium 101 mexicanum 89 minimum 82, 103, 104, 116, 733* miradorense 73, 84, 89, 99, 104, 105, 111, 112, 114, 124, 734* modestum 105, 743* monanthemum 105 monanthes 3, 73, 86, 92, 97, 98, 99, 103, 105, 106, 111, 117, 120, 126, 128, 735*, 746* var. castaneum 86 monodon 83, 106, 107, 738* monteverdense 107 muenchii 107, 124, 737* munchii 107 myapteron 85 myriophyllum 88, 93, 107, 108, 118, 740*, 742* neglectum 261 nesioticum 100, 108, 109, 111, 741* nidus 373 nigricans 103 nigripes 548 obesum 103, 109, 746* obtusifolium 118 var. riparium 117 obtusissimum 105 ocoe¨nse 89 oligosorum 99, 109, 110, 120, 735* olivaceum 99, 109, 110, 120, 744* onopteris 87 onustum 85 otites 104, 115 palmeri 99, 100, 106, 108, 109, 110, 111, 117, 127, 128, 735* partitum 96 parvulum 116, 117 pendulum 99 peruvianum 126 pimpinellifolium 82 pinnatum 109 plantagineum 251, 262 plantaginifolium 262, 263 plenum 89 plumbeum 83

INDEX TO SCIENTIFIC NAMES Asplenium (Continued ) polymeris 111 polymorphum 79, 86 polyphyllum 99, 103, 106, 111, 128, 735* potosinum 84, 99, 104, 105, 111, 112, 121, 124, 737* var. incisum 121 var. semipinnatum 121 praemorsum 73, 112, 113, 123, 132, 733* pringlei 113, 120, 741* progrediens 122 pseuderectum 99, 113, 114, 734* pteropus 81, 99, 104, 112, 114, 115, 736* pulchellum 80, 104, 114, 115, 121, 736* var. subhorizontale 114 pumilum 3, 72, 82, 103, 104, 115, 116, 733* var. laciniatum 81 purdieanum 328 pyramidatum 89 raddianum 105 radicans 88, 96, 116, 129, 740* var. cirrhatum 87 var. dentatum 96 var. partitum 96 var. uniseriale 129 regulare var. latior 111 repandulum 118 resiliens 97, 98, 99, 100, 106, 108, 109, 110, 111, 116, 117, 735* rhizophyllum 116 var. proliferum 107 rigidum 90 riparium 91, 95, 117, 118, 128, 736* rosenstockianum 81 rubinum 86 rupium 90 rutaceum 88, 108, 118, 740* salicifolium 82, 83, 118, 119, 738* var. aequilaterale 119 var. auriculatum 82 sanchezii 86, 87, 98, 113, 119, 120, 741* sanguinolentum 95 sarcodes 95 schlechtendahlianum 104 sciadophilum 93 scolopendrium 72, 73, 120, 123 var. americanum 120, 737* var. lindenii 120 var. scolopendrium 120 seileri 104, 120, 124, 745* semicordatum 82 semipinnatum 88, 121, 124, 746* septentrionale 73, 121, 122, 743* subsp. septentrionale 122 serra 93, 122, 123, 733* serratum 123, 733* sessilifolium 107, 121, 123, 124, 130 var. guatemalense 107, 123, 124 var. occidentale 123, 124, 746* var. sessilifolium 123, 737*

Asplenium (Continued ) shepherdii 254 var. bipinnatum 257 skinneri 138 soleirolioides 100, 120, 124, 125, 126, 128, 739* solmsii 81, 86, 94, 125, 742* sphaerosporum 83, 125, 126, 738* standleyi 104, 105, 112 stolonipes 110, 120, 125, 126, 128, 739* striatum 263, 264 subalatum 96 subvestitum 85 tenerrimum 82, 91, 126, 739* tenue 126 ternatum 265 trichomanes 73, 97, 100, 105, 109, 111, 127, 128, 130, 744* subsp. quadrivalens 127 subsp. trichomanes 127 var. repens 110 var. vespertinum 150 var. viridissimum 111 trichomanes-dentatum 92 triphyllum 126 tryonii 127, 128, 744* tuerckheimii 95, 128, 736* ultimum 94, 128, 743* uniseriale 88, 89, 96, 116, 129, 740* venturae 129, 130, 742* verapax 115 vera-pax 266 verecundum 80, 107 vespertinum 130, 741* viridissimum 111 virillae 80, 130, 131, 745* williamsii 83, 84 yelagagense 88, 116, 131, 736* Astrolepis 64, 132, 443 beitelii 134 cochisensis 132, 133, 134, 748* subsp. arizonica 132, 133 subsp. chihuahuaensis 132, 133 subsp. cochisensis 133 crassifolia 133, 134, 135, 747* integerrima 133, 134, 135, 748* laevis 134, 135, 747* sinuata 132, 134, 135, 186, 187, 747* subsp. mexicana 135 subsp. sinuata 135 windhamii 133, 134 Athyrium 19, 135, 252, 323 achilleifolium 80 arcuatum 136, 137, 750*, 751* barnebyanum 136, 137 bourgaei 137 bourgeaui 136, 137, 750*, 751* conchatum 80 expansum 256 filix-femina 135, 136, 137 var. californicum 137

Athyrium (Continued ) var. cyclosorum 137 galeottii 136 gracile 94 macrocarpon 85 martensii 136 palmense 137, 138, 139, 749* paucifrons 137 pumilio 136 skinneri 136, 137, 138, 139, 749* tejeroi 138, 139, 749* tripinnatum 138 Aulaxina quadrangula 239 Azolla 139, 462, 546 sect. Azolla 139 sect. Rhizosperma 139 caroliniana 140, 141 filiculoides 139, 140, 141, 752* mexicana 140, 141 microphylla 140, 141, 752* Azollaceae 5, 382 Balantium 249 auricomum 249 Blechnaceae 141, 692 Blechnum 141, 466, 692 sect. Diafnia 141 sect. Lomaria 152 sect. Lomaridium 141 sect. Lomariocycas 141 sect. Parablechnum 141 acutum 144 ⫻aduncum 149 angustifrons 150 ⫻antillanum 147, 148 appendiculatum 143, 146, 148, 149, 150, 755*, 756* appendiculatum ⫻ gracile 143 appendiculatum ⫻ meridense 148 appendiculatum ⫻ occidentale 149 appendiculatum ⫻ polypodioides 143 binervatum 141, 144, 146 subsp. fragile 145 blechnoides 150 ⫻caudatum 146, 147, 757* ⫻confluens 149, 153, 757* cordatum 151 danaeaceum 151 divergens 141, 143, 144, 147, 152, 153, 756* ensiforme 141, 144, 145, 146, 754*, 756* ⫻falciculatum 149 falciforme 145, 151, 754* fragile 141, 144, 145, 146, 150, 754* fraxineum 145, 148 ghiesbreghtii 152 glandulosum 143 gracile 143, 146, 147, 148, 149, 755* gracile ⫻ occidentale 146, 757* indicum 152 intermedium 146 jamaicensis 144

INDEX TO SCIENTIFIC NAMES Blechnum (Continued ) l’herminieri 147 subsp. lehmannii 147 lehmannii 141, 144, 147, 152, 153, 753* longifolium 146, 147 maxonii 147 meridense 146, 147, 148, 149, 757* meridionale 143 nudum 141 occidentale 141, 143, 147, 148, 149, 150, 755*, 756* var. intermedium 143 var. minor 143 var. pubirhachis 143 occidentale ⫻ polypodioides 149, 153, 757* orientale 148 penna-marina 3, 149, 150, 756* polypodioides 143, 145, 148, 149, 150, 755* procerum 141 radicans 692 scabrum 150 schiedeanum 141, 145, 151, 754* schlimense 146 serrulatum 141, 151, 152, 755* spicant 141 stoloniferum 141, 149, 150, 152, 153, 753*, 756* subdimorphum 146 triangulare 153 unilaterale 150 varians 151 virginicum 692 wardiae 141, 144, 147, 152, 153, 753*, 756* Blotiella 371 Bolbitis 153, 369, 475 aliena 154, 157, 158, 759* bernoulii 154, 157, 760* cladorrhizans 156 donnell-smithii 154 hastata 155, 157, 760* hemionitis 155, 156, 157, 760* liebmannii 155 mexicana 157 pergamentacea 156, 157, 760* portoricensis 153, 154, 156, 157, 758* serratifolia 153, 157, 759* umbrosa 154, 157, 158, 759* Bommeria 158, 187, 325 ehrenbergiana 158, 761* elegans 159, 761* hispida 159, 160, 161, 761* knoblochii 160 pedata 160, 762* podophylla 158 schaffneri 159 subpaleacea 159, 160, 161, 762* Botrychium 161 sect. Osmundopteris 161 subg. Osmundopteris 161 brachystachys 164 cicutarium 164, 165

Botrychium (Continued ) decompositum 162, 163, 763*, 764*, 765* dissectum 162 f. obliquum 161 subsp. decompositum 162 lunaria 161 var. minganense 164 matricariifolium 164 minganense 162, 164, 763* multifidum 164 pusillum 163 schaffneri 163, 164, 766* var. pusillum 163 silaifolium 164 socorrense 163, 164, 766* tolucaense 163, 164, 763* virginianum 161, 164, 764* subsp. meridionale 164 var. meridionale 164 var. mexicanum 164 ⫻watertonense 164 Botryogramma 368 karwinskii 368 Botrypus 161 virginianus 161 Bryodesma 550, 551 arizonica 559 arsenei 559 asprella 560 bigelovii 563 cinerascens 565 eremophila 568 extensa 569 mutica 583 parishii 586 peruviana 587 rupestris 550 rupincola 592 sartorii 593 sellowii 595 steyermarkii 598 underwoodii 601 viridissima 601 wrightii 602 Byrsopteris 63 aristata 63 denticulata 63 Caenopteris achilleifolia 80 myriophylla 107 Callipteris 252 Calochlaena 233 Campyloneurum 165, 477 amphostenon 166, 167, 168, 169, 172, 767*, 768* var. amphostenon 167 var. irregulare 167 angustifolium 165, 166, 167, 168, 169, 767*, 768* var. amphostenon 166 var. ensifolium 168

Campyloneurum (Continued ) angustipaleatum 167 aphanophlebium 168, 171, 172 brevifolium 169 caudatum 171 costatum 168, 768* densifolium 167, 168 ensifolium 167, 168, 169, 172, 768* fasciale 170, 171 latum 169 lorentzii 166 occultum 171, 172 phyllitidis 165, 169, 767* repens 165, 168, 170, 171, 768* serpentinum 165, 168, 170, 171, 172, 767* tenuipes 171, 172, 767* xalapense 165, 168, 171, 172, 767* Cassebeera cuneata 190 Ceradenia 172, 315, 361, 615 subg. Ceradenia 172 subg. Filicipecten 173, 174 brunneoviridis 173 curvata 172 jungermannioides 172, 173, 316, 317, 769* kookenamae 173 margaritata 173, 361, 769* oidiophora 173, 174, 769* pruinosa 172, 174, 769* Ceratodactylis 368 osmundoides 368 Ceratopteridaceae 175 Ceratopteris 5, 174 cornuta 176 pteridoides 174, 175, 176, 770* richardii 175, 176 thalictroides 174, 175, 176, 770* Ceropteris schaffneri 464 serrata 464 Ceterach 90 dalhousiae 73, 90 Ceterachopsis dalhousiae 90 Cheilanthes 3, 5, 20, 71, 132, 176, 213, 270, 324, 325, 402, 411, 443 aemula 180, 203, 774* affinis 413, 422 alabamensis 180, 181, 203, 780* albida 197 albofusca 71 allosuroides 181, 774* angustifolia 182, 187, 191, 195, 206, 402, 773* apiacea 182, 183, 195, 778* arizonica 183, 208, 773*, 779* aschenborniana 414 aspera 195 aspidioides 204 aurantia 414 aurantiaca 176, 184, 193, 411, 414, 777* aurea 176, 184, 193, 197, 411* var. aurea 184, 185, 772* var. palmeri 184, 185, 777*

INDEX TO SCIENTIFIC NAMES Cheilanthes (Continued ) beitelii 134 bolborrhiza 185, 771* bonariensis 3, 185, 212, 213, 776* brachypus 186, 187, 776* brandegeei 187, 777* californica 72, 416 cancellata 65 candida 416, 417, 422 var. copelandii 417 canescens 687 castanea 192 chaerophylla 182, 187, 191, 192, 772* chipinquensis 187, 188, 784* cinnamomea 192 clevelandii 188, 190, 204, 782* var. clevelandii 188 var. clokeyi 188 cochisensis 132 complanata 183, 188, 189, 191, 192, 195, 208 var. complanata 189, 779* var. interrupta 189, 779* concolor 269 ⫻coruscans 423 covillei 188, 189, 190, 204, 213, 783* crassifolia 133 crenulata 200 cucullans 181, 190, 203, 205, 774* cuneata 182, 187, 190, 191, 772*, 773* davenportii 419 decomposita 182, 187, 191, 206, 779* decurrens 189, 191, 192, 779* deltoidea 134 deserti 416 eatonii 192, 210, 784* f. castanea 192 farinosa 71, 176, 192, 193, 417, 772* feei 193, 194, 785* fendleri 190, 194, 780* ferruginea 186 fimbriata 3, 194, 203, 774* formosa 66 fournieri 203 frigida 196 galeottii 418 glandulifera 196 gracilis 193 grayi 418 greggii 419 gryphus 324 harrisii 189 hintoniorum 183, 194, 195, 206, 208, 778* hirsuta 200, 209 hookeri 424 horridula 195, 780* incana 66 integerrima 133 intermedia 203 intramarginalis 403 jamaicensis 3, 195, 196, 784* jonesii 67

Cheilanthes (Continued ) kaulfussii 196, 211, 775* lanuginosa 196 lemmonii 420 var. australis 421 lendigera 176, 194, 196, 197, 201, 202, 204, 213, 775*, 785* var. minor 196 lerstenii 183, 184 leucopoda 176, 193, 197, 785* limitanea 67 var. mexicana 68 lindheimeri 197, 198, 213, 782* longipila 198, 201 var. brevipila 198, 786*, 789* var. longipila 198, 199, 775*, 789* lozanii 199 lozanoi 199 var. lozanoi 199, 200, 783* var. seemannii 199, 200, 783* lutea 184 marginata 3, 183, 200, 202, 209, 210, 787*, 788* marsupianthes 197, 201, 213, 785* marsupianthes ⫻ mexicana 197 maxoniana 198, 201, 789* meifolia 72 membranacea 183, 201, 202, 773*, 777* mexicana 197, 202, 785* mickelii 202, 203, 774* micromera 190 microphylla 20, 180, 181, 190, 194, 203, 205, 780* var. fimbriata 194 var. micromera 190 micropteris 176 minor 196 moncloviensis 207 myriophylla 203, 204, 775*, 782* nealleyi 423 neglecta 422 newberryi 190, 204, 786* notholaenoides 181, 190, 203, 204, 205, 774*, 789* ochracea 184 paleacea 203 pallens 69 palmeri 185 parryi 190, 194, 205, 786* parvifolia 69 pellaeopsis 183, 195, 205, 206, 208, 778* peninsularis 206 var. insularis 206, 207, 781* var. peninsularis 206, 781* pinkavii 192 potosina 423, 424 pringlei 194, 207, 419 var. moncloviensis 207, 208, 781* var. pringlei 207, 208, 781* pubescens 203 purpusii 195, 206, 208, 778*

Cheilanthes (Continued ) pyramidalis 183, 200, 202, 208, 209, 772*, 788* subsp. arizonica 183 var. arizonica 183 radiata 20 repens 352 rigida 214, 422 rosei 423 rufa 192 rufescens 200 schaffneri 192, 423 seemannii 20, 199 selinoides 201, 202 sinuata 135 var. cochisensis 132 skinneri 3, 185, 209, 324, 325, 327, 771* sonorensis 207 speciosissima 511, 523 var. elata 523 spiculata 210, 787* squamosa var. brachypus 186 sulphurea 425 tomentosa 188, 192, 196, 210, 784* venusta var. aurata 182 villosa 192, 210, 211, 782* viscida 211, 786* viscosa 196 wootonii 204, 211, 212, 213, 783* wrightii 181, 212, 781* yatskievychiana 212, 213, 776* yavapensis 212, 213, 782* Cheiloplecton 213, 411, 443 rigidum 213, 214 var. lanceolatum 214, 215, 790* var. rigidum 214, 215, 790* Cheiroglossa 429 palmata 429, 431 Christella 639 augescens 640 cretacea 641 dentata 642 hispidula 643 lanosa 645 normalis 644 ovata 646 patens 646 puberula 646 Chrysochosma 411 affinis 413 aliena 413 angusta 413 aschenborniana 414 aurantiaca 414 californica 416 candida 417 var. copelandii 417 galeottii 418 grayi 418 greggii 419 hookeri 424

INDEX TO SCIENTIFIC NAMES Chrysochosma (Continued ) lemmonii 420 var. australis 421 leonina 421 neglecta 422 rigida 422 rosei 423 schaffneri 423 var. nealleyi 423 sulphurea 425 Chrysodium 17 lomarioides 18 var. hastatum 18 Chrysopteris 461 dulcis 488 Cibotium 5, 215, 233, 249 chamissoi 215 guatemalense 215 horridum 603 princeps 603 regale 215, 791* schiedei 215, 216, 791* wendlandii 215 Cnemidaria 5, 216 apiculata 216, 217, 792* decurrens 217, 239, 792* speciosa 216 Cochlidium 217 graminoides 217 linearifolium 218, 317, 793* rostratum 218, 317, 793* serrulatum 219, 694, 793* Coniogramme americana 327 subcordata 327 Coptophyllum cicutarium 50 Cornopteris 252 Craspedaria 396 vacciniifolia 396 Ctenitis 63, 219, 358, 379, 386, 475, 609 sect. Subincisae 386 atrogrisea 387 baulensis 221, 229, 795* bullata 221, 224, 232, 796* chiapasensis 222, 797* chiriquiana 222 costaricensis 225 distans 219 effusa 359 equestris 222, 224, 225, 226, 227, 228, 229, 231, 233, 379 var. equestris 222, 223, 798* var. erosa 223, 228, 798* erinacea 223, 224, 226, 796* excelsa 222, 224, 226, 231, 794* grisebachii 225, 226, 796* hemsleyana 223, 224, 225, 226, 798* interjecta 223, 226, 229, 231, 797* karsteniana 388 lanceolata 226, 230, 795* lindenii 230 lunensis 387 melanosticta 223, 226, 227, 794*

Ctenitis (Continued ) mexicana 222, 224, 228, 231, 797* microchlaena 228, 230, 232, 233, 795* molinae 225 nigrovenia 221, 229, 233, 794* pansamalensis 387 pulverulenta 388 var. heydei 388 refulgens 229, 795* salvinii 230, 795* sloanei 222, 229, 230, 797* strigilosa 221, 231, 232, 796* subincisa 388 submarginalis 229, 232, 233, 794* thelypteroides 229, 232, 233, 795* tonduzii 221 ursina 224, 233, 796* Ctenopteris 435, 440 anfractuosa 616 apiculata 361 aromatica 391 asplenifolia 617 cultrata 617, 618 delicatula 618 firma 391 leptostoma 392 megaloura 394, 395 melanosticta 392 mollissima 618 moniliformis 393 pilosissima 394 plumula 441 pruinosa 174 punctata 172, 174 rigens 394 rigescens 391 semihirsuta 619 senilis 620 subtilis 621 Culcita 5, 233, 249, 395 coniifolia 233, 799* macrocarpa 233 Cyathea 5, 234, 603 apiculata 216 arborea 234 aristata 216 articulata 45 bicrenata 235, 236, 800* bourgaei 603 bourgeaui 603 corcovadensis 234 costaricensis 236, 801* decurrens 217 decurrentiloba 217 delicatula 237 divergens 237 divergens var. divergens 237 var. tuerckheimii 236, 801* elegantissima 217 firma 45 fulva 237, 801* glauca 603

Cyathea (Continued ) godmanii 236, 237, 802* guatemalensis 217 horrida 603 jurgensenii 236 liebmanii 217 lucida 217 marginalis 239 mexicana 45, 217, 237 microdonta 238, 802* multiflora 234 muenchii 603 munchii 603 myosuroides 238, 800* polystichoides 45 princeps 603 salvinii 46 schiedeana 239, 800* trejoi 45 tuerckheimii 236 valdecrenata 237 Cyatheaceae 268, 395, 466 Cyclodium 63, 153, 428, 475, 607 Cyclopeltis 239, 369 semicordata 239, 240, 803* Cyclosorus 622, 638, 644 dentatus 642 gongylodes 638, 643 interruptus 643 tottus 643 Cyrtomidictyum 511 Cyrtomiopsis 455 Cyrtomium 455, 511 auriculatum 456 guatemalense 457 juglandifolium 457 macrosorum 457 nobilis 458 pumilum 459 remotispora 459 umbonatum 460 Cystopteris 240, 687 acuta 241 diaphana 241 fragilis 240, 241, 242, 804*, 805* subsp. diaphana 241 var. palmensis 137 membranifolia 4, 241, 804* millefolia 242, 805* reevesiana 241, 242 Danaea 242, 380 cuspidata 242, 243, 806* elata 244 elliptica 243, 806* media 244 moritziana 243 muenchii 242 munchii 242 nodosa 242, 243, 244, 806* Darea thecifera 373

INDEX TO SCIENTIFIC NAMES Davallia divaricata 426 ferulacea 544 inaequalis 545 lindenii 373 multiflora 406 saccoloma 545 schlechtendalii 426 tegularis 346 thecifera 373 theciferum 373 uncinella 425 Davalliaceae 404, 427 Dennstaedtia 244 adiantoides 245 arborescens 245, 247 auriculata 245, 809* bipinnata 245, 546, 807* cicutaria 246, 247, 249, 808* cornuta 246, 247, 249, 807* coronata 247, 809* decomposita 249 dissecta 247 distenta 246, 247, 248, 808* flaccida 244 globulifera 248, 807* gracilis 248 mexicana 248 muenchii 350 munchii 350 obtusifolia 249 rubicaulis 351 rubiginosa 246 spinosa 248, 249, 809* Dennstaedtiaceae 5 Deparia 252 Dicksonia 5, 233, 234, 249, 371 adiantoides 245 var. coronata 247 arborescens 249 bipinnata 245 cicutaria 246 coniifolia 233 cornuta 246 coronata 247 culcita 233 distenta 247 ghiesbreghtii 249 gigantea 249, 250 globulifera 248 obtusifolia 249 rubiginosa 246 schiedei 215 sellowiana 249, 810* umbrosa 246 Dicksoniaceae 5, 466 Dicranopteris 250, 268, 604 sect. Acropterygium 321 subg. Acropterygium 321 bancroftii 268 bifida 605 dichotoma 250 flexuosa 250, 251, 322, 811*

Dicranopteris (Continued ) fulva 605, 606 linearis 251 palmata 606 pectinata 322 speciosa 250 underwoodiana 607 Dictymia 372 Dictyoxiphium 609, 614 panamense 609, 613 Didymochlaena 251 sinuosa 251 truncatula 251, 812* Didymoglossum 669 angustifrons 672 hymenoides 677 krausii 678 muscoides 669, 677 ovale 679 petersii 679 reptans 682 Diphasiastrum 375 complanatum 375 thyoides 376 Diplaziopsis 323 Diplazium 19, 91, 136, 251, 323, 324 acutale 261 altissimum 253, 254, 255, 814* amplum 259 anthraxacolepis 267 arboreum 254 biolleyi 265 camptocarpon 257 cordovense 266 crenulatum 264 cristatum 254, 261, 267, 268, 814* croatianum 255 delitescens 91 denticulosum 254 diplazioides 253, 254, 255, 259, 260, 816* donnell-smithii 255, 258, 815* drepanolobium 255, 256, 258, 261, 267, 814* drepanolobium ⫻ lonchophyllum 258 elongatum 264 entecnum 253 errans 256, 818* expansum 256, 257, 259, 260, 265, 816* falcatum 267 feei 267 flavescens var. proliferum 262 flavoviride 323 franconis 255, 257, 258, 161, 815* franconis ⫻ lonchophyllum 258 gomezianum 258, 267, 817* grandifolium 259, 324, 818* var. andicola 259 var. grandifolium 259 hahnii 255 hellwigii 257, 259, 816* herbaceum 258 hians 257, 259, 260, 814*

Diplazium (Continued ) inaequilaterum 260 induratum 260 ingens 257 klotzschii 254 laciniatum 254, 255 l’herminieri 255 limbatum 323 lindbergii 260, 264, 268, 816* lonchophyllum254, 256, 258, 260, 261, 267, 268, 813* marattiifolium 265 marginatum 323 matudae 264, 265 neglectum 261, 262, 817* obscurum 261, 262, 324, 817* pectinatum 268 plantaginifolium 251, 262, 265, 266, 815* prominulum 261, 268 puberulentum 263, 814* pycnocarpon 323 riedelianum 266 striatastrum 263, 264, 265, 816* striatum 257, 260, 264, 265, 814* ternatum 263, 265, 815* tussacii 260 urticifolium 262, 264, 265, 816* vera-pax 256, 266, 817* villosum 268 wendtii 3, 266, 267, 814* werckleanum 254, 256, 266, 267, 815* Diplopterygium 268, 321, 604 bancroftii 268, 819* farinosum 269 glaucum 268 Diplostachium tenellum 600 Doryopteris 269 concolor 269, 270, 820* fournieri 209 palmata 269, 270, 820* pedata 270 var. palmata 270 Drynaria crassinervata 471 mexicana 472 stenoloma 469 tortulosa 469 vestita 474 Dryopteridaceae 5, 251, 404, 427, 428 subfam. Dryopteridoideae 5 subfam. Lomariopsidoideae 5 subfam. Tectarioideae 5 Dryopteris 19, 63, 219, 241, 270, 455, 475 sect. Cinnamomeae 271 sect. Fibrillosae 271 subg. Ctenitis 219 subg. Ctenitis sect. Subincisae 386 subg. Cyclosorus 638 subg. Dryopteris 271 subg. Goniopteris 648 subg. Leptogramma 667 subg. Meniscium 663 subg. Nephrocystis 271

INDEX TO SCIENTIFIC NAMES Dryopteris (Continued ) subg. Steiropteris 668 subg. Stigmatopteris 607 amplificata 626 angustifolia 664 aperta 278 arguta 272, 274, 822* athyrioides 273 atrogrisea 387 atrovirens 624 augescens 640 var. geropogon 640 var. puberula 646 balbisii 625 blanda 651 caespitosa 651 caucaensis 626 cheilanthoides 626 chiapasensis 222 chrysocarpa 280 cinnamomea 271, 273, 274, 275, 276, 277, 279, 281, 824*, 825* cinnamomea ⫻ knoblochii 271 columbiana 632 concinna 627 davenportii 223 deflexa 628 deltoidea 668 dentata 642 denticulata 63 f. jucunda 63 diplazioides 628 diplazoides 628 domingensis 652 effusa 359 equestris 222 var. mentiens 223 excelsa 224 var. guatemalensis 360 falcata 665 feei 646 filix-mas 270, 272, 274, 280, 822* flaccisquama 279 fournieri 273 futura 271, 274, 275, 822* geropogon 640 ghiesbreghtii 651 glandulifera 273 glandulosa 668 var. brachyodus 668 goldiana 271 grisebachii 225 guadalupensis 652 guatemalensis 360 hemsleyana 225 huatuscensis 233 indecora 273 interjecta 226 juergensenii 665 jungersenii 665 karsteniana 388

Dryopteris (Continued ) var. heydei 388 karstenii 228 karwinskyana 275, 823* knoblochii 271, 273, 276, 279, 281 lanceolata 226 var. deltoideolanceolata 227 var. tricholepis 227 leptocladia 654 liebmannii 655 lindenii 230 lindigii 628 linkiana 628 longicaudata 608 lunensis 387 maxonii 276, 277, 821* melanochlaena 630 melanosticta 227 meniscioides 655 mexicana 273 micradenia 277 microchlaena 228 mollis 642, 651 muenchii 277, 822* munchii 277 nephrodioides var. biolleyi 650 nicaraguensis 657 var. minor 656 nigrovenia 229 normalis 644 var. lindheimeri 645 nubigena 271, 275, 277, 278, 279, 824*, 825* obliterata 657 oligocarpa 632 ovata 646 paleacea 281 pallida 272 panamensis 634 var. gonzalezii 634 var. proxima 634 pansamalensis 387 parallelogramma 280, 281 patens 646 var. lanosa 645 patula 271, 273, 274, 275, 277, 278, 279, 281, 823*, 824* var. chaerophylloides 279 var. moreliae 276 var. rossii 276, 280 var. serrata 279 paucipinnata 658 pilosa 667 var. procurrens 667 pilosohispida 632 pilosula 633 poiteana 658 polyphylla 651 praetermissa 659 pseudofilix-mas 271, 274, 279, 280, 281, 823*

Dryopteris (Continued ) pseudofilix-mas ⫻ wallichiana 271 pseudototta 667 puberula 646 pulverulenta 388 quadrangularis 643 refulgens 229 reptans 659 resiliens 660 resinifera 633 reticulata 665 rosea 273, 274 rossii 271, 276, 277, 279, 280, 825* rudis 635 salvinii 230 sancta 635 sanctiformis 627 scalaris 636 schaffneri 661 semicordata 240 serrata 666 simplicior 278, 279 sordida 608 sprengelii 625 standleyi 666 stuebelii 637 strigilosa 231 var. cookii 231 struthiopteroides 637 stuebelii 637 subincisa 386, 388 submarginalis 232 tablana 637 tetragona 662 thelypteris 669 thomsonii 637 tremula 669 tricholepis 227 tuerckheimii 648 uliginosa 379 wallichiana 3, 271, 280, 281, 823* ⫻Dryostichum singulare 271 Elaphoglossum 3, 153, 282, 369, 475 subsect. Peltapteris 282 acutissimum 298 affine 285, 286, 288, 299, 307, 313, 829* alansmithii 286, 313, 843* albomarginatum 286, 287, 835* apodum var. latum 295 araneosum 300, 315 auricomum 287, 307, 313, 834* blepharodes 289 blumeanum 304 callolepis 291 catherinae 310 chiapense 311 clewellianum 288 conforme 282 crinipes 310 crinitum 282

INDEX TO SCIENTIFIC NAMES Elaphoglossum (Continued ) decursivum 287, 288, 298, 833* dissitifrons 288, 834*, 842* elongatum 311 engelii 288, 289, 837* erinaceum 282, 289, 290, 302 var. erinaceum 289, 829*, 841* var. occidentale 289, 835* eucraspedum 291 eximium 290, 835* fimbriatum 289 glabellum 290, 303, 828* glaucum 291, 826* gratum 291, 292, 293, 300, 834* guatemalense 286, 291, 292, 297, 307, 833*, 840* hartwegii 292, 293, 837* herminieri 293, 839* hirtum 301 hookerianum 300 huacsaro 293, 294, 304, 837* intermedium 300 ipshookense 294, 831* jaliscanum 294, 836* lanceum 295, 832* latifolium 309 latum 295, 296, 835* laxipes 307, 308 leebrowniae 296, 309, 827* leonardii 296, 297, 832* lepidopodum 297, 826*, 840* liebmannii 297 lindenii 282, 297, 298, 300, 305, 833* lingua 309 lonchophyllum 288, 298, 828* manantlanense 298, 299, 838* mathewsii 293 mcvaughii 286, 288, 299, 839* minutum 313 monicae 298, 299, 300, 310, 836* muelleri 292, 300, 305, 315, 831* muscosum 289, 300, 301, 830* nanchititlense 308 nervosum 282, 304 nitidum 289 obscurum 301, 838* paleaceum 3, 287, 299, 301, 302, 307, 308, 313, 830* pallidum 282, 287, 302, 829* parduei 291, 302, 828*, 833* peltatum 282, 303, 307, 311, 832* f. peltatum 303 petiolatum 292, 300, 303, 304, 305, 306, 315, 831* piloselloides 294, 295, 298, 304, 306, 832* pilosius 304, 305, 836* pilosum 300 potosianum 300, 304, 305, 841* pringlei 292, 304, 305, 306, 308, 830* pumilum 297

Elaphoglossum (Continued ) pusillum 306, 836* rampans 306 revolutum 306, 307, 834* roezlii 307 rubescens 307, 839* rufescens 307, 308, 838* rzedowskii 300, 304, 306, 308, 837* salicifolium 304 sartorii 288, 291, 296, 308, 309, 314, 826*, 828*, 840* scapellum 309 semicylindricum 302 seminudum 288, 299, 309, 828* setigerum 300, 309, 315, 832* setosum 310, 836* smithii 295 spatulatum 304 squamipes 303, 307, 310, 311, 829* squamosum 301 squarrosum 311, 833* stellatum 312 stelligerum 304 subaequale 310 tambillense 302 tectum 294, 311, 831* tenuiculum 312, 842* tenuifolium 286, 307, 312, 313, 829* trichomallum 315 trichomidiatum 305 venustum 297, 315 vestitum 287, 301, 307, 313, 830* villosum 310 viride 313, 314, 843* viscosum 303 xanthopodum 309, 314, 827* yourkeorum 309, 314, 827* yunnanense 304 Enterosora 172, 315, 361, 615 campbellii 315 ecostata 316, 844* parietina 173, 316, 844* percrassa 316 trichosora 317, 844* Equisetaceae 5, 434 Equisetum 4, 317 arvense 317 ⫻ferrisii 318, 319, 846* fluviatile 317 giganteum 319, 320 ⫻haukeanum 318, 319, 846* hyemale 318, 319 subsp. affine 318 var. affine 318, 845*, 846* hyemale ⫻ myriochaetum 318 kansanum 318 laevigatum 318, 319, 846* myriochaetum 318, 319, 320, 845*, 846* robustum var. affine 318 ⫻schaffneri 319, 846*

Eriosorus 5, 61, 320, 357, 463 aureonitens 320 flexuosus 174, 320, 321, 426, 427 var. flexuosus 320, 847* var. galeanus 320, 847* hirtus 321, 847* var. hirtus 321 scandens 320 Feea diversifrons 675 Filicula digitata 342 Filix femina 530 Gleichenella 250, 268, 321, 604 pectinata 251, 321, 322, 848* Gleichenia 268, 321, 604 sect. Acropterygium 321 subg. Diplopterygium 268 subg. Mertensia sect. Diplopterygium 268 bancroftii 268, 269 bifida 605 brevipubis 605 circinnata 604 flexuosa 250 furcata 607 glauca 268 liebmannii 605 nitida 322 palmata 606 pectinata 322 underwoodiana 607 Gleicheniaceae 268, 321 Glyphotaenium nesioticum 316 Goniophlebium sect. Lepicystis 477 sect. Lopholepis 396 anisomeron 499 areolatum 461 calaguala 489, 499 chnoodes 487 collinsii 485 eatonii 488 fraxinifolium 491 haenkei 510 inaequale 500 incanum 477, 501 invertens 489 lepidotrichum 494 molestum 499 plectolepis 489 plesiosorum 499 pringlei 488 pyrrholepis 504 revertens 491 rhachipterygium 505 rhagadiolepis 510 sanctae-rosae 508 serratum 511 stramineum 500 thyssanolepis 509

INDEX TO SCIENTIFIC NAMES Goniopteris 622, 648 biolleyi 650 crenata 648, 658 domingensis 652 hatchii 652 imbricata 653 leptocladia 654 mollis 651 obliterata 657 poiteana 658 reptans 659 tetragona 662 toganetra 663 Grammitidaceae 5, 172, 372, 389, 400, 440, 616 Grammitis 217 andicola 391, 392 anfractuosa 616 apiculata 361 aromatica 391 asplenifolia 617 basiattenuata 400 var. valens 400 blepharodes 401 bryophila 217 cultrata 617 delicatula 618 delitescens 362, 363 deltata 390 elongata 470 firma 391 flabelliformis 391, 392, 401 flexuosa 174, 320 graminoides 217 harrisii 316 hellwigii 362 heteromorpha 621 hirta 321 jungermannioides 173 katasophistes 391 kirkbridei 174 lanceolata 372, 470 leptostoma 392 limula 363 linearifolia 218 linkiana 628 margaritata 173 maxoniana 174 melanosticta 392 minuscula 317 mitchellae 363 mollissima 618 moniliformis 393 myosuroides 362, 364 nesiotica 316 oidiophora 173 parietina 316 pilosissima 394 prionodes 363 pruinosa 174 revoluta 470

Grammitis (Continued ) rigens 394 rigescens 391 rostrata 218 rosulata 316 salvinii 372 semihirsuta 619 var. fuscosetosa 619 senilis 620 serrulata 219 skutchii 364 subflabelliformis 620 subtilis 621 taenifolia 401 trichomanoides 401 trichosora 317 xiphopteroides 394, 395 yarumalensis 316 zempoaltepetlensis 395 Gymnocarpium 240 Gymnogramma candida 66 chaerophylla 62 dealbata 464 cheilosora 629 cheilosorus 629 diplazioides 628 ehrenbergiana 158 var. muralis 159 elongata 470 flexuosa 320 hispida 159 leptophylla 62 var. mexicana 62 mexicana 372 pedata 160 pilosa 667 var. major 667 podophylla 158 polypodioides 628 procurrens 667 pumila 323 salvinii 372 sinuata 135 subcordata 327 triangularis 453 viscosa 454 Gymnopteris 325 aliena 154 bernoullii 154 donnell-smithii 154 hastata 155 hispida 159 pedata 160 portoricensis 156 rufa 325, 327 subcordata 327 tuerckheimii 154 Gymnotheca laxa 381

Haplopteris 544, 684 Hecistopteris 322, 544 pinnatifida 322 pumila 322, 323, 849* Hemianemia brandegeea 50 intermedia 53 Hemidictyum 323 marginatum 323, 850* Hemionanthes 324, 327 gryphus 209, 324, 851* Hemionitis 325 cajenensis 526 citrifolia 61 elegans 159 hederifolia 159 lanceolata 527 levyi 325, 852* lineata 525, 528 otonis 325 palmata 325, 326, 327, 852* palmata ⫻ pinnatifida 326 pedata 160 pinnatifida 326, 852* podophylla 158 rufa 325, 326, 327, 853* ⫻smithii 327 subcordata 209, 324, 325, 327, 853* Hemiphlebium krausii 678 membranaceum 678 petersii 679 reptans 682 Hemistegia decurrens 217 elegantissima 217 lucida 217 mexicana 217 Hemitelia 234 apiculata 216 costaricensis 236 decurrens 217 firma 45 guatemalensis 217 lucida 217 mexicana 217 multiflora 234 Heterophlebium grandifolium 537 Hewardia 21, 44 adiantoides 21 Hicriopteris 250 bancroftii 268 speciosa 250 Histiopteris 244, 327 incisa 327, 328, 854* vespertilionis 327 Holodictyum 5, 73, 123, 328 finckii 328, 329 ghiesbreghtii 328, 329, 855* Homalosorus pycnocarpon 323 Huperzia 329, 374, 376 acerosa 335 beiteliana 329, 330, 859*

INDEX TO SCIENTIFIC NAMES Huperzia (Continued ) bradeorum 334 capillaris 330, 333, 860* crassa 331, 358, 856* cuernavacensis 331, 336, 339, 857* dichotoma 331, 337, 860* filiformis 335 hartwegiana 331 hippuridea 332, 334, 856* lechleri 332 linifolia 330, 332, 333, 334, 858* mandioccana 332 mexicana 332 montana 332 myrsinites 333, 857* orizabae 333, 334, 859* pithyoides 332, 334, 856* pringlei 334, 335, 857*, 860* reflexa 335, 856* sargassifolia 336 selago 329 serrata 3, 335, 336, 859* setacea 335 taxifolia 331, 336, 337, 858* tuerckheimii 334, 335, 860* verticillata 335 wilsonii 337, 332, 858* Hydroglossum heterodoxum 377 spectabile 377 Hymenasplenium 73 Hymenodium 282 crinitum 282 Hymenophyllum 337 subg. Mecodium 337, 345 abruptum 339, 865* apterum 347 asplenioides 339, 863* axillare 344 botryoides 345 brevifrons 340 brevistipes 339, 340, 345, 863*, 865* ciliatum 342 var. nudipes 342 contortum 341 crassipetiolatum 340, 346, 866* crispum 340, 861* ectocarpon 341, 342, 348, 864* elegantulum 346 fendlerianum 341, 348, 863* fragile 342, 862* fucoides 341, 342, 348, 864* var. frigidum 347 hirsutum 337, 342, 343, 861* intercalatum 342 jalapense 342 lanatum 343, 862* lindenii 340 maxonii 343, 344, 866* microcarpum 340, 344, 861*

Hymenophyllum (Continued ) millefolium 345 myriocarpum 344, 348, 863*, 865* nigrescens 344 nigricans 344 paucicarpum 345, 865* polyanthos 337, 345, 864* pulchellum 345, 346, 861* schiedeanum 340 sieberi 346, 866* siliquosum 345 tegularis 346, 347, 862* trapezoidale 347, 862* tunbrigense 337, 342, 347, 348, 864* undulatum 341, 348, 865* var. fendlerianum 341 Hymenostachis diversifrons 675 Hypochlamys pectinata 268 Hypolepis 244, 348, 529 sect. Aspidotis 71 blepharochlaena 349, 350, 352, 867* bogotensis 353 brooksiae 351 californica 71, 72 eurychlaena 350 hispaniolica 351 melanochlaena 350, 868* mexicana 350, 352 microchlaena 350, 352, 868* muenchii 350, 351, 353, 868* munchii 350 nigrescens 348, 351, 867* pulcherrima 351, 352, 868* radiata 20 repens 349, 350, 352, 867* stuebelii 352, 868* tenuifolia 348 thysanochlaeana 350, 353, 354, 867* trichochlaena 350, 353, 354, 867* viscosa 353 Isoe¨taceae 5, 551 Isoe¨tes 354, 551, 594 cubana 357 howellii 354, 355, 870* jamaicensis 351 lacustris 354 mexicana 355, 356, 869*, 870* mexicana ⫻ montezumae 870* minima 357 montezumae 354, 355, 356, 870* nuttallii 357 orcuttii 354, 356, 357, 870* pallida 357, 870* pringlei 357, 869*, 870* Jamesonia 5, 320, 357, 463 alstonii 331, 358, 406, 871* pulchra 357

Lastrea ciliata 222 indecora 273 poiteana 658 recedens 358 thelypteris var. pubescens 669 Lastreopsis 219, 358, 379, 386 chontalensis 360 effusa 358, 360 subsp. dilatata 359, 360, 872* subsp. divergens 359, 360, 872* exculta subsp. exculta 360 subsp. guatemalensis 360, 872* subsp. squamifera 360 guatemalensis 360 recedens 358 tenera 358 Lecanium membranaceum 678 Lecanolepis membranacea 678 Lellingeria 219, 360, 389, 615 anamorphosa 364 apiculata 360, 361, 873* delitescens 362, 363, 364, 873* hellwigii 219, 362, 873* limula 362, 363, 364, 873* mitchellae 361, 363, 873* myosuroides 361, 362, 364 prionodes 219, 362, 363, 364, 873* Lepicystis 477 incana 477 Lepidotis 375 cernua 375 Lepisorus 467 Leptochilus alienus 154 bernoullii 154 cladorrhizans 156 donnell-smithii 154 hemiotis 155 liebmannii 155 mexicanus 157 pergamentaceus 156 serratifolius 157 tuerckheimii 154 umbrosus 157 Leptogramma 667 totta 667 Leptopteris 434 Lindsaea 364, 603 arcuata 365, 366, 874* curvans 365 falcata 366 galeottii 365 imbricata 366 klotzschiana 365, 875* lancea 366, 874* var. arcuata 365 var. falcata 366 var. lancea 364 var. quadrangularis 366 microphylla 367 multifrondosa 367

INDEX TO SCIENTIFIC NAMES Lindsaea (Continued ) panamensis 613 portoricensis 367, 368 quadrangularis 366 subsp. antillensis 367 subsp. mexiae 367, 875* subsp. subalata 367, 874* schlechtendalii 426 stricta 367, 368, 874* f. moritziana 367 trapeziformis 364 var. arcuata 365 Lindsaeaceae 5 Lindsayopsis divaricata 426 Litobrochia 533 ampla 533 galeottii 542 grandifolia 537 grandis 534 hemipteris 539 incisa 328 kunzeana 534 mexicana 541 orizabae 539 setifera 539 tripartita 542 Llavea 5, 368 cordifolia 368, 876* Lomagramma 153 Lomaria 141 sect. Lomaridium 141 sect. Plagiogyria 466 acrodonta 151 arguta 466 danaeacea 151 deflexa 145 divergens 143 ensiformis 144 euphlebia 466 exaltata 143 falciformis 145 fragilis 145 ghiesbreghtii 152 juglandifolia 370 longifolia 151 mexicana 145 nuda 141 onocleoides 145 pectinata 466 plumieri 141 polypodioides 145 schiedeana 151 spectabilis 151 stolonifera 152 varians 151 Lomaridium 141 plumieri 141 semicordatum 466 Lomariocycas 141 Lomariopsidaceae 5, 369

Lomariopsis 282, 368 cochinchinensis 368 fendleri 370 japurensis 368 latiuscula 370 maxonii 370 mexicana 369, 877* recurvata 369, 370, 877* sorbifolium 370 vestita 369, 370, 877* Lonchitis 5, 328, 370 hirsuta 370, 371, 878* pedata 540 repens 352 tenuifolia 348 Lophidium 548 elegans 548 latifolium 548 Lopholepis 396 piloselloides 396 Lophosoria 371, 395, 396 pruinata 371, 372 quadripinnata 371, 372, 879* Lorinseria areolata 692 Lotzea diplazioides 254 Loxogramme 372 lanceolata 372 mexicana 372, 373, 550, 880* salvinii 372 Loxoscaphe 5, 373 concinna 373 var. concinna 373 lindenii 373 thecifera 373, 881* theciferum 373 Lycopodiella 329, 374, 376 alopecuroides 374, 882* caroliniana 374, 882* var. meridionalis 374, 375 cernua 374, 375, 882* inundata 374 Lycopodiaceae 5, 551 Lycopodium 329, 374, 375 alopecuroides 374 apodum 558 arcanum 337 capillare 330 carolinianum 374 var. meridionale 374 cernuum 374, 375 clavatum 375, 376, 883* complanatum 375, 377 var. validum 376 convolutum 566 crassum 331 cuernavacense 331 cuspidatum 586 dichotomum 331 fawcettii 377 flabellatum 550

Lycopodium (Continued ) var. strictum 585 gigas 334 guatemalense 330 hippurideum 332 inundatum 374 lepidophyllum 576 linifolium 333 marginatum 579 meridionale 374 mexicanum 332 myrsinites 333 nitens 336 nudum 529 orizabae 333 pallescens 586 pithyoides 334 pringlei 334 reflexum 335 rupestre 550 sargassifolium 336 schwendeneri 337 selaginoides 550 selago 329, 330 var. recurvatum 330 serratum 335 skutchii 333 taxifolium 336 tenellum 600 thyoides 376, 377, 883* trichodendron 337 tuerckheimii 334 wilsonii 337 Lygodictyon heterodoxum 377 Lygodium 18, 48, 377 flexuosum 377, 378 heterodoxum 377, 378, 884* mexicanum 378 micans 379 polymorphum 378 scandens 376 venustum 378, 874* volubile 378, 874* Macrothelypteris 379, 622 torresiana 379, 380, 875* Marattia 380, 434 alata 380, 381 excavata 380, 381, 886* laxa 381, 886* weinmanniifolia 380, 381, 886* Marattiaceae 5, 242 Marginaria 477 ceteraccina 477 incana 501 karwinskyana 499 polypodioides 501 thyssanolepis 509 Marsilea 4, 139, 381, 462 ancylopoda 382, 383, 385, 386, 888*

INDEX TO SCIENTIFIC NAMES Marsilea (Continued ) crotophora 383, 385, 887* deflexa 383, 384, 385, 887* drummondii 382 macropoda 384, 888* mexicana 382 minuta 386 mollis 382, 384, 385, 889* mucronata 386 natans 546 polycarpa 383, 384, 385, 887* var. mexicana 385 quadrifolia 381 vestita 384, 385, 386, 889* subsp. tenuifolia 386 subsp. vestita 386 Marsileaceae 382, 462, 546 Matteuccia 428 Maxonia 153, 428, 475 Mecodium 337 abruptum 339 asplenioides 339 contortum 341 fendlerianum 341 myriocarpum 344 nigricans 344 polyanthos 345 siliquosum 345 undulatum 348 Megalastrum 358, 386 atrogriseum 387, 389, 890* lunense 387, 389, 890* pansamalense 387 pulverulentum 387, 388, 389, 890* var. heydei 388 subincisum 386, 387, 388, 389, 472, 890* villosum 386 Melpomene 361, 362, 389, 615, 617 anfractuosa 389, 616 deltata 390, 392, 393, 395, 892* firma 361, 391, 891* flabelliformis 389, 390, 391, 392, 393, 395, 401, 892* leptostoma 392, 395, 892* melanosticta 390, 392, 393, 891* moniliformis 389, 390, 392, 393, 395, 892* pilosissima 392, 393, 394, 395, 892* xiphopteroides 361, 391, 392, 394, 395, 891* zempoaltepetlensis 390, 392, 395, 891* Meniscium 622, 663 angustifolium 664 cristatum 254 falcatum 665 juergensenii 665 jungersenii 665 reticulatum 663, 665 serratum 666 Meringium fucoides 342

Mertensia bancroftii 268 var. vitellina 268 bifida 605 dichotoma 250 farinosa 269 ferruginea 605 flexuosa 250 fulva 605, 606 furcata 607 glaucescens var. mexicana 322 gleichenioides 605 laevigata 604 palmata 606 pectinata 321, 322 Metaxya 395 rostrata 395, 396, 893* Microgonium petersii 679 schaffneri 677 Microgramma 165, 396, 467, 468, 477 acatallela 397, 399 ciliata 399 lycopodioides 396, 397, 398, 894* mauritiana 397 muenchii 498 nitida 397, 894* owariensis 397 percussa 398, 894* persicariifolia 396 piloselloides 396, 398, 399, 894* reptans 397, 399, 894* tecta var. nana 399, 894* var. tecta 399 vacciniifolia 396 Microlepia caudata 545 galeottii 545 schlechtendalii 426 Microphlebodium 477, 498 muenchii 396, 477, 498 Micropolypodium 361, 363, 400, 615 basiattenuatum 400, 401, 402, 895* okuboi 400 pseudotrichomanoides 400 taenifolium 401, 402, 895* trichomanoides 401, 402, 895* Mildella 5, 176, 402, 443 fallax 402, 896* intramarginalis 402, 403, 896* var. serratifolia 403 Mohria 48, 377 intermedia 51 Monogramma linearifolia 218 rostrata 218 Myriopteris cheiloglyphis 213 fendleri 194 frigida 196 gracilis 193 intermedia 203 lindheimeri 197 marsupianthes 201 rufa 192

Myriopteris cheiloglyphis (Continued ) tomentosa 210 villosa 196 Neottopteris 373 Nephelea 45 mexicana 45 polystichoides 45 tryoniana 46 Nephrodium acrostichoides 513 caucaense 626 cinereum 627 deflexum 628 fendleri var. paucipinnatum 658 fournieri 273 grisebachii 225 guatemalense 360 karwinskyanum 275 kunthii 644 lanceolatum 226 macrophyllum var. viviparum 612 mexicanum 273 nigrovenium 229 paludosum 643 panamense 634 pilosohispidum 632 polyphyllum 520 puberulum 646 quadrangulare 643 salvinii 230 schaffneri 661 tricholepis 227 tuerckheimii 648 Nephrolepidaceae 5, 404 Nephrolepis 403 ⫻averyi 405 biserrata 403, 404, 405, 407, 898* cordifolia 405, 406, 408, 898* cv. Duffii 406 exaltata 403, 405, 406, 407, 898* cv. Bostoniensis 406 cv. Whitmanii 406 falcata cv. Furcans 404 hirsutula 405, 406, 407, 897*, 898* cv. Superba 407 intermedia 409 mollis 404 multiflora 406, 407 neglecta 408 occidentalis 409 pectinata 406, 407, 408, 409, 897* pendula 406, 408, 898* rivularis 408, 897* undulata 408, 409, 897* Neurodium 409 lanceolatum 409, 410, 899* Nevrodium 409 Niphidium 165, 410, 477, 483 americanum 410

INDEX TO SCIENTIFIC NAMES Niphidium (Continued ) crassifolium 410, 411, 900* longifolium 410 Notholaena 3, 132, 176, 187, 193, 197, 214, 411 sect. Argyrochosma 64 sect. Chrysochosma 411 affinis 413, 415, 422, 906* aliena 413, 908* angusta 413 arsenii 418 aschenborniana 413, 414, 418, 903* atropurpurea 445 aurantiaca 414, 907* aurea 186 bipinnata 414 bonariensis 186 brachycaulis 414, 415, 906* brachypus 186 brevistipes 415, 901* bryopoda 415, 419, 904* californica 416 subsp. californica 416, 904* subsp. leucophylla 416 subsp. nigrescens 416 candida 193, 416, 417, 425, 905* var. copelandii 417 var. quinquifido-palmata 424 chalcolepis 135 chiapensis 186 cinnamomea 203 cochisensis 132 copelandii 417, 420, 905* crassifolia 133 cretacea 425 delicatula 65 deltoidea 134 fendleri 65 ferruginea 186 formosa 66 galeottii 414, 417, 418, 424, 903* grayi 413, 418, 419, 420, 908* subsp. sonorensis 418, 419 greggii 416, 419, 904* hookeri 424 hyalina 418 hypoleuca 418 incana 66 integerrima 133 jacalensis 417, 419, 420, 905* jonesii 67 laevis 134 lemmonii 420, 421, 422 var. australis 420, 421, 902* var. lemmonii 420, 421, 902* var. straminea 423 leonina 421, 902* limitanea 67 subsp. mexicana 68 lumholtzii 68 meridionalis 413, 420, 421, 422, 901*

Notholaena (Continued ) nealleyi 423, 424 var. mexicana 423 neglecta 422, 904* newberryi 204 nivea 64 var. mexicana 68 pallens 69 palmeri 70 parryi 205 parvifolia 69 peninsularis 70 pilifera 71 pringlei 419 pruinosa 133 rigida 422, 901* rosei 423, 902*, 908* schaffneri 419, 423, 424, 903* var. mexicana 423 var. nealleyi 423 sinuata 135 var. cochisensis 132 var. integerrima 133 var. integra 134 var. madriensis 134 var. pinnatifida 135 var. pruinosa 133 var. robusta 135 standleyi 424, 425, 906* sulphurea 411, 425, 906* var. quinquifido-palmata 424 trichomanoides 411 weatherbiana 413, 425, 908* Odontoloma schlechtendalii 426 Odontosoria 320, 364, 373, 425, 603, 604 clavata 604 divaricata 426 guatemalensis 426, 909* schlechtendalii 320, 426, 427, 909* uncinella 425 Oleandra 427 articulata 427, 910* bradei 427 guatemalensis 427 lehmannii 427 neriiformis 427 nodosa 427 Oleandraceae 5, 427 Olfersia 153, 427, 475, 607 alata 428 caudata 476 cervina 427, 428, 475, 911* corcovadensis 427 Onoclea 428 nuda 141 Onocleopsis 428 hintonii 428, 429, 912* Onychium angustifolium 182 Ophioderma palmata 431 Ophioglossaceae 5

Ophioglossum 429 californicum 430, 914* crotalophoroides 430, 914* dudadae 433 ellipticum 431 engelmannii 430, 431, 432, 913* nudicaule 430, 431, 914* var. tenerum 431 palmatum 429, 431, 913* petiolatum 433 polyphyllum 431, 432, 914* pringlei 433 pycnostichum 433 reticulatum 432, 433, 913* scandens 377, 378 tenerum 431 vulgatum 429, 431, 433, 913* var. pycnostichum 433 Ornithopteris adiantifolia 49 hirsuta 52 mexicana 55 speciosa 60 Orthiopteris 544 ferulacea 544 Osmunda 321, 433 adiantifolia 49 asplenifolia 49 cervina 428 cicutaria 164 cinnamomea 434, 435, 915* hirsuta 52 humilis 56 lunaria 161 mexicana 435 oblongifolia 56 palmeri 435 peltata 282, 303 phyllitidis 48, 58 polypodioides 145 procera 141 regalis 433, 434, 435 var. spectabilis 435, 915* spectabilis 435 virginiana 161, 164 Osmundaceae 5, 434, 466, 528 Osmundastrum 434 cinnamomeum 434 Osmundopteris 161 Paesia 348, 529 Palhinhaea 374 cernua 374, 375 Paltonium 410 lanceolatum 410 Parablechnum 141 procerum 141 Parapolystichum 358 effusum 358, 359 Parkeria 174 pteridoides 174, 175 Parkeriaceae 5

INDEX TO SCIENTIFIC NAMES Pecluma 435, 477, 487 alfredii 436, 438, 440, 443 var. alfredii 437 var. cupreolepis 437, 438, 443, 916*, 917* var. occidentalis 437, 440, 441, 916* atra 438, 439, 916* boliviana 439 chiapensis 484 consimilis 436, 438, 439, 442, 916*, 917* var. pastazensis 439 cupreolepis 437 dispersa 438, 439, 917* divaricata 436, 439, 440, 917* eurybasis 440 ferruginea 437, 438, 440, 916* hygrometrica 440, 917* pastazensis 439 pectinata 435 plumula 435, 437, 438, 439, 441, 442, 916* ptilodon 436, 438, 439, 440, 442 var. bourgeauana 442, 916* var. caespitosa 442 var. ptilodon 442 sursumcurrens 437, 442, 917* Pellaea 64, 132, 176, 213, 443 alabamensis 181 allosuroides 181, 443 andromedifolia 444, 445, 446, 919* var. pubescens 444 angustifolia 182 var. elongata 208 arsenei 181 aspera 195 atropurpurea 443, 445, 920* brandegeei 451 calomelanos 269 candida 66 cardiomorpha 445 cordata 445 cordifolia 445, 446, 450, 918* cuneata 190 fendleri 65 flavescens 209 flexuosa 448 formosa 66 fournieri 209 greggii 419 intermedia 446, 919* var. pubescens 446 intramarginalis 403, 443 var. serratifolia 403 jonesii 67 limitanea 67 longimucronata 446 lozanii 199 lozanoi 199, 440, 443 membranacea 201 microphylla 69 mucronata 446, 920* var. californica 447

Pellaea (Continued ) notabilis 445, 447, 919* oaxacana 447, 448, 918* ornithopus 446 ovata 3, 448, 918* pallida 181 pringlei 448, 449, 919* pulchella 66 pulcherrima 192 ribae 3, 449, 452, 921* rigida 214, 443 sagittata 446, 448, 449, 450, 918* var. cordata 445 seemannii 199, 443 skinneri 209, 443 ternifolia 3, 450, 451, 452 subsp. arizonica 450, 451, 453, 921* subsp. brandegeei 451, 921* subsp. ternifolia 450, 451, 921* subsp. villosa 452 var. wrightiana 452 truncata 451, 452, 453, 920* villosa 451, 452, 921* wrightiana 452, 453, 921* var. longimucronata 446 Peltapteris 282, 303 peltata 282, 303 Pentagramma 453, 463 triangularis 453 subsp. maxonii 454, 455, 922* subsp. semipallida 453, 454 subsp. triangularis 453, 454, 455, 922* subsp. viscosa 454, 455, 922* Pessopteris 410 crassifolia 411 Phanerophlebia 271, 455, 475 auriculata 456, 923* gastonyi 456, 457, 459, 924* guatemalensis 456 haitiensis 455 juglandifolia 456, 457, 458 lindenii 459 macrosora 457, 458, 459, 924* nobilis 455, 456, 457, 458, 459, 460, 923* var. nobilis 458 var. remotispora 458, 459 pumila 459, 924* remotispora 457, 458, 459, 460, 924* umbonata 456, 460, 923* Phegopteris 379 amplificata 626 blanda 651 caespitosa 651 elongata 627 hirsuta 388 impressa 635 inaequalis 233 nicaraguensis 657 pilosula 632 platyphylla 521 refulgens 229

Phegopteris (Continued ) stenolepis 388 subincisa 388 Phlebodium 460, 461, 477 araneosum 461, 462 areolatum 461, 462, 925* astrolepis 470 aureum 461, 462 var. araneosum 461 decumanum 461, 462, 925* inaequale 500 nitidum 397 Phyllitis 72, 73 fernaldiana 120 japonica 120 subsp. americana 120 lindenii 120 scolopendrium 72, 120 var. americana 120 var. lindenii 120 Physematium canescens 687 molle 689 Pilularia 462 americana 462, 926* globulifera 462 Pityrogramma 61, 62, 453, 463 calomelanos 464, 927* chrysophylla 463 dealbata 464, 928* ebenea 464, 465, 928* guatemalensis 62 schaffneri 464 tartarea 465 triangularis 453, 463 var. maxonii 454 var. semipallida 453 var. viscosa 454 trifoliata 463, 465, 928* tripinnata 465 viscosa 454 Plagiogyria 466 aequidentata 466 arguta 466 biserrata 466 costaricensis 466, 467 euphlebia 466 fialhoi 467 maxonii 467 pectinata 466, 467, 929* semicordata 466 serrulata 467 truncata 466, 467 Plananthus myrsinites 333 Platyloma atropurpurea 445 flexuosa 448 sagittata 449 Plecosorus 511 speciosissimus 511, 523 Pleopeltis 396, 410, 467, 477, 498 angusta 467, 469 var. angusta 469, 931*

INDEX TO SCIENTIFIC NAMES Pleopeltis (Continued ) var. stenoloma 469, 931* astrolepis 470, 472, 473, 930* conzattii 471, 473, 930* crassinervata 471, 472, 473, 930* crassinervata ⫻ fallax 472 elongata 470 erythrolepis 468, 474 fallax 469, 470, 472, 930* interjecta 475 lanceolata 470 ⫻leucospora 473 macrocarpa 473 var. interjecta 475 var. trichophora 472 macrocarpa ⫻ Polypodium polypodioides subsp. ecklonii 467 macrocarpa ⫻ Polypodium thyssanolepis 473 ⫻melanoneuron 472, 930* mexicana 471, 472, 473, 475, 930* muenchii 498 peltata 474 percussa 398 pleopeltifolia 469 polylepis 472, 473, 474 var. erythrolepis 468, 474, 475, 931* var. interjecta 475, 931* var. polylepis 474, 475, 931* polypodioides 501 var. acicularis 502 var. michauxiana 503 repanda 410 revoluta 470 riograndensis 506 ⫻sordidula 470, 472, 930* ⫻Pleopodium 467, 468 bartlettii 474 fallacissimum 468, 490 simianum 467 tricholepis 473, 937* Pleuroderris 609 michleriana 609, 615 Polybotrya 153, 428, 475, 607 aucuparia 476 caudata 476, 932* cervina 428 osmundacea 475, 476 polybotryoides 476, 932* scandens 476 villosula 476 Polypodiaceae 5, 175, 372 subfam. Ceratopteroideae 175 subfam. Pleopeltoideae 5 subfam. Polypodioideae 5 tribe Ceratopterideae 175 Polypodium 3, 5, 165, 239, 396, 435, 441, 461, 467, 468, 469, 472, 474, 475, 477 sect. Phlebodium 460 adelphum 509 affine 497

Polypodium (Continued ) alansmithii 468, 480, 481, 938* alavae 481, 945* alfredii 436, 437, 443 alsophiloides 223 americanum 410 amphostenon 166 anfractuosum 616 angustifolium 167 var. amphostenon 166 var. ensifolium 168 angustifrons 281 angustum 469 anisomeron 499 var. abortivum 500 var. elongatum 500 var. genuinum 500 var. molestum 499 var. pectinatum 500 apiculatum 360, 361 araneosum 461 arboreum 234 arcanum 481, 482, 486, 488 var. arcanum 482, 483, 940* var. bakeri 482, 947* var. septentrionale 482, 483, 947* areolatum 461 aristatum 63 aromaticum 391 arthropodium 511 asplenifolium 615, 617 astrolepis 470 atrum 438 var. areolatum 461 balbisii 625 bartlettii 474, 930* basiattenuatum 400 bernoullii 486 biauritum 496 biserratum 509 blandulum 399 blepharodes 401 bolivianum 439, 440 brachyodus 668 bradeorum 477, 483, 508, 933*, 934*, 943* californicum 483, 484, 940* callolepis 497 cancellatum 495 caudatum 171 ceteraccinum 477 cheilostictum 499 chiapense 435, 484, 487, 946* chnoodes 487 christensenii 484, 485, 945* ciliatum 399 collinsii 468, 485, 937* colpodes 482, 483, 485, 486, 500, 940* colysoides 483 concinnum 627 confluens 486, 499 consimile 438

Polypodium (Continued ) var. bourgaeanum 442 var. bourgeauanum 442 conterminans 486, 946* conzattii 471 cordifolium 405, 610 costatum 168 crassifolium 410, 411 crassinervatum 471 crenatum 648, 658 var. ghiesbreghtii 651 cryptocarpon 468, 486, 487, 492, 499, 936* cultratum 617 cupreolepis 437 curvatum 172 cyathicola 442, 443 decumanum 462 delicatulum 402, 618 delitescens 362 deltoideum 481, 668 dentatum 642 denticulatum 63 diaphanum 241 dichotomum 250 dilatatum 359 diplotrichum 435, 484, 487, 937* dispersum 439 dissimile 487, 488, 941* divaricatum 439, 440 divergens 359 domingense 652 donnell-smithii 505 duale 219 dulce 488, 934* eatonii 482, 488, 947* echinolepis 488, 489, 504, 511, 937* ecostatum 316 effusum 358, 359 ehrenbergianum 504 ellipsoideum 493 elongatum 470 ensifolium 168 eperopeutes 489, 941* erythrolepis 474 exaltatum 406 excelsum 224 falcaria 477, 489, 490, 940* fallacissimum 468, 475, 490, 507, 944* fallax 468, 472 fasciale 170 faucium 657 feei 500 ferrugineum 440 filix-femina 135 filix-mas 269, 274 firmulum 509 firmum 391 fissidens 490, 491, 945* flabelliforme 391 flavopunctatum 607

INDEX TO SCIENTIFIC NAMES Polypodium (Continued ) flexuosum 174 fournieri 504 fragile 240 fraternum 491, 504, 511, 939* fraxinifolium 491, 942* friedrichsthalianum 496 fulvum 411 furfuraceum 468, 477, 486, 487, 492, 499, 511, 936*, 943* var. coronulatum 492 var. rufum 486, 492 fuscopetiolatum 485 galeottii 388, 472, 504 ghiesbreghtii 488 gibbosum 402 glaberulum 504 glanduliferum 273 glandulosum 668 glaucinium 461 glaucum 268, 371, 372 globuliferum 248 griseum 495 guadalupense 652 guatemalense 483, 500 guilleminianum 509 guttatum 468, 475, 490, 492, 946* haenkei 510 hahnii 509 harrisii 316 hartwegianum 435, 487, 493, 495, 939* hemsleyanum 225 hesperium 484, 493, 945* hirsutissimum var. sericeum 507 hirsutulum 406 hispidulum 486, 494, 937* hygrometricum 440 imbricatum 653 inaequale 500 incanum 501 var. fimbriatum 510 var. umbrosum 486 induens 616 insigne 489 isomeron 506 jamesonii 364 juglandifolium 457 jungermannioides 173 karstenianum 388 karwinskyanum 499 lanceolatum 470 var. crassinervatum 471 var. elongatum 470 var. trichophorum 472 latipes 490 laxifrons 617 lepidotrichum 468, 494, 495, 935* leptophyllum 61, 62 leptostomum 392 lesourdianum 506 leucatomas 462

Polypodium (Continued ) leucosticton 505 liebmannii 493, 495, 946* limulum 363 lindenianum 468, 495, 943* litigiosum 634 lobatum 610 lonchitis 511 longepinnulatum 487, 488, 489, 496, 511, 939*, 946* longicaudatum 608 loriceum 490 var. plesiosorum 499 lowei 500 lycopodioides 396, 397 madrense 468, 493, 496, 497, 499, 938* margallii 492 martensii 497, 939* maxonii 509 medullare 603 melanostictum 392 meniscioides 655 mickelii 485 microdontum 238 microgrammoides 3, 497, 498, 942* minusculum 317 mitchellae 363 mollissimum 618 moniliforme 389, 393 var. major 393 montigenum 481 muenchii 3, 396, 468, 477, 498, 933* munchii 498 muricatum 519 myosuroides 364 nesioticum 316 nitidum 397 nivosum 492 obliteratum 657 oligocarpum 632 olivaceum 504 var. elatum 504 oulolepis 496 pachyrhizon 500 palmeri 397 parietinum 316 patens 646 pectinatum 435 var. caespitosum 442 peltatum 474 var. interjectum 475 penna-marina 149 percussum 396, 397, 468 persicariifolium 396 petiolatum 491 phyllitidis 169 piloselloides 396, 397 pilosissimum 394 platylepis 468, 498, 499, 936* plebeium 468, 477, 481, 497, 499, 938* plectolepis 489

Polypodium (Continued ) var. incisum 489 pleolepis 508 plesiosorum 477, 482, 484, 494, 499, 500, 504, 505, 507, 508, 940*, 947* var. bakeri 482 pleurosorum 500, 501, 940*, 941* plumula 441 polylepis 473, 483 polypodioides 477, 499, 501, 502 var. aciculare 468, 502, 933* var. knoblochii 468, 502, 503, 933* var. michauxianum 468, 502, 503, 933* var. polypodioides 502, 933* praetermissum 500, 503, 504, 942* productum 392 prolongilobum 493 pruinatum 174, 371, 372 pruinosum 174 pseudoaureum 462 pseudotrichomanoides 400 ptilodon var. caespitosum 442 puberulum 481, 485, 489, 504, 937*, 939* pubescens 493, 504 pulchrum 441 var. minus 441 pulverulentum 388 purpusii 510 pyrrholepis 468, 504, 935* quadripinnatum 371, 372 refulgens 229 remotum 468, 505, 944* repens 165, 170 repletum 317 reptans 659 resiniferum 633 reticulatum 663, 665 revertens 491 rhachipterygium 505, 941* rhagadiolepis 510 rhodopleuron 494, 500, 506, 940* rigens 394 rigescens 391 riograndense 468, 490, 506, 507, 944* rivulare 408 rosei 468, 495, 505, 507, 508, 944* rossii 500 rostratum 395, 396 rosulatum 316 rude 635 rzedowskianum 468, 507, 508, 938* salicifolium 664 sanctae-rosae 468, 485, 507, 508, 935* schaffneri 509 scouleri 484, 508, 509, 947* semicordatum 239, 240 semihirsutum 619 var. fuscosetosum 619 senile 620 serpentinum 170 serratum 511

INDEX TO SCIENTIFIC NAMES Polypodium (Continued ) serrulatum 219 sessilifolium 489 skinneri 486 skutchii 364 sloanei 231 sordidulum 470, 472 sororium 488 sporadocarpum 462 squamatum 495, 505, 508, 510 stenoloma 505 subflabelliforme 620 subincisum 388 submarginale 232 subpetiolatum 477, 481, 485, 489, 509, 937*, 939* subtile 621 surcurrens 486 sursumcurrens 442, 443 tablazianum 436, 437 taenifolium 401 tectum 399 tenuipes 171 teresae 509 tetragonum 662 thomsonii 637 thyssanolepis 468, 473, 490, 501, 506, 509, 510, 936*, 944* var. riograndense 506 tottum 643, 667 trichomanoides 401, 402 trichosorum 317 trifoliatum 609 triseriale 492, 501, 510, 511, 941* tuerckheimii 472 vacciniifolium 396 van-heurckii 500 verapax 495 villagranii 468, 511, 943* villosum 386 vulgare 477, 484 subsp. columbianum 493 var. columbianum 493 var. hesperium 493 weatherbyanum 171 xalapense 171 xiphopteroides 394 yarumalense 316 Polystichopsis 63, 455 Polystichum 19, 63, 271, 455, 475, 511 sect. Parapolystichum 358 acrostichoides 512, 513, 519, 949* aculeatum var. alfaroi 513 var. distans 525 var. gracile 525 var. incisum 525 alfarii 514 alfaroi 514, 520, 525, 950* alfaroi ⫻ platyphyllum 514 bicolor 521 concinnum 524

Polystichum (Continued ) confluens 522 cystopteroides 273 denticulatum 63 distans 514, 515, 516, 517, 518, 522, 523, 524, 525, 952* drepanoides 517, 518 echinatum 519 erythrosorum 515, 516, 517, 521, 523, 951* fournieri 516, 517, 520, 521, 525, 951* furfuraceum 516, 517, 950* glandulosum 19 grande 517 hartwegii 512, 514, 515, 516, 517, 518, 520, 521, 523, 525, 951*, 952* hartwigii 517 lanosum 522 lonchitis 271, 511 mickelii 512, 518, 525, 951*, 952* montevidense 518 muelleri 516 muenchii 512, 518, 949* munitum 512, 514, 519, 949* subsp. solitarium 519 muricatum 519, 520, 521, 525, 950* orbiculatum 512, 518, 520, 950* ordinatum 515, 517, 518, 520, 521, 525, 951* pallidum 514 platyphyllum 512, 518, 520, 521, 522, 952* polyphyllum 520 polystichiforme 522 rachichlaena 515, 516, 517, 522, 523, 524, 525, 948* schizophyllum 3, 512, 523, 948* sellowianum 512 semicordatum 240 smithii 524, 525 solitarium 519 speciosissimum 511, 512, 518, 520, 523, 524, 949* talamancanum 524 torresianum 379 trejoi 517 turrialbae 517, 524, 525, 949* Polytaenium 61, 525, 544, 549, 550 anetioides 527 brasilianum 526 cajenense 526, 527, 953* chlorosporum 526, 527, 953* ensiforme 550 feei 527, 953* guayanense 527 lanceolatum 525, 527, 528 lineatum 48, 525, 527, 528, 953* Pomatophytum pocillatum 196 Pseudocolysis 477 bradeorum 477, 483 Pseudolycopodiella 374 caroliniana 374 meridionalis 374 Pseudophegopteris 379

Psilogramme chiapensis 321 Psilotaceae 5, 434, 528 Psilotum 528 complanatum 528, 529, 954* nudum 528, 529, 954* triquetrum 528, 529 Pteridaceae 5, 175, 544 subfam. Adiantoideae 21 subfam. Cheilanthoideae 5 subfam. Pteridoideae 5 subfam. Taenitoideae 5 Pteridanetium citrifolium 60, 61 Pteridium 44, 328, 348, 529 aquilinum 529, 530 subsp. caudatum 532 var. arachnoideum 532 var. caudatum 532 var. feei 529, 530, 531, 955* var. latiusculum 529, 531, 532, 955* var. pseudocaudatum 533 var. pubescens 530, 531, 532, 533, 955* arachnoideum 529, 532, 533, 955* caudatum 529, 530, 532, 533, 955* var. arachnoideum 532 feei 530 Pteris 5, 17, 533 sect. Histiopteris 327 aequalis 537 alabamensis 181 altissima 534, 535, 957* ampla 533 andromedifolia 444 angustifolia 47 apicalis 539 aquilina 529, 530 var. arachnoidea 532 var. caudata 532 var. mexicana 532 var. pubescens 531 arachnoidea 532 arborescens 541 atropurpurea 443, 445 aurantiaca 184 aurea 186 biaurita 535, 542, 956* cartilaginea 214 caudata 532 var. mexicana 532 chiapensis 535, 536, 961* concolor 269 cordata 445 cretica 536, 956* erosa 536, 539, 540, 957* fallax 403 farinosa 192 feei 530 flexuosa 192, 448 galeottii 542 grandifolia 533, 537, 956* hemipteris 539

INDEX TO SCIENTIFIC NAMES Pteris (Continued ) incisa 328 inflexa 540 interrupta 643 intramarginalis 402, 403 kunzeana 534 laciniata 371 lanceolata 409, 410 latiuscula 531 lendigera 196 lineata 683, 686 longifolia 533, 537, 538, 543, 956*, 961* lutea 134 mexicana 541 mollis 538 muricata 538, 543, 956* muricella 538, 543, 959* nemoralis var. major 542 nivea 64 notholaenoides 204 orizabae 535, 537, 539, 540, 541, 957* ovata 448 palmata 270 paucinervata 539, 540, 542, 959* pedata 540 plumula 542 podophylla 537, 539, 540, 957* polita 539 propinqua 537, 539, 540, 541, 960* protea 534 pulchra 536, 541, 956* pungens 540, 541, 542, 949* quadriaurita 535, 542, 956* rigida 213, 214 sagittata 449 sulphurea 411, 425 ternifolia 450 tremula 543, 960* trichomanoides 411 trifoliata 536 tripartita 542, 958* triphylla 536 vespertilionis 327 vittata 533, 538, 543, 961* Pteropsis angustifolia 469 Radiovittaria 47, 61, 322, 544, 685 remota 47, 544 stipitata 544, 962* Regnellidium 462 Rhipidopteris 282 peltata 303 Rumohra 358, 386, 512 adiantiformis 512 denticulata 63 Saccoloma 5, 244, 544 elegans 544, 545, 963* ferulaceum 544 inaequale 545, 546, 963*

Sagenia lobata 610 mexicana 613 Salvinia 139, 462, 546 auriculata 546, 547, 964* minima 547, 964* molesta 546 natans 546, 547 rotundifolia 546, 547 Salviniaceae 382 Sceptridium 161 decompositum 162 obliquum 161 pusillum 163 schaffneri 163 Schaffneria 5, 73, 123, 547 nigripes 547, 548, 965* Schizaea 18, 48, 377, 548 dichotoma 548 elegans 548, 549, 966* germanii 18 pennula 18 poeppigiana 549, 966* Scoliosorus 47, 525, 544, 549 ensiformis 372, 373, 549, 550, 967* Scolopendrium 72 lindenii 120 nigripes 548 vulgare 72 Selaginella 3, 550 series Articulatae 551, 595, 601 series Circinatae 588 series Eremophilae 562, 602 series Sartorii 602 subg. Heterostachys 550, 551 subg. Selaginella 551 subg. Stachygynandrum 550, 551 subg. Tetragonostachys 550, 551 acutifolia 552, 558, 986* albonitens 600 ambigua 570 var. fallax 570 anisoclada 588 apoda 558, 987* applanata 577 apus 558 arizonica 551, 559, 568, 587, 976* arsenei 551, 559, 566, 978* arsiclada 559, 560, 577, 986* asprella 551, 560, 975* barnebyana 560, 561, 565, 575, 985* basipilosa 551, 561, 562, 568, 576, 968* bernoullii 551, 562, 563, 574, 578, 589, 974* bigelovii 551, 563, 602, 977* binervis 589 breedlovei 563, 564, 987* bulbifera 589 carioi 564, 986* carnerosana 551, 564, 565, 568, 975* cavifolia 589 chiapensis 561, 565, 568, 572, 971* chrismarii 592

Selaginella (Continued ) var. karwinskyana 592 var. neeana 592 cinerascens 551, 565, 566, 975* cladorrhizans 600 contigua 565 convoluta 552, 566, 577, 981* corrugis 565, 566, 567, 571, 575, 599, 970* coryi 602 cruegeri 581 crugeri 581 cuneata 567, 576, 585, 969* cuspidata 586 var. elongata 586 delicatissima 567, 568, 594, 970* denudata 565, 575 diffusa 597 disticha 588 eclipes 558 eremophila 551, 568, 976* eurynota 551, 568, 569, 595, 596, 599, 979* extensa 569, 551, 978* faucium 571 fendleri 601 finitima 551, 569, 570, 578, 582, 590, 989* flabellata 550 flacca 589 flagellata 551, 564, 570, 575, 590, 971* flexuosa 561, 565, 567, 568, 571, 575, 593, 599, 970* galeottii 579, 597 guatemalensis 561, 565, 568, 571, 572, 575, 985* gypsophila 552, 572, 573, 588, 981* hansenii 602 harrisii 552, 573, 586, 588, 591, 969*, 984* heterostachys 550 hintonii 593 hirtifolia 573, 574, 577, 971*, 973* hoffmannii 574, 575, 578, 969*, 971* homaliae 577 huehuetenangensis 560, 561, 565, 572, 575, 985* humilis 600 idiospora 565, 572, 575 illecebrosa 575, 576, 581, 583, 974* incurvata 597 jungermannioides 577 krugii 565 kunzeana 599 landii 551, 562, 568, 576, 595, 976* lehmannii 571 lepidophylla 552, 576, 577, 584, 592, 968* lindenii 574, 577, 985* lineolata 551, 570, 577, 578, 580, 582, 590, 988* longispicata 566 ludoviciana 558 macrathera 551, 578, 978* macrostachya 565 maracasensis 600

INDEX TO SCIENTIFIC NAMES Selaginella (Continued ) marginata 551, 579, 979* martensii 568, 572, 579, 580, 585, 972* mendocae 565 mickelii 576, 580, 581, 970* microdendron 573 microphylla 564 millspaughii 573 minima 551, 581, 597, 973* miradorensis 597 mixteca 551, 581, 582, 594, 973* mollis 581, 600 moritziana 551, 582, 989* var. moritziana 582, 589 mosorongensis 582, 583, 585, 970* mutica 551, 583, 980* var. mutica 583 var. limitanea 583 nicaraguensis 595 nivea 566 nothohybrida 552, 577, 583, 584, 592, 982* novoleonensis 552, 577, 584, 981* oaxacana 567, 576, 583, 585, 969* oregana 569 orizabensis 563, 564, 585, 987* pallescens 552, 558, 573, 586, 588, 969* var. acutifolia 558 pansamalensis 571 parishii 551, 568, 576, 586, 976* pauciciliata 595 peruviana 551, 559, 568, 587, 975* pilifera 552, 573, 587, 588, 982* var. pringlei 587 polyptera 552, 573, 586, 588, 591, 984* popayanensis 551, 574, 588, 589, 973* porphyrospora 551, 562, 563, 574, 578, 589, 971* porrecta 593 pringlei 587 prolifera 551, 570, 578, 582, 590, 972* pulcherrima 552, 573, 586, 588, 590, 591, 983* reflexa 560, 564, 591, 986* revoluta 573, 574 rhizophora 570, 590 ribae 552, 577, 591, 592, 980* rupestris 550 f. peruviana 587 f. wrightii 602 var. fendleri 601 var. sartorii 593 rupincola 551, 563, 592, 602, 968* rzedowskii 592, 593, 982* saccharata 567 sartorii 551, 593, 598, 977* schaffneri 593, 594, 980* schiedeana 560, 564, 591, 594, 973* schizobasis 551, 569, 594, 595, 596, 599, 979* schultesii 577 selaginoides 550 sellowii 551, 559, 593, 595, 977*

Selaginella (Continued ) sertata 551, 569, 595, 596, 599, 979* sheldonii 587 silvestris 551, 596, 599, 601, 972* simplex 551, 581, 596, 597, 988* sintenisii 600 solmsii 579 spinosa 550 stellata 551, 579, 596, 597, 599, 972* stenophylla 551, 578, 580, 597, 598, 987* steyermarkii 551, 598, 977* subcaulescens 600 subrugosa 565, 567, 571, 575, 598, 599, 985* tarapotensis 571 tarda 551, 596, 599, 974* tenella 551, 575, 577, 600, 989* tropidophora 581 truncata 577 tuberosa 551, 582, 590, 600, 601, 988* underwoodii 551, 601, 978* var. dolichotricha 601 viridissima 551, 601, 602, 977* wendtii 575 wrightii 551, 576, 595, 602, 968* Selaginellaceae 4, 551 Selenodesmium rigidum 682 Selliguea mexicana 372 Sinephropteris 547 Sinopteris 71 Sphaerocionium 337 ciliatum 342 crispum 340 elegantulum 346 hirsutum 337. 342 lanatum 343 microcarpum 344 nigricans 344 pulchellum 345 schiedeanum 340 sieberi 346 Sphaeropteris 268, 603 horrida 603, 990* integra 268 marginalis 239 medullaris 603 myosuroides 238 Sphenomeris 426, 603 clavata 3, 603, 604, 652, 991* Stachygynandrum 550 flabellatum 550 Stegnogramma 622, 667 aspidioides 667 pilosa 667 Steiropteris 622, 668 glandulosa 668 Stenochlaena palustris 370 pittieri 370 recurvata 369 vestita 370 Stenogramma ehrenbergiana 158 Stenoloma clavatum 604 schlechtendalii 426

Sticherus 250, 268, 269, 321, 604 bifidus 605, 606, 607, 992* brevipubis 605, 606, 607, 993* brevitomentosus 606 farinosus 269 ferrugineus 605 fulvus 606 furcatus 606 jamaicensis 607 laevigatus 604 palmatus 605, 606, 607, 993* truncatus 604 underwoodianus 606, 607, 993* Stigmatopteris 455, 475, 607 chimalapensis 608, 609 longicaudata 608, 609, 994* rotundata 607 sordida 608, 609, 994* Stromatopteris 321, 604 Struthiopteris 141 ensiformis 144 exaltata 143 falciformis 145 maxonii 147 spicant 141 stolonifera 152 Taenitis angustifolia 47 lanceolata 410 Tectaria 219, 609 buchtienii 615 ⫻bulbifera 613 cicutaria 613 cinnamomea 273 dilacerata 613 fimbriata 610, 995* heracleifolia 610, 612, 613, 614, 615, 996* var. maxima 611 var. trichodes 615 incisa 611, 612, 614 f. incisa 612, 995* f. vivipara 612, 613, 996* subsp. transiens 614 incisa ⫻ panamensis 615 latifolia 613 lobata 610 martinicensis 612 var. vivipara 612 mexicana 611, 613, 996* var. pilosula 613 minima 610 nicotianifolia 609 panamensis 609, 613, 614, 996* pilosa 615 plantaginea 609 pubens 615 ramkissoonii 614 rheosora 609 rivalis 609, 613 transiens 611, 614, 995* trichodes 611, 615, 995* trifoliata 609

INDEX TO SCIENTIFIC NAMES Tectaria (Continued ) vivipara 612, 614 Tectariaceae 609 Terpsichore 5, 360, 389, 400, 615 achilleifolia 400 alfarii 619 anfractuosa 616, 617, 620, 998* asplenifolia 615, 616, 617, 618, 999* chrysleri 617 cretata 621 cultrata 360, 363, 616, 617, 618, 619, 620, 999* delicatula 616, 618, 997* elastica 615 flexuosa 174 heteromorpha 621 jamesonioides 618 lanigera 618, 620 lehmanniana 617 longisetosa 400 mollissima 616, 618, 619, 997* saxicola 616 saxicolum 616 semihirsuta 616, 617, 619, 620, 997* senilis 616, 618, 619, 620, 998* spathulata 616, 620, 621, 999* subscabra 618 subtilis 616, 621, 998* taxifolia 616, 620 Thelypteridaceae 379, 622 Thelypteris 219, 379, 607, 621 sect. Apelta 630 sect. Goniopteris 648 sect. Lepidoneuron 630 sect. Macrothelypteris 379 sect. Phacelothrix 630 sect. Uncinella 630 subg. Amauropelta 623 subg. Cyclosorus 638 subg. Cyclosorus sect. Meniscium 663 subg. Goniopteris 648 subg. Macrothelypteris 379 subg. Meniscium 663 subg. Stegnogramma 667 subg. Steiropteris 668 subg. Thelypteris 668 albicaulis 639, 640, 647, 1006* angustifolia 664, 666, 667, 1013* asterothrix 649 atrovirens 624, 625, 1004* augescens 3, 640, 654, 1007* balbisii 625, 629, 634, 637, 1003* biolleyi 650, 651, 653, 1008* blanda 651, 662, 1008* blepharis 641, 1007* brachyodus 668 burksiorum 667, 668 caucaensis 625, 1004* cheilanthoides 626, 637 var. cheilanthoides 626, 627, 1000* var. mucosa 626, 627, 1000* cinerea 627, 628, 637, 1001*

Thelypteris (Continued ) columbiana 632 concinna 627, 628, 638, 1002* var. elongata 627 confluens 669 conspersa 644 cretacea 641, 642, 1007* cruciata 635 darwinii 652, 660 decussata 668 deflexa 628, 630, 1002* deltoidea 668 dentata 640, 642, 643, 1006* diplazioides 628 falcata 665, 666, 667, 1013* feei 646 ghiesbreghtii 649, 651, 653, 655, 659, 1011* glandulosa 668 var. brachyodus 668, 1016* var. glandulosa 668 var. longipilosa 668 gongylodes 643 grandis 640 var. pallescens 640 guadalupensis 652, 654, 1009* hatchii 651, 652, 653, 1008* hispidula 640, 642, 643, 644, 1006* var. hispidula 643 var. inconstans 643 var. versicolor 643 hondurensis 649, 653, 655, 1011* imbricata 653, 654, 656, 657, 658, 663, 1008* interrupta 638, 643, 644, 1007* kunthii 640, 641, 644, 645, 1006* lanosa 644, 645, 1007* leptocladia 654, 1012* limbata 623 lindigii 628 linkiana 625, 628, 1002* loreae 629, 630, 1005* martinezii 654, 655, 658, 659, 1010* melanochlaena 630, 636, 638, 1003* meniscioides 652, 653, 655, 661, 1011* var. meniscioides 656 var. ternata 656 minor 654, 655, 656, 657, 659, 1009* mollis 651 mortonii 629, 630, 637, 1001* muenchii 656, 657, 1009* munchii 656 nephrodioides 651 nicaraguensis 654, 655, 656, 657, 659, 1009* normalis 644 nubigena 631, 1003* oaxacana 631, 632, 637, 1003* obliterata 655, 657, 658, 663, 1009* oligocarpa 632, 633, 1002* ovata 639, 640, 644, 646 var. lindheimeri 645, 646, 648, 1006* var. ovata 646

Thelypteris (Continued ) pachyrhachis 632 palustris 621, 622, 669 var. palustris 669 var. pubescens 669, 1017* panamensis 634 patens 642, 644, 645, 646, 1006* var. dissimilis 646 var. patens 646 var. smithiana 646 paucipinnata 658, 1008* pellita 649 pilosa 632, 667, 1014*, 1015* var. alabamensis 667 var. major 667 var. pilosa 667 pilosohispida 632, 633, 635, 1002* pilosula 632, 633, 1003* poiteana 648, 652, 655, 658, 659, 1011* polyphylla 651 pozoi 667 praetermissa 655, 659, 1010* puberula 522, 639, 640, 644, 646, 647, 648 var. puberula 646, 647, 1006* var. sonorensis 647, 648, 1007* quadrangularis 643 reptans 652, 659, 1010* resiliens 657, 660, 1010* resinifera 625, 629, 631, 633, 634, 636, 637 var. proxima 634, 635, 1004* var. resinifera 634, 1003* reticulata 663, 665, 666, 1013* retrorsa 633 rhachiflexuosa 660, 661, 1011* rudis 626, 632, 633, 635, 1002* salicifolia 664 sancta 630, 635, 636, 637, 1001* sanctiformis 627 scalaris 630, 636, 638, 1003* schaffneri 658, 661, 662, 1011* schippii 649 serrata 665, 666, 1013* skinneri 649, 652 sprengelii 625 standleyi 666, 667, 1013* stegnogramma 667 stolzeana 662, 1010* struthiopteroides 625, 629, 634, 637, 1001* tablana 637, 1001* tetragona 653, 654, 655, 657, 658, 662, 663, 1008* thelypteroides 621 thomsonii 631, 637, 638, 1001* toganetra 658, 663, 1008* torresiana 379 totta 643 tuerckheimii 647, 648, 1007* uliginosa 379 venturae 638, 1004* Thyrsopteris 395 Todea 434 Trachypteris 269

INDEX TO SCIENTIFIC NAMES Trichipteris 234 bicrenata 235 corcovadensis 234 costaricensis 236 excelsa 234 mexicana 237 microdonta 238 scabriuscula 235, 236 schiedeana 239 Trichomanes 669 sect. Didymoglossum 673, 679 angustatum 671, 672, 673, 1019* angustifrons 672, 1020* apodum 677 asplenioides 339 boschianum 682 bucinatum 672, 682, 1024* capillaceum 671, 672, 673, 1023* cavifolium 681 ciliatum 342 collariatum 673, 682, 1018* crispum 669, 674, 676, 683, 1022* curtii 674, 675, 1020* diaphanum 677 diversifrons 675, 1019* ekmanii 675, 676, 1020* elegans 675 flaccidum 244 fragile 342 fucoides 342 galeottii 674, 675, 676, 677, 1019* godmanii 675, 676, 1024* hirsutum 342 holopterum 676, 677, 1021* hymenoides 669, 673, 677, 1024* hymenophylloides 677, 681, 1023* hypnoides 673 krausii 677, 678, 1024* kunzeanum 681 leptophyllum 677 lucens 678, 1022* martinezii 673 membranaceum 678, 1019* mexicanum 681 muscoides 669, 677 ovale 673, 674, 675, 679, 680, 1019* petersii 673, 675, 679, 680, 1020* pinnatum 680, 1022* plumosum 683 polyanthos 337, 345 polypodioides 680, 681, 1022* pyxidiferum 677, 681, 1023* radicans 674, 681, 682, 1018*, 1021* reptans 673, 678, 682, 1021* rigidum 682, 683, 1023* scandens 681 schaffneri 677, 679 schiedeanum 673 sinuosum 680 tenellum 673 tenerum 671

Trichomanes (Continued ) tenerum 671 trichodes 673 trichoideum 673 tuerckheimii 683, 1021* tunbrigense 347 undulatum 348 Trichopteris 234 Trichosorus 371 densus 372 frigidus 372 glaucescens 372 pruinatus 371 Trismeria 463, 465 aurea 463 trifoliata 465 Urostachys 329 capillaris 330 crassus 331 cuernavacensis 331 cuernavensis 331 dichotomus 331 gigas 334 guatemalensis 330 hippurideus 332 linifolius 333 myrsinites 333 orizabae 333 pithyoides 334 pringlei 334 reflexus 335 serratus 335 var. japonica-neotropicus 336 taxifolius 336 trichodendron 337 tuerckheimii 334 wilsonii 337 Vandenboschia angustata 671 capillacea 673 hymenophylloides 677 martinezii 673 pyxidifera 681 radicans 681 tenera 671 Vittaria 47, 525, 526, 544, 683 subg. Radiovittaria 544 angustifolia 47 appalachiana 684 bradeorum 684, 685, 686, 1025* costata 47 deppeana 686 dimorpha 685 filifolia 685 flavicosta 685, 1025* graminifolia 685, 686, 1025* isoetifolia 684 lanceolata 528 lineata 528, 683, 685, 686, 1025* remota 544

Vittaria (Continued ) stipitata 544 Vittariaceae 5, 21, 544 Wolffia 139 Wolffiella 139 Woodsia 3, 686 canescens 687, 688, 1026* cochisensis 688, 1027*, 1028* cystopteroides 688, 689, 1027* fragilis 689 guatemalensis 689, 690 ilvensis 686 mexicana 689, 690, 1028* mollis 687, 688, 689, 690, 691, 1026* montevidensis 690, 1026* neomexicana 688, 690, 1028* obtusa var. glandulosa 691 var. plummerae 691 oregana ⫻ phillipsii 690 phillipsii 688, 689, 691, 692, 1028* plummerae 689, 691, 692, 1027* pusilla 689 var. glandulosa 691 var. mexicana 689 Woodsiaceae 19 Woodwardia 141, 692 areolata 692 chamissoi 692 fimbriata 692, 693, 695, 1030* japonica 693 kempii 693 martinezii 693, 694, 695, 1030* semicordata 694, 695, 1030* spinulosa 693, 694, 695, 1029* radicans 692 virginica 692 Xiphopteris 217, 361 basiattenuata 400 blepharodes 401 delitescens 362 flabelliformis 391 jamesonii 364 limula 363 mitchellae 363 myosuroides 364 rigescens 391 serrulata 217, 219 skutchii 364 taenifolia 401 trichomanoides 401 Zaluzianskia 381 ancylopoda 382 deflexa 383 macropus 384 marsiloides 381 mexicana 382 polycarpa 385 vestita 386 Zygophlebia 172

John T. Mickel and Alan R. Smith Dr. John T. Mickel (left) was born in Hudson, Ohio, in 1934. He received his B.A. from Oberlin College in 1956 and then went to the University of Michigan, where he earned his M.A. in 1958 and Ph.D. in 1961 as a student of Dr. Warren H. Wagner, Jr. Dr. Mickel taught at Iowa State University from 1961 to 1969, with one year off as Visiting Professor at the University of Costa Rica, leading a course in tropical pteridophytes for the Organization for Tropical Studies (OTS) in the summer of 1967, and as Senior Visiting Research Associate at the Smithsonian Institution. In 1969 he was appointed Curator of Ferns at the New York Botanical Garden, where he has carried on research in tropical American ferns with emphasis on Mexico, Hispaniola, and Trinidad, and the genera Anemia and Elaphoglossum. His published books include How to Know the Ferns and Fern Allies (the first book since 1906 to include all species of North America), Trinidad Pteridophytes, Pteridophyte Flora of Oaxaca, Mexico (with Joseph Beitel), and Pteridophytes of Nueva Galicia (western Mexico). Dr. Mickel founded the New York Chapter of the American Fern Society in 1973 and has served as its secretary-treasurer to the present. He has been president of the American Fern Society and established its bulletin, Fiddlehead Forum, in 1974, for which he was editor for 20 years. In addition to his scientific studies, Dr. Mickel has a strong interest in fern cultivation. His work on testing of cold-hardiness has led to the introduction of many temperate fern species to cultivation and resulted in the publication of Ferns for American Gardens (1994; reprinted 2003). Dr. Alan R. Smith (right) was born in Sacramento, California, in 1943. He obtained his B.S. degree in Botany from Kansas State University in 1965, and then went to Iowa State University, where he earned his Ph.D. in 1969 as a student of Dr. John T. Mickel. Dr. Smith has been a Curator and Research Botanist in the University Herbarium, University of California, Berkeley, from 1969 to the present, and has taught several courses and many fern workshops there. A pivotal moment in his career came as a result of being a participant in an Organization of Tropical Studies (OTS) summer course in Pteridology, in Costa Rica, in 1967, a course coordinated by Dr. Mickel with other leading pteridologists. Dr. Smith began work on the Mexican pteridophytes in the state of Chiapas, shortly after arriving in Berkeley, and has continued floristic work on American ferns, especially those from California, Mexico, Central America, Venezuela, and Bolivia. He has contributed treatments of Thelypteridaceae, one of the largest and most complex fern families, to a number of other floras, including those of North America north of Mexico, Guatemala, Mesoamerica, Ecuador, Peru, and the Guianas. He has also conducted monographic and revisionary work on Thelypteris, grammitids, and other groups. Fern cytology and phytogeography are two additional areas of long-standing interest. Dr. Smith was Pteridophyte Editor for the Flora of North America North of Mexico, Vol. 2 (1993), and also served on the Editorial Board, Management Committee, and finally as Chairperson of the project. In recent years, he has been extensively involved in collaborative phylogenetic work on relationships of the basal (evolutionarily older) ferns and on the more derived families Polypodiaceae, Grammitidaceae, and Thelypteridaceae. Dr. Smith has served the American Fern Society as President and as Editor of the American Fern Journal.