Flora of the Venezuelan Guayana, Volume 8: Poaceae-Rubiaceae [8] 1930723369, 9781930723368

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Volume 8 is enhanced by 659 drawings of plants, mostly by noted botanical artist Bruno Manara. The entire flora will contain close to 5,000 drawings, or roughly half of the nearly 10,000 species treated. Mr. Manara worked closely with Dr. Steyermark and accompanied him on a number of his collecting expeditions.

Left to right: Bruce Holst, Kay Yatskievych, and Paul Berry. JACKET ILLUSTRATIONS: (front) a typical landscape in the eastern part of the Venezuelan Guayana showing part of the Chimantá-massif and Angasima-tepui, photo by Karl Weidmann; (back, clockwise from upper left) Stegolepis hitchcockii subsp. morichensis (Rapateaceae) from a cultivated plant, photo by Claudia Lipke; Isertia verrucosa (Rubiaceae) along the Yavita-Maroa road, Amazonas state, photo by Paul E. Berry; and Rhyncholacis penicillata (Podostemaceae) along Salto Hacha, Canaima, Bolívar state, photo by Paul E. Berry.

FLORA OF THE VENEZUELAN GUAYANA

The Flora of the Venezuelan Guayana was initiated by Dr. Julian A. Steyermark, a botanist and one of the most prolific plant collectors of all time, who personally gathered more than 135,000 different specimens. Dr. Steyermark completed many important publications during his life, including the Flora of Missouri and the Flora of Guatemala (as coauthor), but his death in 1988 left the present flora at an early stage. The task of organizing and completing this flora was placed in the capable hands of Dr. Paul E. Berry, now at the University of Wisconsin at Madison, who has spent much of his career studying the Venezuelan flora. Dr. Berry was joined by two other colleagues to produce these volumes. Kay Yatskievych at the Missouri Botanical Garden in St. Louis provided editorial expertise, a knowledge of desktop publishing and graphic design, and is the coordinating editor for the project. Bruce K. Holst, at the Marie Selby Botanical Gardens in Sarasota, Florida, was an important collaborator of Dr. Steyermark for many years and gained an intimate knowledge of the area’s flora as a result.

VOLUME 8

POACEAE -RUBIACEAE

This is the eighth book in a nine-volume flora that is the first full scientific account of the plants of a botanically rich and geologically ancient part of northern South America, the Venezuelan Guayana. This area is dominated by massive table mountains, tepuis, that tower over surrounding rain forest and savannas and provide a wealth of habitats for nearly 10,000 species of vascular plants. Volume 1 provides a general introduction to the southeastern half of Venezuela that encompasses the flora area, with chapters on geography, history of botanical exploration, vegetation types, endemism, and conservation. Keys to the families of seed plants are also provided. The present volume continues the alphabetical sequence of family treatments begun in Volume 2, which treated the ferns and their allies and the first 11 families of seed plants. Keys, descriptions, and illustrations of more than half the species treated — a feature rarely found in floras of even more familiar areas — make this work an enduring reference that will be useful well beyond the borders of the vast region covered by the flora.

VOL 8

More than 200 botanists from around the world have collaborated on this project, which is based at the Missouri Botanical Garden.

POACEAE RUBIACEAE

GENERAL EDITORS

Missouri Botanical Garden Press P.O. Box 299 St. Louis, Missouri 63166-0299 U.S.A. www.mbgpress.org ® Printed in U.S.A.

FLORA OF THE VENEZUELAN GUAYANA

Flora of the Venezuelan Guayana VOLUME 8

Missouri Botanical Garden Press

Julian A. Steyermark, Paul E. Berry, Kay Yatskievych, and Bruce K. Holst

ISBN 1-930723-36-9

Flora of the Venezuelan Guayana

GENERAL EDITORS Julian A. Steyermark, Paul E. Berry, Kay Yatskievych, and Bruce K. Holst

Flora of the Venezuelan Guayana VOLUME 8 POACEAE–RUBIACEAE

VOLUME EDITORS Paul E. Berry, Kay Yatskievych, and Bruce K. Holst Illustrated by Bruno Manara

MISSOURI BOTANICAL GARDEN PRESS St. Louis

Copyright © 2004 by the Missouri Botanical Garden Press All rights reserved. ISBN 1-930723-36-9 Library of Congress Control Number: xxxxxxxxxxxx Printed in U.S.A. Published on xxxxxxxxxxxx by Missouri Botanical Garden Press P.O. Box 299 St. Louis, Missouri 63166-0299, U.S.A.

Contents Preface and Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . vii Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ix Contributors . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xiii Poaceae (Gerrit Davidse, Emmet J. Judziewicz, and Fernando O. Zuloaga) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 Podocarpaceae (David J. de Laubenfels) . . . . . . . . . . . . . . . . . . . . . . . . . 297 Podostemaceae (Paul E. Berry) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 301 Polygalaceae (Gerardo A. Aymard C., Paul E. Berry, and Bente Eriksen) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 316 Polygonaceae (Gerardo A. Aymard C. and Richard A. Howard) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 347 Pontederiaceae (Charles N. Horn) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 371 Portulacaceae (Julian A. Steyermark) . . . . . . . . . . . . . . . . . . . . . . . . . . . 376 Primulaceae (James S. Miller and Paul E. Berry) . . . . . . . . . . . . . . . . . . 383 Proteaceae (Julian A. Steyermark) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 384 Quiinaceae (Julio V. Schneider and Georg Zizka) . . . . . . . . . . . . . . . . . . 393 Rafflesiaceae (George Yatskievych and Willem Meijer) . . . . . . . . . . . . . . 407 Ranunculaceae (Paul E. Berry) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 410 Rapateaceae (Paul E. Berry) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 413 Rhamnaceae (Julian A. Steyermark and Paul E. Berry) . . . . . . . . . . . . . 473 Rhizophoraceae (Julian A. Steyermark) . . . . . . . . . . . . . . . . . . . . . . . . . . 484 Rosaceae (Gerardo A. Aymard C. and Nidia L. Cuello A.) . . . . . . . . . . . 490 Rubiaceae (Charlotte M. Taylor, Julian A. Steyermark, Piero G. Delprete, Alberto Vincentini, Rocio Cortés, Daniela Zappi, Claes Persson, Cristina Bestetti Costa, and Elisete Araujo da Anunciação) . . . . . . . . . . . . . . . . . . . . . . . . . 497

Appendix: List of new names and emendations published in this volume . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 848 Index (compiled by George Thornburgh) . . . . . . . . . . . . . . . . . . . . . . . . . 849

v

Preface and Acknowledgments This volume continues the taxonomic treatments of all native and naturalized vascular plant families known to occur in the Venezuelan Guayana. It includes 17 families of flowering plants, from Poaceae through Rubiaceae. The remaining families will be published in alphabetical order in the following ninth volume. The Flora of the Venezuelan Guayana project has been strongly supported since its inception by the Missouri Botanical Garden and especially by its director, Peter H. Raven. The Herbario Nacional de Venezuela, now part of the Fundación Instituto Botánico de Venezuela, has supported this project in many ways; it was there that Julian Steyermark conceived the idea of the flora and did most of his background research. The Julian A. Steyermark Fund, established by the late Dr. Steyermark at the Missouri Botanical Garden, provided major funding for the flora. The Wisconsin State Herbarium and the Botany Department at the University of Wisconsin–Madison has also supported this project in its later stages. This volume is based upon work supported by the National Science Foundation under Grants Nos. BSR-8717303, BSR-9045532, and BSR-9201044. The Foundation provides awards for research in the sciences and engineering. The awardee is wholly responsible for the conduct of such research and preparation of the results for publication. The Foundation, therefore, does not assume responsibility for such findings or their interpretation. Any opinions, findings, conclusions, or recommendations expressed in this publication are those of the authors and do not necessarily reflect the views of the National Science Foundation. Besides the illustrations by Bruno Manara, Cathy Pasquale made the drawing of Spartina alterniflora (Fig. 221). This was originally published in the Flora of the Guianas (A.R.A. Görts-van Rijn, ed. Series A: Phanerogams, Fascicle 8. 1990) and is used with the permission of the Smithsonian Institution, Washington, D.C. Kandis Elliot, artist at the University of Wisconsin–Madison Botany Department, redrew several of the Podostemaceae figures and added details important to their identification. She also assisted with the selection and layout of the photographs on the back of the dust jacket. This project has benefited greatly from the knowledge and assistance of Otto Huber, the main contributor to Volume 1 and an expert on the flora and vegetation of the Venezuelan Guayana. We are grateful to Masahiro Kato and Thomas Philbrick for their reviews of the Podostemaceae. Numerous institutions and individuals in Venezuela have contributed significantly to different aspects of the flora. These include the Corporación Venezolana de Guayana and its affiliate Electrificación del Caroní, C.A.; the Ministerio del Ambiente y de los Recursos Naturales Renovables; the Instituto Nacional de Parques and the Dirección General de Parques Nacionales; the Servicio Autónomo vii

viii P REFACE

AND

A CKNOWLEDGMENTS

para el Desarrollo Ambiental del Estado Amazonas; and the Consejo Nacional de Investigaciones Científicas y Tecnológicas. The Fuerzas Aéreas de Venezuela, Fundación Terramar, Charles Brewer-Carías, and the late Parker Redmond also gave important logistical support on various trips to southern Venezuela. Several Venezuelan herbaria and associated botanists provided considerable assistance over an extended period of time, including the Herbario Nacional de Venezuela, especially through Francisco Delascio, Francisco Guánchez, Gilberto Morillo, and Zoraida Luces de Febres; the Universidad Central de Venezuela, through the Herbario Ovalles at the Facultad de Farmacia and its director, Stephen Tillett, and the herbarium of the Facultad de Agronomía in Maracay, particularly with the help of Carmen Emilia Benítez de Rojas; the Universidad Nacional Experimental de los Llanos Ezequiel Zamora in Guanare and its active team of botanists including Gerardo Aymard, Nidia Cuello, and Basil Stergios; the herbarium at Puerto Ayacucho, now associated with the Centro Amazónico de Investigaciones Ambientales Alejandro de Humboldt of the Servicio Autónomo para el Desarrollo Ambiental del Estado Amazonas; and the Universidad de Los Andes in Mérida, mainly through the herbaria of the Facultad de Ciencias Forestales and the Facultad de Farmacia. In the United States, the National Geographic Society provided grants for several explorations associated with this project. Three major institutions, the Missouri Botanical Garden, the New York Botanical Garden, and the Smithsonian Institution, lent their facilities, specimens, and the expertise of their staff to assist in the preparation of numerous floristic treatments. The project has also benefited from the efforts of two postdoctoral researchers, Denis Kearns and John MacDougal. Carlos Vargas worked for a year as a full-time Research Assistant for the project. Other people who worked part time or volunteered to assist on the flora include William Betz, Lois Brako, Germán Carnevali, Kandis Elliot, Luther Raechal, Ronald Liesner, Ivón Ramírez, Erika Rohrbach, and George Yatskievych. A special thanks to Shirley Flavin and George Thornburgh, whose help as volunteers was instrumental in seeing this volume through to completion.

Introduction The Flora of the Venezuelan Guayana is an illustrated, synoptical treatment of the native and naturalized vascular plants that occur in the southern Venezuelan states of Amazonas, Bolívar, and Delta Amacuro (see endpaper map). This area includes spectacular tabletop mountains called tepuis and is known for its high vascular plant endemism. The flora area covers almost 500,000 square kilometers and includes about half the area of the geologically ancient Guayana Shield. The project is centered at the Missouri Botanical Garden and is a collaborative effort with about 200 contributing authors worldwide. It is significant because it is the first effort to produce a comprehensive inventory and identification guide for the plants of such an extensive region of northern South America. Volume 1 of the Flora of the Venezuelan Guayana provides extensive background information for the taxonomic treatments that began in Volume 2. It includes chapters on the physical geography of the region and its human occupation, the history of botanical exploration, the classification and altitudinal zonation of vegetation types in the flora area, floristic and phytogeographical analyses, and the conservation importance of the region. There is also a section of color photographs of landscapes, vegetation types, and plants, as well as a key to the families of seed plants in the flora area. Volume 1 is accompanied by two oversize maps of the Venezuelan Guayana at a scale of 1:2,000,000, one a topographical map with an index to place names, the other a multicolor map of vegetation types. Volume 2 of the Flora of the Venezuelan Guayana treated the ferns and fern allies, or pteridophytes. It also included an introduction to the pteridophytes and a key to the pteridophyte families, followed by the corresponding family treatments in alphabetical order. The volume then continued with the first 11 seed plant families, again in alphabetical order, beginning with Acanthaceae and ending with Araceae. Volume 3 contained 20 families from Araliaceae through Cactaceae; Volume 4 contained 35 families from Caesalpiniaceae through Ericaceae; Volume 5 contained 29 families from Eriocaulaceae through Lentibulariaceae; Volume 6 contained 28 families from Liliaceae to Myrsinaceae; and Volume 7 contained 18 families from Myrtaceae to Plumbaginaceae. Family names of flowering plants in the Flora of the Venezuelan Guayana mostly follow those used by A. Cronquist in An Integrated System of Classification of Flowering Plants (Columbia University Press, New York. 1981). Also included are certain families published after Cronquist’s book, such as Euphroniaceae and

ix

x INTRODUCTION

FAMILIES INCLUDED IN THIS VOLUME Genera Species Poaceae Podocarpaceae Podostemaceae Polygalaceae Polygonaceae Pontederiaceae Portulacaceae Primulaceae Proteaceae

96 1 8 7 6 3 2 1 3

409 9 26 63 45 7 13 1 16

Genera Species Quiinaceae Rafflesiaceae Ranunculaceae Rapateaceae Rhamnaceae Rhizophoraceae Rosaceae Rubiaceae

4 1 1 14 5 3 3 86

22 1 2 72 20 8 10 524

244

1248

Monotaceae. A complete listing of the families to be covered in the Flora of the Venezuelan Guayana with the numbers of genera and species currently recorded from the flora area appears in Appendix A of Volume 1. The 17 families that appear in Volume 8 and their numbers of native or naturalized genera and species in the Venezuelan Guayana are given above. Organization and Scope of the Floristic Treatments The floristic treatments in this flora are organized alphabetically by family, first within the ferns and fern allies, then within the seed plants. The gymnosperms, monocotyledons, and dicotyledons are not treated separately within the seed plant category. The authors for each family treatment are cited below the family heading; in multi-authored families, if different parts were contributed by different authors, the respective authors are cited separately under the parts they were responsible for (such as a particular genus or the key to the genera). Each family treatment contains a complete description of the family and the genera that are present in the flora; these descriptions are worldwide in scope. Within each family, genera are arranged in alphabetical order. At the end of each family or genus description, a separate paragraph lists the overall distribution of the family or genus. This is followed by the author’s estimate of the number of subordinate taxa worldwide, as well as the number of taxa present in the Venezuelan Guayana. For each genus, the estimated number of species in all of Venezuela is also given. All keys used in the flora are dichotomous and bracketed. In each couplet, the number in parenthesis refers to the couplet that led to it. The keys to the genera are designed primarily to cover the species that occur in the Venezuelan Guayana and not necessarily species found elsewhere. Slightly more than half the species in the flora are illustrated by black-andwhite line drawings. Volume 8 includes 659 figures. The line drawings are consecu-

I NTRODUCTION

xi

tively numbered within each volume, and they are usually grouped together by genus near the end of the corresponding species entries to provide easy visual comparisons between species. Since the illustrations and the keys are designed to enable identification of plants from the flora area, the flora does not provide full species descriptions. Species entries are organized alphabetically within their respective genus and begin with the accepted species name, author(s), and place of publication. If the accepted name is a combination, it is immediately followed by its basionym and then by other synonyms based on the same type. Common or vernacular plant names that are known to be used within the Venezuelan Guayana are listed at the end of this paragraph. Synonyms based on a different type from the accepted name are listed in separate paragraphs in chronological order of basionym publication. Synonymy is not intended to be exhaustive, but rather includes synonyms pertinent to the Venezuelan Guayana and adjacent areas. We have attempted to include all names originally based on collections from the flora area. Following the nomenclatural portion of each species entry, a new paragraph indicates the plant’s habit and sometimes includes diagnostic characters of the plant that were not included in the keys. This is followed by a listing of the habitats and the elevational range where the taxon occurs in the flora area only (not worldwide), as well as its geographical distribution within the Venezuelan Guayana. Distribution outside the flora area is listed next, beginning with any states or regions of Venezuela north of the Río Orinoco where the taxon occurs, then any other countries or regions of occurrence. Whenever a taxon is illustrated by a black-and-white line drawing, the symbol “◆” followed by the figure number appears at the end of the paragraph. A separate paragraph is sometimes used to provide taxonomic discussion and/or notes on the uses of the plant in the flora area. Any varieties or subspecies present in the flora area appear below their corresponding species entry, and they contain a similar level of information as regular species entries. A key is included if more than one subspecies or variety is present. Forms found in the flora area do not have separate entries, but they are discussed under the next higher rank. Throughout the flora, author abbreviations follow Authors of Plant Names (R. K. Brummitt and C. E. Powell, editors. Royal Botanic Gardens, Kew. 1992). The only exception is the use of the traditional abbreviation “H.B.K.” for the authors of the publication Nova Genera et Species Plantarum (A. von Humboldt, A. Bonpland, and K. S. Kunth. Librairie Grecque-Latine-Allemande, Paris. 1815–1825). This series of seven volumes was published in two versions (quarto and folio), each with different page numbers. Page numbers cited in this flora refer to the quarto version, which is more widely available than the folio version, except for volume four, which is the only volume in which the folio edition was distributed before the quarto. Book abbreviations follow Taxonomic Literature: A Selective Guide to Botanical Publications and Collections with Dates, Commentaries and Types, 2nd edition (F. A. Stafleu and R. S. Cowan. Bohn, Scheltema & Holkema, Utrecht, Netherlands. 1976–1988), and journal abbreviations follow Botanico-Periodicum-Huntianum (G. H. M. Lawrence, A. F. G. Buchheim, G. S. Daniels, and H. Dolezal, editors. Hunt Botanical Library, Pittsburgh, Pennsylvania. 1968) or B-P-H/S: BotanicoPeriodicum/Supplementum (G. D. R. Bridson and E. R. Smith, editors. Hunt Institute for Botanical Documentation, Pittsburgh, Pennsylvania. 1991). When the effec-

xii

INTRODUCTION

tive publication date of a reference is different from the imprint date, the effective publication date is placed in brackets following the imprint date. Herbarium abbreviations follow Index Herbariorum, 8th edition (P. K. Holmgren, N. H. Holmgren, and L. C. Barnett, editors. New York Botanical Garden, Bronx, New York. 1990). For each family treated in the flora we have prepared lists of selected voucher specimens of the accepted taxa. These are specimens that have been seen and verified by the authors of the different treatments and are arranged alphabetically by genus, species, and collector. These exsiccatae lists are not included in the flora, but are available upon request from the Missouri Botanical Garden, c/o Flora of the Venezuelan Guayana.

Contributors Elisete Araujo da Anunciação Instituto de Botânica (SP) Seção de Fanerógamas Caixa Postal 4005 01061-970 São Paulo, SP BRAZIL

Gerrit Davidse Missouri Botanical Garden P. O. Box 299 St. Louis, Missouri 63166

Gerardo A. Aymard C. Herbario Universitario PORT UNELLEZ–Guanare Mesa de Cavacas 3323, Portuguesa VENEZUELA

Piero Delprete National Herbarium of the Netherlands Utrecht University branch (U) Plant Systematics Heidelberglaan 2 3584 CS Utrecht THE NETHERLANDS

Paul E. Berry Botany Department University of Wisconsin–Madison 132 Birge Hall 430 Lincoln Drive Madison, Wisconsin 53706

Bente Eriksen Herbarium Botanical Museum Carl Skottsbergs Gata 22 S-413 19 Göteborg SWEDEN

Rocio Cortés Herbario Forestal (UDBC) Universidad Distrital Carrera 8, No. 40-78/ 40-50 Santafé de Bogotá, D.C. COLOMBIA Cristina Bestetti Costa Instituto de Botânica (SP) Seção de Fanerógamas Caixa Postal 4005 01061-970 São Paulo, SP BRAZIL Nidia L. Cuello A. Herbario Universitario PORT UNELLEZ–Guanare Mesa de Cavacas 3323, Portuguesa VENEZUELA

Bruce K. Holst Marie Selby Botanical Gardens 811 South Palm Ave. Sarasota, Florida 34236 Charles N. Horn Department of Biology Newberry College Newberry, South Carolina 29108 Richard A. Howard (deceased) c/o Harvard University Herbaria 22 Divinity Avenue Cambridge, Massachusetts 02138 Emmet J. Judziewicz Department of Biology CNR Building UW-Stevens Point Stevens Point, Wisconsin 54481 xiii

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CONTRIBUTORS

David J. de Laubenfels Department of Geography Syracuse University Syracuse, New York 13244 Willem Meijer (deceased) c/o School of Biological Sciences 101 Morgan Building University of Kentucky Lexington, Kentucky 40506 James S. Miller Missouri Botanical Garden P. O. Box 299 St. Louis, Missouri 63166 Claes Persson Botanical Institute (GB) Goteborg University P.O. Box 461 SE 40530 Goteborg SWEDEN

Charlotte M. Taylor Missouri Botanical Garden P. O. Box 299 St. Louis, Missouri 63166 Alberto Vincentini Missouri Botanical Garden P. O. Box 299 St. Louis, Missouri 63166 George Yatskievych Missouri Botanical Garden P. O. Box 299 St. Louis, Missouri 63166 Kay Yatskievych Missouri Botanical Garden P. O. Box 299 St. Louis, Missouri 63166 Daniela Zappi Royal Botanic Gardens, Kew Richmond, Surrey TW9 3AE ENGLAND, U.K.

Julio V. Schneider Botanik/Palaeobotanik Forschungsinstitut Senckenberg Senckenberganlage 25 D-60325 Frankfurt am Main GERMANY

Georg Zizka Botanik/Paläobotanik Forschungsinstitut Senckenberg und J.W. Goethe-Universität Senckenberganlage 25 D-60325 Frankfurt am Main GERMANY

Julian A. Steyermark (deceased) c/o Missouri Botanical Garden P. O. Box 299 St. Louis, Missouri 63166

Fernando O. Zuloaga Instituto de Botánica Darwinion Casilla de Correo 22 1642, San Isidro, Buenos Aires ARGENTINA

POACEAE by Gerrit Davidse, Emmet J. Judziewicz, and Fernando O. Zuloaga Herbaceous annuals or perennials, or in some genera the culms strongly lignified; habit various, either cespitose, rhizomatous, stoloniferous, scandent, or aquatic. Culms hollow, pithy, or solid, cylindrical, branching or not at the middle and upper nodes, the branches usually solitary at the nodes (in some genera apparently several per node), bearing bicarinate prophylls as their first appendages. Leaves alternate, estipulate, 2-ranked (rarely spirally arranged); petiole modified into a leaf sheath, the margins usually free; ligule(s) usually present at junction of sheath and blade; inner ligule present (rarely absent) on adaxial surface, membranous and/or ciliate; outer ligule present on abaxial surface in some genera; blades linear to less commonly ovate, flat to inrolled, entire, parallel-veined (rarely obliquely veined), the base of the blade sometimes narrowed into a pseudopetiole. Inflorescences compound, terminal and/or axillary, open to contracted, of few to many discrete, reduced partial inflorescences (spikelets) arranged in 1–several spikes, racemes, or panicles. Spikelets very diverse in structure but typically consisting of an axis (the rachilla) bearing at its base 2 sterile bracts (glumes), these persistent or deciduous, veinless to several-veined; rachilla above the glumes alternately bearing florets, these persistent or deciduous; florets consisting of a subtending bract (lemma) and the first appendage of the floral axis (palea), these enclosing a reduced flower; lemma 1– many-veined, awned or not; palea typically bicarinate, sometimes absent. Flower bisexual or unisexual, the perianth reduced, represented by 2 or 3 (uncommonly none) minute, fleshy or scale-like tepals (lodicules). Androecium of (1, 2)3 or 6 (–many) stamens. Gynoecium with ovary unilocular, uniovulate; styles 2; stigmas (1)2 or 3, hispid to plumose. Fruit usually a caryopsis with the seed coat adnate to the pericarp; endosperm abundant; embryo small to large. Worldwide in tropical, temperate, and arctic regions; ca. 750 genera and 10,000 species, 96 genera and 409 species in the flora area. All 5 subfamilies of grasses occur in the flora area1. The Bambusoideae (21 genera and 80 species in the flora area) consists of the woody bamboos, the herba1

For an alternative, updated suprageneric classification that incorporates numerous recent molecular studies see the Catalogue of New World Grasses [http://mobot.mobot.org/W3T/Search/nwgc.html; http://mobot.mobot.org/W3T/Search/nwgclass.html] and Grass Phylogeny Working Group. 2001. Phylogeny and subfamilial classification of the grasses (Poaceae). Ann. Missouri Bot. Gard. 88: 373–457.

1

2

P OACEAE

ceous bamboos, and the rice-like grasses, all with the C3 photosynthetic pathway and a distinctive leaf anatomy that frequently includes the presence of fusoid and arm cells. The woody bamboos (Bambuseae, 9 genera and 48 species) are characterized by their generally large stature, lignified culms with distinctive bud and branching patterns, dimorphic leaves (culm, foliage) usually with oral setae (stiff bristles produced at the summit of the leaf sheath), pseudopetioles, and outer (as well as inner) ligules; and often long intervals between flowering episodes. As evidenced by the 12 unnamed species designated in this treatment by letters (in Arthrostylidium, Aulonemia, Chusquea, Myriocladus, Neurolepis, and Rhipidocladum), the group is diverse and still not well known in the flora area, and more new species are expected from forested river sides and tepui slopes. The “herbaceous bamboos” (Olyreae, Phareae, and Streptogyneae, with 9 genera and 24 species) are mostly monoecious, lowland forest plants with broad leaf blades. Molecular evidence now indicates that the Phareae are a “basal grass” group, and that the Streptogyneae are more closely related to the marsh-loving or aquatic rice-like members of the Oryzeae (3 genera and 8 species), which have spikelets with the lowest florets often reduced and resembling glumes. The Pooideae are a large, predominantly temperate and montane tropical group represented in the flora area by only one species in the introduced genus Briza. All pooid grasses have the C3 photosynthetic pathway. Several genera and about 25 species occur in the Coastal Cordillera of Venezuela at elevations approximately equivalent to those of the tepui summits, and the complete absence of native taxa in the Guayana Highlands is noteworthy. The Arundinoideae are a large, heterogeneous (both the C3 and C4 photosynthetic pathways are present) subfamily that is sparingly represented (7 genera and 16 species) in the flora area in 4 tribes. The Aristideae (Aristida) are grassland and savanna plants, the Arundineae are found near water (Gynerium, Phragmites) or on tepui summits (Cortaderia), the Centotheceae (Orthoclada, Zeugites) are pseudopetiolate, broad-leaved grasses of lowland forest understories, and the Steyermarkochloeae (Steyermarkochloa) are restricted to white-sand savannas in the flora area and adjacent Brazil. The Chloridoideae (17 genera and 38 species) are mostly lowland plants adapted to seasonally dry grasslands and savannas; all have the C4 photosynthetic pathway. Two tribes occur in the flora area, the Pappophoreae (1 species) and the Cynodonteae (16 genera and 37 species). Morphologically, the chloridoids often have ciliate ligules, inflorescences with digitately arranged racemose branches, and laterally compressed spikelets with the uppermost floret(s) reduced in size. The Panicoideae (50 genera and 276 species) are the dominant subfamily in the flora area, accounting for 1/2 of the genera and 2/3 of species-level diversity. The C3 and C4 photosynthetic pathways are both found in this group. The Paniceae (31 genera and 232 species) are found in many habitats, including tepui summits, but are most characteristic of lowland savannas and forest edges. The tribe includes the large genera Panicum (53 species), Paspalum (46 species), and Axonopus (34 species), and is most easily distinguished by its plump, terete or plano-convex, 2-flowered spikelets, with the lower floret sterile or staminate and the upper floret bisexual and coriaceous to indurate in texture. The Andropogoneae (17 genera and 41 species) is characterized by its paired sessile and pedicellate spikelets and prefers

P OACEAE 3

savannas, where some genera and species are ecological dominants. Two other panicoid tribes present in the flora area are the Arundinelleae (1 species) and the Isachneae (1 genus and 2 species). This treatment does not include grasses that occur in the flora area solely in cultivation, such as Saccharum officinarum L. (Caña de azúcar, Sugar cane) and Zea mays L. (Maize, Corn). There is also an unconfirmed report of the adventive Poa annua L. from the summit of Roraima-tepui at a site where hikers often camp on the summit.

Key to the Tribes and Genera of Poaceae by Emmet J. Judziewicz 1.

1. 2(1). 2. 3(2). 3.

4(2).

4.

5(4). 5. 6(5).

6.

7(5). 7.

Inflorescence subtended by a hollow, bony, usually white bead 9–12 mm long that conceals the pistillate spikelets, the staminate spikelets exserted through an apical pore ..... Panicoideae: Andropogoneae (19. Coix) Inflorescence not subtended by a bony bead ............................................ 2 All or some spikelets partially or completely concealed in spiny burs or deciduous fascicles of flat, indurate bracts ........................................... 3 Spikelets borne in panicles, racemes, or spikes, not concealed in spiny burs or by flat, indurate bracts ............................................................. 4 Inflorescence a spike of spikelet groups concealed within indurate, fascicled bracts, on a zigzag rachis ...... Panicoideae: Paniceae (4. Anthephora) Inflorescence a spike of readily deciduous, spiny burs, these each concealing 1–several spikelets, on a straight rachis .......................................... ......................................................... Panicoideae: Paniceae (15. Cenchrus) Leaves all bladeless except for the uppermost one which has a tiny, reduced blade; plants robust, reed-like; the inflorescence a spicate panicle with pistillate spikelets above, staminate below ...................... ................... Arundinoideae: Steyermarkochloeae (88. Steyermarkochloa) Leaves all with well-developed blades; plants of various habits, but never producing a spicate panicle with pistillate spikelets above and staminate ones below ...................................................................................... 5 Leaves with an outer ligule present (in addition to the inner ligule), the blades ultimately deciduous from the sheaths ..................................... 6 Leaves without an outer ligule (inner ligule usually present), the blades not or rarely deciduous .......................................................................... 7 Plants herbaceous, < 1 m tall, unbranched above the base; leaves all alike; summit of leaf sheath never bearing oral setae; herbaceous perennials ....................................... Bambusoideae: Streptogyneae (89. Streptogyna) Plants lignified, (1–)2–30 m tall, the branches often several at each node; leaves dimorphic, those of the culm different from those of the foliage complements; summit of leaf sheath often bearing oral setae; woody bamboos ............ Bambusoideae: Bambuseae (see Subkey 3, page 6) Spikelets strictly 1-flowered, the floret subtended by 2 or fewer glumes, the rachilla internode not prolonged past the floret ............................ 8 Spikelets 2–many-flowered, or, if 1-flowered, then with the rachilla inter-

4

P OACEAE

8(7). 8. 9(8).

9.

10(9).

10.

11(10).

11. 12(8). 12. 13(12). 13. 14(12).

14. 15(7).

15.

16(15).

16. 17(16).

node prolonged past the floret or with 2 glumes and 2 glume-like sterile lemmas present ............................................................................... 15 Spikelets unisexual, dimorphic ................................................................. 9 Spikelets bisexual, monomorphic ............................................................ 12 Pseudopetioles 8–60 mm long, twisted at the summit to invert the blade; blades with oblique venation; pistillate spikelets with florets covered with hooked hairs ............................. Bambusoidae: Phareae (70. Pharus) Pseudopetioles 1–7 mm long, not twisted at the summit; blades with parallel venation; pistillate spikelets with florets never covered with hooked hairs ......................................................................................... 10 Inflorescence spicate, of many deciduous whorls, each whorl composed of 4–6 staminate spikelets surrounding and concealing a single pistillate spikelet; summit of leaf sheaths often with oral setae evident ............. ........................................................ Bambusoideae: Olyreae (65. Pariana) Inflorescence open, not spicate, the staminate spikelets not in whorls surrounding and concealing the pistillate spikelets; oral setae absent .............................................................................................................. 11 Pistillate spikelets with well-developed glumes, these mostly exceeding the floret; leaf blades usually broad; plants often found in shaded forests ...................... Bambusoideae: Olyreae (see Subkey 4, page 7) Pistillate spikelets with glumes absent; leaf blades narrow; plants often aquatic ............................................. Bambusoideae: Oryzeae (51. Luziola) Florets with midvein of lemma prolonged into a prominently 3-parted awn ....................................................................................................... 13 Florets awnless or awned, but the midvein of the lemma never prolonged into a prominently 3-parted awn ........................................................ 14 Glumes awned; spikelets in triads; forest edges at high elevations on tepuis ..................................... Chloridoideae: Cynodonteae (2. Aegopogon) Glumes awnless; spikelets solitary, not in triads; mostly in lowland savannas ...................................... Arundinoideae: Aristideae (6. Aristida) Spikelets laterally compressed, gibbous, the veins often hispidulous; palea 3-veined; often aquatic .................................................................. ................................ Bambusoideae: Oryzeae (see Subkey 5, page 8) Spikelets terete, ovoid to lanceoloid, the veins not hispidulous; palea 2-veined; terrestrial ........... Chloridoideae: Cynodonteae (85. Sporobolus) Spikelets 2-flowered, or if 1-flowered, then with the rachilla internode prolonged past the floret, or with 2 glumes and 2 glume-like sterile lemmas present .................................................................................... 16 Spikelets 3–many-flowered (some spikelets in a given inflorescence may be 2-flowered), or 2-flowered with the rachilla internode prolonged past the upper floret ............................................................................ 23 Plants 4–8 m tall, the culms solid; inflorescence a large panicle (often to 1 m long) with drooping branches ........................................................... ................................................ Arundinoidae: Arundineae (36. Gynerium) Plants < 4 m tall, the culms hollow to solid; inflorescence various, < 1 m long ....................................................................................................... 17 Leaves all basal, the blades (50–)100–165 × (2.5–)4–6.5 cm; plants 1–4(–10) m tall; spikelets with 2 glumes and 2 glume-like sterile lemmas below the solitary florets ............... Bambusoideae: Bambuseae (56. Neurolepis)

P OACEAE 5

17.

18(17).

18.

19(18). 19. 20(19). 20. 21(19).

21.

22(21).

22.

23(15).

23.

24(23). 24. 25(24). 25. 26(25). 26. 27(26).

Leaves cauline (or some basal), the blades < 100 cm long, usually < 4 cm wide; plants of various sizes; spikelets never with 2 glumes and 2 glume-like sterile lemmas present below the floret ........................... 18 Spikelets laterally compressed, the glumes and sterile lower floret persistent, awnless; upper floret bisexual, deciduous, awned; glumes slightly recurved at the apex ............ Panicoideae: Arundinelleae (8. Arundinella) Spikelets without the above combination of characters, most characteristically lacking laterally compressed spikelets, or the glumes not recurved, or the upper floret awnless .................................................... 19 Both florets indurate and hemispherical, the spikelets globose; lower floret staminate or bisexual, the upper floret pistillate ......................... 20 Both florets not indurate and hemispherical, the spikelets globose or not; florets with sexuality various .............................................................. 21 Lower glume present, nearly as long as the spikelet; upper floret strawcolored .............................................. Panicoideae: Isachneae (44. Isachne) Lower glume absent; upper floret brown .................................................... .................................. Panicoideae: Paniceae (68. Paspalum, P. convexum) Spikelets usually paired, one (sub)sessile and the other pedicellate, often dimorphic and with the pedicellate spikelet much reduced; if not paired, then spikelets either in readily disarticulating racemes or the hyaline upper floret geniculately awned ................................................ ..................... Panicoideae: Andropogoneae (see Subkey 7, page 10) Spikelets not paired and dimorphic; or if paired, then neither in readily disarticulating racemes, the upper floret not geniculately awned .............................................................................................................. 22 Spikelets with lower floret usually bisexual, not reduced, the upper floret usually sterile and reduced; floret(s) falling from the persistent glumes ........................ Chloridoideae: Cynodonteae (see Subkey 6, page 8) Spikelets with lower floret reduced, either staminate or sterile (rarely pistillate or bisexual), the upper floret bisexual; spikelets usually falling entire from the summit of the pedicel .............................................. ................................. Panicoideae: Paniceae (see Subkey 8, page 12) Pseudopetioles 8–50 mm long; leaf blades lanceolate-ovate to elliptical, strongly cross-veined beneath; in forest shade ...................................... ...................... Arundinoideae: Centotheceae (see Subkey 2, page 6) Pseudopetioles absent or < 5 mm long; leaf blades linear to narrowly lanceolate, never cross-veined beneath; in open habitats or uncommonly in forest shade ...................................................................................... 24 Dioecious grasses 0.5–1 m tall with bushy panicles; usually above 2000 m elevation .............................. Arundinoideae: Arundineae (20. Cortaderia) Plants not dioecious, of various habits; usually below 2000 m elevation .............................................................................................................. 25 Lowest (bisexual) floret 11–15-awned ......................................................... ..................................... Chloridoideae: Pappophoreae (63. Pappophorum) Lowest floret awnless or 1(–3)-awned ..................................................... 26 Coarse, reed-like grasses with culms 1–8 m tall and 1–3 cm diameter ......................... Arundinoideae: Arundineae (see Subkey 1, page 6) Slender grasses with culms < 1 m tall and 1 cm diameter .................... 27 Spikelets about as long as wide; glumes and lemmas circular, spreading

6

P OACEAE

27.

at right angles to the rachilla; rare weed ...... Pooideae: Poeae (14. Briza) Spikelets longer than wide; glumes and lemmas ovate to linear, ascending ....... Chloridoideae: Cynodonteae (see Subkey 6, page 8)

Subkey 1 to the Genera of Arundinoideae: Arundineae 1. 1. 2(1).

2.

Leaf blades 1–4 mm wide, in a basal cluster, the culms few-leaved; tepui summits ................................................................................ 20. Cortaderia Leaf blades 15–60 mm wide, disposed all along culm or in a fan-shaped, terminal cluster; below 200 m elevation .............................................. 2 Culms solid; leaves all terminal in a fan-shaped cluster, the blades 100– 200 × 3–6 cm, the margins serrulate; spikelets 2-flowered, the staminate spikelets glabrous; commonly flowers, to 1 m long ...... 36. Gynerium Culms hollow; leaves disposed all along culm, the blades 50–75 × 1.2– 2 cm, the margins smooth to scaberulous; spikelets 3–many-flowered, the staminate spikelets often pubescent; rarely flowers in the tropics, < 0.5 m long ......................................................................... 71. Phragmites

Subkey 2 to the Genera of Arundinoideae: Centotheceae 1. 1.

Leaf blades 2–4 cm wide; spikelets 2- or 3-flowered, the florets bisexual, all alike................................................................................. 59. Orthoclada Leaf blades 6–9 cm wide; spikelets 5–8-flowered, the florets unisexual, only the lowest one pistillate, the upper ones all staminate .......... 96. Zeugites

Subkey 3 to the Genera of Bambusoideae: Bambuseae 1.

1.

2(1). 2. 3(2).

3.

4(2).

Culms usually over (2–)4 cm diameter, often armed with thorns or spines; inflorescence indeterminate (i.e., composed of pseudospikelets); paleas with keels winged; stamens 6 .................................................. 34. Guadua Culms seldom exceeding 2 cm diameter, always unarmed; inflorescence determinate (i.e., composed of typical spikelets); paleas with keels unwinged; stamens usually 3 ................................................................ 2 Spikelets with 1 functional floret, this subtended by 2 glumes and 2 glume-like bracts ................................................................................... 3 Spikelets with 2–several functional florets, or if only 1 functional floret then this subtended by 2 glumes and only 1 glume-like bract ............ 4 Culms strongly lignified, solid, abundantly branching at the middle and upper nodes; leaves with both outer and inner ligule present, the blades 2–25 cm long ............................................................... 17. Chusquea Culms barely lignified, solid or hollow, in vegetative state unbranched above the base; leaves lacking an outer ligule, the blades (40–)100– 250 cm long ........................................................................... 56. Neurolepis Culms with an extremely long first internode (usually > 1 m long), followed by 1–many short internodes (ca. 1 cm long) .............................. 5

P OACEAE 7

4. 5(4).

Culms with all internodes of about the same length ............................... 6 Culms 12–15 m long, branched above the base, with 50–100 branches per node; spikelets 4–7 cm long, the lemmas with awns 3–10 mm long; Cerro Marahuaka, Sierra de Maigualida ............................................... ......................................................... 7. Arthrostylidium (A. schomburgkii) 5. Culms 0.5–5 m long, unbranched above the base; spikelets 0.5–1.3 cm long, the lemmas awnless; widespread ............................. 55. Myriocladus 6(4). Dwarf plants < 1.5 m tall; inflorescence < 3.5 cm long; Macizo del Chimantá ........................ 55. Myriocladus (M.involutus, M. steyermarkii) 6. Plants > 1.5 m tall; inflorescence > 3.5 cm long ....................................... 7 7(6). Branch complement at midculm nodes usually consisting of only 1 branchlet; foliage leaves often with reflexed blades and oral setae > 2 cm long; inflorescence an open panicle, the spikelets not borne on racemose branches ............................................................... 10. Aulonemia 7. Branch complement at midculm nodes consisting of 3–many branchlets; foliage leaves with spreading to reflexed blades and oral setae usually < 2 cm long; inflorescence a contracted, usually sparsely flowered panicle or composed of a secund raceme(s) .......................................... 8 8(7). Midculm nodes with branches arising from a promontory, not from an appressed, triangular or fan-shaped meristem .................................... 9 8. Midculm nodes with branches arising from the margin of a triangular or fan-shaped meristem that is appressed and adnate to the culm; inflorescence frequently 1-sided ................................................................. 10 9(8). Spikelets 3–7-flowered (although lowermost and uppermost florets rudimentary) ......................................................................... 7. Arthrostylidium 9. Spikelets 1(2)-flowered, the upper floret if present rudimentary ............. .............................................................................................. 9. Atractantha 10(8). Culm leaves with blades often constricted at the base, reflexed; midculm nodes with branches arising from a triangular meristem; spikelets stout, broadly lanceolate, unawned .................................. 53. Merostachys 10. Culm leaves with blades broadest at base, erect; midculm nodes with branches arising from a fan-shaped meristem; spikelets slender, narrowly lanceolate, often awned ................................ 75. Rhipidocladum

Subkey 4 to the Genera of Bambusoideae: Olyreae 1.

1.

2(1).

2.

Inflorescence spiciform, disarticulating and falling in segments, the segments each consisting of 4–6 staminate spikelets surrounding and concealing a single pistillate spikelet; oral setae frequently present at summit of leaf sheath ............................................................... 65. Pariana Inflorescence an open or contracted panicle, if the latter then the spikelets not in whorls of staminate ones each concealing a pistillate spikelet; oral setae absent .............................................................................. 2 Culms dimorphic, the inflorescences borne on leafless, floriferous culms (uncommonly terminal inflorescences on leafy shoots also present) ................................................................................................................ 3 Culms monomorphic, the inflorescences borne on regular leafy shoots ................................................................................................................ 4

8

P OACEAE

3(2). 3. 4(2). 4.

5(4). 5. 6(4).

6.

7(6). 7. 8(7). 8. 9(6). 9.

Plants 100–200(–400) cm tall; floriferous culms erect to scandent ................................................................................ 57. Olyra (O. ecaudata) Plants 15–40 cm tall, floriferous culms decumbent, often buried in the leaf litter .................................................................................... 72. Piresia Leaf blades 1–2 cm long; plants delicate, the erect portions of culms < 30 cm tall; pistillate florets often falling attached to the glumes .... 5 Leaf blades 2.5–40 cm long; plants delicate to robust, the erect portions of culms (30–)50–300 cm tall; pistillate florets disarticulating from the persistent glumes .................................................................................. 6 Pistillate spikelets black at maturity, the glumes ovate, indurate, closely enveloping the floret ...................................... 66. Parodiolyra (P. lateralis) Pistillate spikelets greenish, the glumes lanceolate, membranous, not closely enveloping the floret .................................................. 73. Raddiella Pistillate spikelets without a distinct internode between the glumes and floret; inflorescences 1(–3) from the uppermost node only, generally open and conspicuous, never produced from the middle nodes of the culm ........................................................................................................ 7 Pistillate spikelet with a distinct internode present between the glumes and floret; inflorescences produced from both terminal and axillary nodes, generally contracted and inconspicuous ................................... 9 Pistillate florets 5–10 mm long ........................................................ 57. Olyra Pistillate florets 1.8–3 mm long ................................................................ 8 Pistillate florets 2.5–3 mm long, deeply pitted ...... 57. Olyra (O. micrantha) Pistillate florets 1.8–2 mm long, smooth ..... 66. Parodiolyra (P. luetzelburgii) Pistillate floret dorsally compressed or terete, lanceolate to elliptical, not separated from the glumes by a swollen internode ................ 5. Arberella Pistillate floret laterally compressed, obtriangular or helmet-shaped, separated from the glumes by a swollen internode ............. 50. Lithachne

Subkey 5 to the Genera of Bambusoideae: Oryzeae 1. 1. 2(1). 2.

Spikelets unisexual, borne in different inflorescences or different parts of the same inflorescence; plants strictly aquatic ........................ 51. Luziola Spikelets bisexual, borne in the same inflorescence; plants aquatic or terrestrial .................................................................................................... 2 Spikelets without glume-like sterile florets below the fertile floret .................................................................................................... 47. Leersia Spikelets with 2 glume-like sterile lemmas below the fertile floret ...................................................................................................... 60. Oryza

Subkey 6 to the Genera of Chloridoideae: Cynodonteae 1. 1.

Spikelets all 1-flowered; inflorescence often a ± open panicle ................... .............................................................................................. 85. Sporobolus Spikelets (1)2–many-flowered; inflorescence paniculate or composed of 1–several spicate racemes (if spikelets 1-flowered then the inflores-

P OACEAE 9

cence composed of racemes) .................................................................. 2 Inflorescence a ± open panicle, if spikelets appressed to branches then not arranged in 2 regular rows ............................................................. 3 2. Inflorescence of 1–many, ± spicate racemes, the spikelets regularly and often densely arranged in 1 or 2 rows................................................... 5 3(2). Apex of lemmas minutely tridentate ............................................ 93. Tridens 3. Apex of lemmas obtuse to acuminate, never tridentate ........................... 4 4(3). Cleistogamous spikelets rarely present in lower leaf sheaths; paleas with keels unwinged; lemmas with apex obtuse or acute to less commonly subaristate; rachilla internodes not bearing tufts of hairs near the base of each floret; widespread ............................................ 30. Eragrostis 4. Cleistogamous spikelets often present in lower leaf sheaths; paleas with keels winged; lemmas with apex attenuate to subaristate; rachilla internodes bearing tufts of hairs near the base of each floret ............................................................................................... 87. Steirachne 5(2). Raceme 1 .................................................................................................... 6 5. Racemes 2–many ....................................................................................... 7 6(5). Spikelets 1-flowered, borne in deciduous clusters of 3 on one side of the rachis, the central, pistillate spikelet larger than the lateral, staminate or sterile spikelets; glumes emarginate, awned ........... 2. Aegopogon 6. Spikelets 5–7-flowered, not borne in clusters of 3, the spikelets all alike in size and sexuality; glumes entire, awnless ........................ 94. Tripogon 7(5). Spikelets falling entire at maturity, 1-flowered, strongly laterally compressed; brackish marshes and dunes .................................... 84. Spartina 7. Spikelets not falling entire, usually disarticulating between the florets, the glumes persistent; spikelets 1–several-flowered, laterally compressed or not ......................................................................................... 8 8(7). Spikelets with 2–many bisexual florets (in addition to any sterile ones that are found above the bisexual ones) ............................................... 9 8. Spikelets with only a single bisexual floret (in addition to any sterile ones found above the bisexual one) ............................................................. 12 9(8). Inflorescence with racemes not digitate, borne singly or in pairs all along the axis ................................................................................................. 10 9. Inflorescence of 1–several digitate racemes at the apex of the culm, or occasionally with another raceme placed just below the terminal cluster ......................................................................................................... 11 10(9). Spikelets 9–13 mm long, the uppermost floret or rudiment with a terminal awn ...................................................................................... 33. Gouinia 10. Spikelets 3–4.5 mm long, the uppermost floret or rudiment awnless .............................................................................................. 48. Leptochloa 11(9). Racemes 4–8 times as long as wide, the rachis prolonged past the uppermost spikelet as a prominent sterile bristle ................ 22. Dactyloctenium 11. Racemes 15–75 times as long as wide, the rachis not prominently prolonged as a sterile bristle ........................................................ 27. Eleusine 12(8). Inflorescence with racemes borne singly or in pairs all along the axis, not digitate ................................................................................................. 13 12. Inflorescence of 1–several digitate racemes at apex of culm, or occasionally with another raceme placed just below the terminal cluster ..... 15 2(1).

10

P OACEAE

13(12). Lowest floret with apex of lemma trilobed, the midvein prolonged into an awn ca. 2 mm long ................................................................. 12. Bouteloua 13. Lowest floret with apex of lemma bifid, the midvein prolonged into an awn 3–15 mm long ............................................................................... 14 14(13). Florets and caryopsis dorsally flattened; awn of lowest lemma 3–6 mm long ..................................................................................... 29. Enteropogon 14. Florets and caryopsis laterally compressed; awn of lowest lemma 8– 15 mm long ........................................................................ 35. Gymnopogon 15(12). Glumes as long as or slightly longer than the florets (excluding their awns); leaf blades lacking a midvein ............................... 35. Gymnopogon 15. Glumes (at least the lower), distinctly shorter than the florets; leaf blades with an evident midvein ...................................................................... 16 16(15). Racemes 8–25; lemmas awned; spikelets 2–several-flowered ..... 16. Chloris 16. Racemes 3–5; lemmas awnless; spikelets 1-flowered and with the rachilla internode prolonged as a sterile rudiment ............................. 21. Cynodon

Subkey 7 to the Genera of Panicoideae: Andropogoneae 1. 1. 2(1).

2.

3(1). 3.

4(3).

4. 5(4). 5. 6(4). 6. 7(6). 7.

8(3).

Staminate and pistillate spikelets separated in different parts of the same inflorescence; plants coarse, maize-like ...................................... 2 Staminate and pistillate spikelets mixed in the same inflorescence ...... 3 Inflorescence subtended by an inflated, bony, white to dark bead, this containing the pistillate spikelets, the deciduous, pedicellate staminate spikelets exserted through the terminal pore of the bead .... 19. Coix Inflorescence of 1–many spicate racemes, not subtended by a bony bead, each raceme with the indurate pistillate spikelets below and the persistent, papery staminate spikelets above .......................... 95. Tripsacum Inflorescence with numerous (> 10) branches or racemes, the individual branches or racemes never subtended by spathe-like bracts .............. 4 Inflorescence with 1–10 branches or racemes, if > 10 then each branch or raceme subtended by 1–many spathe-like bracts (and thus forming a compound inflorescence)........................................................................ 8 Sessile and pedicellate spikelets dissimilar, the sessile spikelet bisexual, larger than the reduced or absent, staminate or sterile pedicellate spikelet ................................................................................................... 5 Sessile and pedicellate spikelets both similar and fertile ........................ 6 Pedicellate spikelets absent, only the pedicel present; leaf blades 4–8 mm wide; savannas .................................................................. 82. Sorghastrum Pedicellate spikelets present but reduced in size, staminate; leaf blades (5–)10–50 mm wide; weedy places ......................................... 83. Sorghum Inflorescence with hairs golden yellow to chestnut brown .... 32. Eriochrysis Inflorescence with hairs white to pinkish or purplish ............................. 7 Rachis of racemes tough, persistent; both spikelets of a pair pedicellate; inflorescence narrowly contracted ......................................... 43. Imperata Rachis of racemes fragile, readily disarticulating along with attached sessile and pedicellate spikelets; inflorescence not or only loosely contracted .................................................................................. 78. Saccharum Spikelets all awnless ................................................................................. 9

P OACEAE 11

8. 9(8). 9. 10(9). 10. 11(10). 11. 12(11). 12. 13(12).

13.

14(8). 14. 15(14). 15. 16(15). 16. 17(15).

17.

18(14). 18. 19(18). 19. 20(19).

20.

Spikelets (at least some of them) awned ................................................ 14 Individual inflorescences of 2–7 racemes ............................... 3. Andropogon Individual inflorescences of 1 raceme ..................................................... 10 Inflorescence densely white-pubescent .................................... 28. Elyonurus Inflorescence glabrous ............................................................................. 11 Sessile and pedicellate spikelets similar ........... 3. Andropogon (A. virgatus) Sessile and pedicellate spikelets dissimilar ........................................... 12 Sessile spikelet globose, inflated, deeply pitted, 1–1.2 mm long ............... .......................................................................................... 37. Hackelochloa Sessile spikelet lanceolate to ovate, not inflated, not deeply pitted, 3.5– 6.5 mm long .......................................................................................... 13 Sessile spikelet with lower glume prominently winged above the middle and at the deeply notched apex; lower floret sterile, the palea rudimentary or absent; pedicel of pedicellate spikelet with a prominent lateral wing near the summit; inflorescence tardily disarticulating ............................................................................................. 18. Coelorachis Sessile spikelet with lower glume not winged, not notched at the apex; lower floret staminate, with a well-developed palea; pedicel of pedicellate spikelet lacking a lateral wing; inflorescence readily disarticulating ............................................ 77. Rhytachne (R. guianensis) Individual inflorescences of 1 raceme ..................................................... 15 Individual inflorescences of 2–9 racemes ............................................... 18 Pedicellate spikelets absent or reduced to < 2/3 the length of the sessile spikelets ............................................................................................... 16 Pedicellate spikelets present, > the length of the sessile spikelets ....... 17 Lower glume of sessile spikelet with corrugated, sculptured margins; inflorescence glabrous ................................. 77. Rhytachne (R. gonzalezii) Lower glume of sessile spikelet with smooth margins; inflorescence glabrous to pubescent ......................................................... 80. Schizachyrium Spikelets with glumes awned; callus hairs white; lower glume of pedicellate spikelet large and flattened; awn of upper lemma of sessile spikelet 3–5 cm long ............................................ 3. Andropogon (A. fastigiatus) Spikelets with glumes awnless; callus hairs coppery brown; lower glume of pedicellate spikelet not large and flattened; awn of upper lemma of sessile spikelet 6–10 cm long ............................................ 38. Heteropogon Sessile spikelet staminate or sterile, awnless, the pedicellate spikelet bisexual, awned .................................................................... 92. Trachypogon Sessile spikelet bisexual, awned, the pedicellate spikelet sterile or staminate, awnless ....................................................................................... 19 Lower floret of sessile spikelet staminate, with a well-developed palea ............................................................................................. 45. Ischaemum Lower floret of sessile spikelet sterile, the palea usually rudimentary or absent ................................................................................................... 20 Lower glume of sessile spikelet bicarinate, sulcate; rachis with white cilia; lower glume of pedicellate spikelet awned .................................... .................................................................... 3. Andropogon (A. angustatus) Lower glume of sessile spikelet convexly rounded on the back, not bicarinate or sulcate; rachis with golden brown cilia; lower glume of pedicellate spikelet awned ............................................... 41. Hyparrhenia

12

P OACEAE

Subkey 8 to the Genera of Panicoideae: Paniceae (also includes Isachne of Panicoideae: Isachneae) 1.

Spikelets borne in fascicles, each fascicle falling entire along with numerous attached bristles and/or flattened spines ....................................... 2 1. Spikelets not borne in fascicles that fall attached to numerous subtending bristles or spines .............................................................................. 3 2(1). Spikelet fascicles enclosed in a spiny bur, at least some of the spines conspicuously flattened and connate ..................................... 15. Cenchrus 2. Spikelet fascicles not enclosed in burs, subtended by bristles but not by flattened, connate spines .................................................... 69. Pennisetum 3(1). Spikelets falling attached to a branchlet that terminates in a sterile bristle; rare, forming mats in wet sand along rivers ......... 64. Paratheria 3. Spikelets not falling attached to a branchlet that terminates in a sterile bristle ..................................................................................................... 4 4(3). Spikelets subtended by 1–many bristles that remain attached to the branch after the spikelet falls ................................................... 81. Setaria 4. Spikelets not subtended by bristles (occasionally by hairs in some species of Axonopus) ........................................................................................... 5 5(4). Inflorescence a spicate panicle .................................................................. 6 5. Inflorescence an open panicle or panicle of racemes, not greatly contracted, rarely a solitary raceme bent downward .............................. 10 6(5). Spikelets borne in fascicles enclosed by indurate, ovate bracts with slightly recurved apices; rachis strongly zigzag .................. 4. Anthephora 6. Spikelets not borne in fascicles enclosed by indurate bracts; rachis straight ................................................................................................... 7 7(6). Spikelets placed alternately in 2 rows along the rachis, strongly laterally flattened; dry savannas ....................................................... 54. Mesosetum 7. Spikelets not placed alternately in 2 rows along the rachis, not or only weakly laterally flattened; generally in wet areas .............................. 8 8(7). Leaf blades 2–5.5 cm wide; plants (50–)100–300 cm tall .... 40. Hymenachne 8. Leaf blades 0.1–1 cm wide; plants 15–75 cm tall ..................................... 9 9(8). Inflorescence 0.5–2.5 cm long; spikelets with neither the glumes nor lemmas saccate at the base ......... 62. Panicum (P. caricoides, P. stenodes) 9. Inflorescence 3–40 cm long; spikelets with the upper glume and/or the lower lemma saccate at the base, or in 1 species with none of the bracts saccate .................................................................................. 79. Sacciolepis 10(5). Spikelets conspicuously awned ............................................................... 11 10. Spikelets awnless ..................................................................................... 14 11(10). Upper glume bifid, awned from between the teeth; foliage strongly viscid and aromatic .............................................................................. 52. Melinis 11. Upper glume not bifid, awned from the apex if an awn is present; foliage neither aromatic nor viscid ................................................................. 12 12(11). Raceme solitary, bent downward, the rachis prolonged past the last spikelet as a sterile bristle; spikelets tuberculate-ciliate .............. 26. Echinolaena 12. Racemes several to many, spreading to ascending, never bent downward or terminated by a sterile bristle; spikelets variously pubescent but never strongly tuberculate-ciliate ....................................................... 13

P OACEAE 13

13(12). Spikelets terete; leaf blades linear; lower lemma awnless or with a scabrous awn ...................................................................... 25. Echinochloa 13. Spikelets laterally flattened; leaf blades lanceolate; lower lemma with long, usually smooth and viscid (in 1 species scabrous) awn ................ ............................................................................................. 58. Oplismenus 14(10). Glumes similar, both nearly to as long as the spikelet ........... 39. Homolepis 14. Glumes unequal, the lower distinctly shorter than the spikelet ........... 15 15(14). Spikelets black at maturity .......................................................... 46. Lasiacis 15. Spikelets green at maturity .................................................................... 16 16(15). Lower glume represented by a bead-like or knob-like swelling at the base of the spikelet ......................................................................... 31. Eriochloa 16. Lower glume present or absent, but not reduced to a bead-like swelling at the base of the spikelet ........................................................................ 17 17(16). Base of spikelet prolonged downward to form a circular flange ................ ........................................................................................ 90. Streptostachys 17. Base of spikelet not toroidally prolonged into a downward-pointing flange .............................................................................................................. 18 18(17). Spikelet bracts with their apices beaked and laterally flattened; apex of upper palea with a pair of blisters .......................................... 1. Acroceras 18. Spikelet bracts with their apices not beaked and laterally flattened; apex of upper palea lacking a pair of blisters ............................................. 19 19(18). Rachises of racemes each ending in a sterile, pointed prolongation or bristle ................................................................................ 67. Paspalidium 19. Rachises of racemes not ending in a pointed bristle .............................. 20 20(19). Spikelets placed very obliquely on their pedicels, the upper floret appearing to fit into a pocket formed by the somewhat inflated lower glume; lower palea often strongly winged at maturity .................61. Otachyrium 20. Spikelets not placed obliquely on their pedicels, the upper floret not fitting into a pocket formed by the lower glume; lower palea never strongly winged .................................................................................... 21 21(20). Spikelets biconvex, with both florets hemispherical .............................. 22 21. Spikelets usually not biconvex, but if so then with both florets not hemispherical ............................................................................................... 23 22(21). Lower glume present, nearly as long as the spikelet; upper floret strawcolored .............................................. (Panicoideae: Isachneae) 44. Isachne 22. Lower glume absent; upper floret brown .......... 68. Paspalum (P. convexum) 23(21). Inflorescence of racemes, each raceme with the spikelets gradually reduced in size from the base of the rachis toward the apex; spikelets with glumes completely absent ...................................... 74. Reimarochloa 23. Inflorescence of racemes or not, but if racemes present then the spikelets never gradually reduced in size from the base of the rachis toward the apex; spikelets with at least the upper glume usually present ......... 24 24(23). Panicle open, the branches not strongly raceme-like ............................. 25 24. Panicle composed of racemes or raceme-like branches .......................... 33 25(24). Spikelets covered with elongate silky hairs ........................................... 26 25. Spikelets not covered with elongate silky hairs ..................................... 27 26(25). Lower glume absent; spikelet hairs tawny ..................... 49. Leptocoryphium 26. Lower glume present, ca. 1 mm long; spikelet hairs usually reddish, less

14

P OACEAE

commonly white ............................................................ 76. Rhynchelytrum 27(25). Upper lemmas with fleshy appendages, blisters, or scar-like depressions along their margins near the base; spikelets with glumes and lemmas usually strongly keeled, acuminate to attenuate .............. 42. Ichnanthus 27. Upper lemma without appendages, blisters, or scars along margins near the base; spikelets usually not strongly keeled, obtuse to acuminate .............................................................................................................. 28 28(27). Upper glume strongly saccate at the base ........... 79. Sacciolepis (S. striata) 28. Upper glume not saccate ......................................................................... 29 29(28). Upper lemma papery, the margins not enveloping the apex of the palea; robust aquatics; leaf blades 2–5.5 cm wide, cordate-based ................... ........................................................................................... 40. Hymenachne 29. Upper lemma indurate, the margins inrolled over the entire length of the edges of the palea; plants of various habitats, most with blades < 1.5 cm wide and not cordate-based .................................................. 30 30(29). Spikelets prominently cross-veined, at least toward the apex .................. .................................................... 13. Brachiaria (B. fasciculata, B. mollis) 30. Spikelets not cross-veined ....................................................................... 31 31(30). Lower palea developing enlarged keels or flanks at maturity; upper floret with compound papillae regularly disposed all over the lemma and palea ................................................................................... 86. Steinchisma 31. Lower palea not expanded at maturity; upper floret without compound papillae regularly disposed all over the lemma and palea ................ 32 32(31). Terminal inflorescences with chasmogamous spikelets; axillary inflorescences with cleistogamous spikelets; plants usually with foliar dimorphism .................................................................... 23. Dichanthelium 32. Terminal and axillary inflorescences with chasmogamous spikelets; plants without foliar dimorphism ........................................... 62. Panicum 33(24). Inflorescence a solitary, often arcuate raceme ....................................... 34 33. Inflorescence of 2–many racemes ............................................................ 35 34(33). Spikelets positioned so that the backs of the lower lemmas of successive spikelets face away from each other ..................................... 68. Paspalum 34. Spikelets positioned so that the backs of the lower lemmas of successive spikelets face toward each other .............................................. 91. Thrasya 35(33). Backs of lower lemmas turned toward the rachis .................................. 36 35. Backs of lower lemmas turned away from the rachis ............................ 37 36(35). Upper floret soft, flexible, the edges of the lemma thin and exposed, folded over the palea .............................................................. 24. Digitaria 36. Upper floret rigid, the edges of the lemma inrolled over the margins of the palea ................................................................................. 68. Paspalum 37(35). Lower glume absent or very small; upper floret smooth .......... 11. Axonopus 37. Lower glume well-developed; upper floret transversely rugulose ......... 38 38(37). Spikelets obtuse to acute, the veins glabrous to pubescent, but never hispid with curved hairs ...................................................... 13. Brachiaria 38. Spikelets acuminate to long-awned, the veins often hispid with curved hairs ................................................................................... 25. Echinochloa

Acroceras 15

1. ACROCERAS Stapf in Prain, Fl. Trop. Afr. 9: 621. 1920. [Subfamily Panicoideae, Tribe Paniceae] by Emmet J. Judziewicz Annuals or perennials, typically with sprawling, decumbent culms. Leaves with ligules membranous; blades linear to lanceolate. Inflorescence and open panicle with few, usually stiff and ascending branches. Spikelets obovate to elliptic, firmly membranous, laterally compressed, 2-flowered; bracts with small, laterally flattened apices; lower glume distinctly shorter than spikelet, 3–5-veined; internode between glumes conspicuous; upper glume and lower lemma as long as spikelet, 3– 5-veined, separated by a distinct internode; lower floret staminate or sterile, the palea well developed or not; upper floret slightly shorter than spikelet, indurate, pale, smooth to papillose, the lemma apiculate, with a depressed area on the back near the slightly narrowed and somewhat stipitate base, the margins thick, not strongly enclosing the edges of the palea, the palea flat, rigid, minutely bifid and beaked, its apex with a pair of scars or blisters. Tropical and subtropical areas worldwide; 22 species, 2 in Venezuela, both in the flora area. Key to the Species of Acroceras 1. 1.

Spikelets 2.8–3 mm long, the bracts all tipped with minute tufts of hairs, weakly beaked; leaf blades acute at base ............................. A. excavatum Spikelets 5–6 mm long, glabrous, the bracts strongly beaked; leaf blades subcordate to slightly clasping at base ................................. A. zizanioides

Acroceras excavatum (Henrard) Zuloaga & Morrone, Darwiniana 28: 195. 1987. —Panicum excavatum Henrard, Repert. Spec. Nov. Regni Veg. 23: 179. 1926. Perennial to 75 cm tall; leaf blades 15–20 × 1.5–2.5 cm; branches of inflorescence filiform, scabrous; spikelets strongly veined. Forest openings, 200–400 m; Bolívar (lower Río Paragua, near Upata). Southern Brazil, Bolivia, Paraguay, Argentina.

Acroceras zizanioides (Kunth) Dandy, J. Bot. 69: 54. 1931. —Panicum zizanioides Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 100. 1815 [1816]. Sprawling, decumbent, clone-forming perennial to 75 cm tall; leaf blades 8–20 × 1–3 cm; branches of inflorescence few, stout, scaberulous. Open stream sides, swamps, ditches, near sea level to 500 m; Delta Amacuro (common), Bolívar (Río Parguaza, Río Upata, Río Uruyen, Tumeremo), Amazonas (Río Negro, Río Orinoco, Río Yatúa). Anzoátegui, Carabobo, Distrito Federal, Guárico, Lara, Miranda, Portuguesa, Táchira, Zulia; Neotropics, Africa. ◆Fig. 1.

16

P OACEAE

Fig. 1. Acroceras zizanioides

Fig. 2. Aegopogon cenchroides

Andropogon 17

2. AEGOPOGON Humb. & Bonpl. ex Willd., Sp. Pl. 4: 899. 1806. [Subfamily Chloridoideae, Tribe Cynodonteae] by Emmet J. Judziewicz Weak, sprawling or stoloniferous herbs of indefinite duration. Leaves with ligules membranous; blades narrow. Inflorescence a terminal raceme, the rachis triquetrous and alternately bearing 2 rows of short, deciduous branches from 2 of the 3 sides; branches bearing a triad of spikelets, the central spikelet large, subsessile, bisexual, the lateral spikelets smaller, short-pedicellate, sterile or staminate. Central (bisexual) spikelet 1-flowered, with no extension of the rachilla; glumes equal, 1-veined, shorter than the floret, bifid, long-awned; lemma 3-veined, each vein extending into an awn, the central awn most prominent; palea slightly shorter than lemma, 2-keeled, the keels prolonged into awns. Lateral spikelets similar to central spikelet but florets smaller. Southwestern U.S.A., Mexico, Central America, Colombia, Venezuela, Ecuador, Peru, southern Brazil, Bolivia, Argentina; several poorly defined species, 1 in Venezuela. Aegopogon cenchroides Humb. & Bonpl. ex Willd., Sp. Pl. 4: 899. 1806. Plant straggling, from slender, rigid stolons, erect portions of culms 20–35 cm tall; inflorescence 4–6 cm long; spikelets 2.5–4 mm long, with central awn ca. 10 mm long.

Forest margins on tepui summits, 1800–2300 m; uncommon in Bolívar (Sororopán-tepui, Roraima-tepui). Widespread in Venezuelan Andes and Coastal Cordillera; Mexico, Central America, Colombia, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 2.

3. ANDROPOGON L., Sp. Pl. 1045. 1753. [Subfamily Panicoideae, Tribe Andropogoneae] Diectomis Kunth, Mém. Mus. Hist. Nat. 2: 69. 1815. Hypogynium Nees in Mart. et al., Fl. Bras. Enum. Pl. 2: 364. 1829. by Gerrit Davidse Perennials or annuals, usually cespitose, sometimes short-rhizomatous. Stems branched from the middle to upper nodes. Sheaths keeled or rounded; ligule a glabrous or ciliolate membrane; blades linear. Inflorescence 1 or more racemes, terminal and axillary, usually paired, sometimes solitary or when 3 or more digitate, often aggregated into complex compound panicles; peduncle spatheate with bladeless sheaths; racemes composed of numerous pairs of spikelets; disarticulation along the rachis between spikelet pairs; rachis internodes and pedicels filiform to clavate, often somewhat flattened, cupulate at the tip. Spikelets dimorphic, with 2 florets, dorsally or laterally compressed, paired, each pair with 1 sessile and 1 pedicellate spikelet. Sessile spikelets bisexual, dorsally compressed, narrowly lanceolate, acute, awned or awnless; callus obtuse; glumes subequal, membranous to chartaceous, completely concealing the florets; lower glume flattened or concave, with 2 submarginal keels, the margins inflexed over the margins of the upper glume; upper glume convex or keeled, with or without awn; lower floret sterile, the lower lemma hyaline, the lower palea absent; upper floret usually bisexual, sometimes pistillate; upper lemma hyaline, entire when awnless, or awned between 2 lobes; upper palea absent; lodicules 2; stamens 1 or 3; pedicellate spikelet rudimentary and sterile to enlarged and staminate, awned or awnless; pedicels free. Cosmopolitan in warm and tropical climates; ca. 100 species, ca. 18 in Venezuela, 13 of these in the flora area.

18

P OACEAE

Key to the Species of Andropogon 1. 1. 2(1). 2. 3(2).

3.

4(2).

4.

5(4). 5. 6(5).

6.

7(1). 7. 8(7). 8. 9(8). 9. 10(7). 10. 11(10). 11. 12(11). 12. 13(12).

Raceme 1 per peduncle .............................................................................. 2 Racemes 2 or more per peduncle ............................................................... 7 Sessile spikelets awnless ........................................................................... 3 Sessile spikelets awned ............................................................................. 4 Rachis internodes and pedicels pilose; racemes usually completely exserted from the subtending spathe; sessile spikelets bisexual or pistillate ..................................................................................... A. insolitus Rachis internodes and pedicels glabrous; racemes usually exserted < 1/4 the length of the subtending spathe; sessile spikelets pistillate .................................................................................................... A. virgatus Lower glume of pedicellate spikelet at least twice as long and wide as the lower glume of the sessile spikelet; the awn > 5 mm long; annuals ................................................................................................ A. fastigiatus Lower glumes of the pedicellate spikelet about as long as the lower glume of the sessile spikelet and slightly narrower, the awn mostly 0.5–2 mm long; perennials ..................................................................................... 5 Plants 20–65 cm tall; leaf blades of the middle of the culm 0.5–5 mm wide; ligules 1.5–1.7 mm long; anthers 1–1.5 mm long ........ A. diuturnus Plants 100–400 cm tall; leaf blades of the middle of the culm 10–20 mm wide; ligules 2–5 mm long; anthers 1.5–2.5 mm long .......................... 6 Racemes borne on peduncles long-exserted from the subtending spathes; spathes narrow, never enclosing the peduncle at maturity ............................................................................................A. longiramosus Racemes mostly borne on peduncles not fully exserted from the subtending spathes; spathes broader, usually enclosing the base of the raceme or the raceme laterally exserted .................................................... A. vetus Sessile spikelets awnless ........................................................................... 8 Sessile spikelets awned ........................................................................... 10 Plants 1–2 m tall; inflorescences densely aggregated at the top of the stem forming a large broom-like mass ...................................... A. bicornis Plants usually < 1 m tall; inflorescences diffusely arranged in the upper leaf of the stem....................................................................................... 9 Leaf blades 1–3 mm wide, sessile spikelets 2.7–4 mm long, acute to acuminate .......................................................................... A. leucostachyus Leaf blades 3–6.5 mm wide; sessile spikelets 3.5–4.5 mm long, shortly acute ......................................................................................... A. selloanus Annuals; upper glume of sessile floret awned 4–10 mm ..........A. angustatus Perennials; upper glume of sessile floret awnless .................................. 11 Leaf blades 10–20 mm wide; plants 100–250 cm tall; inflorescences > 15 per culm ......................................................................... A. indetonsus Leaf blades < 9 mm wide; plants 30–100 cm tall; inflorescences > 1– 5(–10) per culm .................................................................................... 12 Racemes 4–16 per peduncle ...................................................... A. hypogynus Racemes 2 or 3 per peduncle ................................................................... 13 Pedicellate spikelets enlarged, with staminate flowers; apex of rachis segment shallowly and evenly cupulate ................................. A. carinatus

Andropogon 19

13.

Pedicellate spikelets narrow, with staminate flowers; apex of rachis segment deeply and obliquely, lacerate cupulate ....................... A. diuturnus

Andropogon angustatus (J. Presl) Steud., Syn. Pl. Glumac. 1: 370. 1855 [1854]. —Diectomis angustata J. Presl in C. Presl, Reliq. Haenk. 1: 333. 1830. Diectomis laxa Nees in Mart. et al., Fl. Bras. Enum. Pl. 2: 340. 1829, non Andropogon laxus Willd. 1806. Erect annual to ca. 85 cm tall; lower glume of pedicellate spikelets narrow. Disturbed areas in savannas, near sea level to 500 m; widespread in Delta Amacuro, Bolívar, and Amazonas. Anzoátegui, Aragua, Barinas, Cojedes, Distrito Federal, Guárico, Lara; common in Mexico, Guatemala, Honduras, El Salvador, Nicaragua, Costa Rica, Panama, Caribbean (Cuba), Colombia, Guyana, Suriname, Brazil, Bolivia. ◆Fig. 4. Andropogon bicornis L., Sp. Pl. 1046. 1753. Robust annual to 100–150 cm tall; inflorescences dense, prominent, whitish hairy. Open, herbaceous, secondary vegetation, near sea level to 2300 m; widespread weed in Delta Amacuro, Bolívar, and Amazonas. Anzoátegui, Apure, Aragua, Distrito Federal, Falcón, Guárico, Lara, Mérida, Miranda, Monagas, Nueva Esparta, Portuguesa, Sucre, Táchira, Trujillo, Yaracuy, Zulia; Mexico, Central America, Caribbean, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Chile, Paraguay, Argentina; 2 varieties, both in the flora area. Key to the Varieties of A. bicornis 1. Pedicellate spikelets with 1 or 2 of the terminal ones of each raceme conspicuously larger than the rest ....... var. bicornis 1. Pedicellate spikelets all enlarged and of approximately the same size .................. ..................................... var. burchellii A. bicornis var. bicornis All except the uppermost of the pedicellate spikelets rudimentary and never bearing staminate flowers. Open, herbaceous, secondary vegetation, near sea level to 2300 m; widespread weed in Delta Amacuro, Bolívar, and Amazonas. Anzoátegui, Apure, Aragua, Distrito Federal, Falcón, Guárico,

Lara, Mérida, Miranda, Monagas, Nueva Esparta, Portuguesa, Sucre, Táchira, Trujillo, Yaracuy, Zulia; Mexico, Central America, Caribbean, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Chile, Paraguay, Argentina. A. bicornis var. burchellii Hack. in Mart., Fl. Bras. 2(4): 285. 1883. All or most of the pedicellate spikelets enlarged and bearing staminate flowers. Savannas, 100–200 m; Amazonas (Río Parucito). Barinas, Monagas, Sucre; Colombia, Brazil. Andropogon carinatus Nees in Mart. et al., Fl. Bras. Enum. Pl. 2: 330. 1829. Strongly cespitose perennial, mostly < 60 cm tall; inflorescence usually 1 per culm. Savannas, 100–200 m; Amazonas (Rincones de Chacorro). Brazil, Bolivia. Andropogon diuturnus Sohns, Mem. New York Bot. Gard. 9(3): 406, fig. 77. 1957. Perennial 20–90 cm tall; racemes 1 or 2 per peduncle. Open areas in rock, herbaceous or shrubby vegetation, 1200–2100 m; Bolívar (Cerro Jaua, Macizo del Chimantá [Chimantá-tepui]), Amazonas (Cerro Duida, Cerro Guanay, Cerro Marahuaka, Cerro Yaví, Cerro Yutajé). Endemic. ◆Fig. 5. Andropogon fastigiatus Sw., Prodr. 1788. —Diectomis fastigiata (Sw.) P. Beauv., Ess. Agrostogr. 132, 160. 1812. Erect annual to 150 cm tall; lower glume of pedicellate spikelets broad and prominent; plants often quite noticeable shortly after fire. Savannas, often in disturbed areas, near sea level to 600 m; widespread in Bolívar and Amazonas. Anzoátegui, Apure, Aragua, Barinas, Carabobo, Guárico, Lara, Sucre, Zulia; Mexico, Guatemala, Honduras, El Salvador, Nicaragua, Costa Rica, Panama, Caribbean, Colombia, Guyana, Brazil, Bolivia, Old World tropics. ◆Fig. 3. Andropogon hypogynus Hack. in Mart., Fl. Bras. 2(4): 290, pl. 66. 1883. Perennial 0.6–2 m tall; sheaths markedly distichous and usually > 5 racemes per pe-

20

P OACEAE

duncle. Marshy areas in savannas, 50–500 m; Bolívar (Cerro Ichún), Amazonas (Puerto Ayacucho). Apure, Barinas, Guárico, Portuguesa; Guatemala, Colombia, Brazil, Bolivia, Paraguay, Argentina, Uruguay. ◆Fig. 7. Andropogon indetonsus Sohns, Mem. New York Bot. Gard. 9(3): 269. 1957. Perennial 1–2.5 m tall, with broad blades and numerous inflorescences in compound panicle, each inflorescence with 2, sometimes 3, racemes per peduncle. Boulder-strewn, open shrubby formations, openings in low, open forests, savannas with Pteridium, especially near escarpments, 700–1900 m; Bolívar (Auyán-tepui, Cerro Guaiquinima, Gran Sabana, Río Uaiparú, Uaipán-tepui). Brazil (Roraima: Serra Tepequem). Andropogon insolitus Sohns, Mem. New York Bot. Gard. 9(3): 271, fig. 9. 1957. Andropogon crucianus Renvoize, Gram. Bolivia 596, fig. 142. 1998, syn. nov. Perennial to 1 m tall; inflorescences usually dense, large, compound, with solitary, pilose racemes and elongated peduncles extending the spikelets beyond the spathes. Mauritia palm swamps and marshy areas in savannas, 50–200 m; Bolívar (near Ciudad Piar). Apure; Brazil (Bahia), Bolivia. Occasionally some peduncles in an inflorescence bear a short, second raceme (Davidse & González 15710, MO, from Apure). Andropogon insolitus is closely related to A. virgatus, and one intermediate collection (Huber & Entralgo 7401, MO) from the Gran Sabana is known. These intermediate plants have the pilose callus as in A. insolitis, but lack the hairs on the rachis segments and pedicels as in A. virgatus. The pilose callus is also the character that distinguishes the closely related, African A. festuciformis Rendle from A. virgatus, so there is a chance that Huber & Entralgo 7401 represents an introduction from Africa, although the remote location argues against this. Andropogon leucostachyus Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 187. 1815 [1816]. Perennial to 1 m tall; inflorescences white-hairy. Disturbed areas in savannas, roadsides, 50–200 m; Delta Amacuro (expected), widespread in Bolívar and Ama-

zonas. Anzoátegui, Apure, Aragua, Barinas, Distrito Federal, Falcón, Guárico, Lara, Mérida, Miranda, Monagas, Nueva Esparta, Portuguesa, Táchira, Yaracuy, Zulia; Canada, Mexico, U.S.A., Central America, Caribbean, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Chile, Paraguay, Argentina, Uruguay. ◆Fig. 6. Andropogon longiramosus Sohns, Mem. New York Bot. Gard. 9(3): 272, fig. 10. 1957. Perennial ca. 2 m tall; lower leaves with long pseudopetioles, the blades broad; racemes solitary, borne on remarkably long-exserted peduncles. Open areas on dry tepui slopes, 900–1100 m; Amazonas (Cerro Guanay). Endemic. Andropogon longiramosus, so far only known from the type collection, is closely related to A. vetus in the broad sense adopted in this treatment. Besides the very long peduncles and narrow spathes, A. longiramosus seems to have modally longer racemes and rachis segments. A specimen from Serra Aracá, Amazonas, Brazil, is similar to A. longiramosus in all respects except for its possession of two racemes per peduncle. Andropogon selloanus (Hack.) Hack., Bull Herb. Boissier, sér. 2, 4: 266. 1904. —Andropogon leucostachyus subsp. selloanus Hack. in A. DC. & C. DC., Monogr. Phan. 6: 420. 1889. Similar to Andropogon leucostachyus but coarser in all of its parts. Disturbed areas in savannas, roadsides, clearings, 50–2000 m; Delta Amacuro (expected), widespread in Bolívar and Amazonas. Anzoátegui, Apure, Aragua, Barinas, Carabobo, Cojedes, Distrito Federal, Falcón, Guárico, Lara, Mérida, Monagas, Portuguesa, Sucre, Yaracuy, Zulia; Mexico, Central America, Caribbean, Colombia, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia, Paraguay, Argentina, Uruguay. Andropogon vetus Sohns, Mem. New York Bot. Gard. 9(3): 277, fig. 12. 1957. Andropogon crassus Sohns, Mem. New York Bot. Gard. 9(3): 271, fig. 11. 1957, syn. nov.

Andropogon 21

Fig. 3. Andropogon fastigiatus

Fig. 4. Andropogon angustatus

22

P OACEAE

Fig. 5. Andropogon diuturnus

Fig. 6. Andropogon leucostachyus

Andropogon 23

Fig. 7. Andropogon hypogynus

24

P OACEAE

Fig. 8. Andropogon vetus

Andropogon 25

Fig. 9. Andropogon virgatus

26

P OACEAE

Andropogon perdignus Sohns, Mem. New York Bot. Gard. 9(3): 274, fig. 10. 1957, syn. nov. Robust perennial 1–4 m tall; blades broad; racemes solitary, white-pilose, partially exserted from the broad spathes. Rocky open escarpments, open woodlands and thickets, 500–1400 m; Bolívar (Cerro Guiaquinima, Macizo del Chimantá [Chimantá-tepui]), Amazonas (Cerro Duida, Cerro Guanay, Cerro Yapacana, Cerro Yutajé, Sierra de la Neblina). Colombia (Caquetá, Sierra de Chiribiquete). ◆Fig. 8. Andropogon vetus is a very characteristic species with its numerous inflorescences of solitary racemes and the lower leaves with prominent pseudopetioles but broad blades. These pseudopetioles are not developed on the leaves arising from the middle and upper nodes. Andropogon crassus was based on specimens in which these pseudopetioles were particularly well developed, but the over-mature inflorescence racemes and spikelets are similar to those of A. vetus. Andropogon perdignus was originally distinguished by its tendency to have its slightly shorter racemes less exserted from the subtending spathes, but with the additional collections now available, this does not seem to be a useful distinction. Included here are also some unusually shorter

plants (Liesner & Holst 21483, MO; Davidse & Miller 27345, MO) that look distinctive because they have relatively narrow cauline leaf blades but numerous pseudopetiolate basal leaves. Andropogon virgatus Desv. in W. Ham., Prodr. Pl. Ind. Occid. 9. 1825. —Hypogynium virgatum (Desv.) Dandy, J. Bot. 69: 54. 1931. Perennial to 1 m tall; inflorescences typically dense, large, compact, compound, the solitary racemes glabrous and mostly only a little longer than the spathes. Mauritia palm swamps, marshy areas in savannas, 50–1200 m; Delta Amacuro (expected), widespread in Bolívar and Amazonas. Anzoátegui, Apure, Guárico; Mexico, Guatemala, Belize, Honduras, Nicaragua, Costa Rica, Caribbean, Panama, Colombia, Guyana, Suriname, French Guiana, Brazil, Bolivia, Paraguay, Argentina, Uruguay. ◆Fig. 9. The compound inflorescence is usually elongated but is sometimes contracted into a short, tightly compact, nearly spheroidal shape. Atypical plants with extremely slender inflorescences with widely spaced racemes (Davidse et al. 17471, MO; Davidse et al. 17274, MO) are known from the seasonally flooded white-sand savannas at the base of Cerro Yapacana in Amazonas state).

4. ANTHEPHORA Schreb., Beschr. Gräs. 2: 105. 1779. [Subfamily Panicoideae, Tribe Paniceae] by Emmet J. Judziewicz Cespitose annuals. Leaves with ligules membranous, the blades linear. Inflorescence a densely spicate panicle of deciduous fascicles on a zigzag rachis; fascicles stipitate, of 4 indurate, broadly lanceolate to ovate bracts surrounding and concealing 2–4 spikelets, several of these rudimentary. Spikelets lanceolate, 2-flowered; lower glume absent; upper glume about as long as spikelet, needle-like, 1-veined; lower floret as long as spikelet, sterile, the lemma hyaline, 3–5-veined; upper floret about as long as spikelet, thinly coriaceous, the lemma faintly 3–5-veined, its margins loosely enclosing the edges of the palea. Neotropics, Africa, southwestern Asia; ca. 13 species, 1 in Venezuela. Anthephora hermaphrodita (L.) Kuntze, Revis. Gen. Pl. 2: 759. 1891. —Tripsacum hermaphroditum L., Syst. Nat. ed. 10, 1261. 1759. Cespitose annual ca. 30 cm tall; inflores-

cence 3–5 cm long; spikelet fascicles 4.5–6 mm long. Dry, open ground, ca. 100 m; Bolívar (Cerro Picacho, Moitaco, Río Aro, Santa María del Vapor). Widespread in dry, coastal Venezuela; Neotropics. ◆Fig. 10.

Anthephora 27

Fig. 10. Anthephora hermaphrodita

28

P OACEAE

5. ARBERELLA Soderstr. & C.E. Calderón, Brittonia 31: 433. 1979. [Subfamily Bambusoideae, Tribe Olyreae] by Emmet J. Judziewicz Monoecious, cespitose perennials. Leaves forming complements; ligules short, membranous; pseudopetioles short, turgid; blades linear to elliptic, usually somewhat asymmetrical. Inflorescences borne at all nodes or absent from middle nodes, loosely paniculate, few-flowered, inconspicuous; individual inflorescences each bearing a single terminal pistillate spikelet on a clavate pedicel, and several pedicellate staminate spikelets below. Spikelets one-flowered, unisexual. Pistillate spikelets much larger than the staminate ones; glumes lanceolate to ovate, subequal, whitish, strongly 5–9-veined, acuminate to short-aristate; rachilla internode between glumes and floret present or not; floret shorter than the glumes, ellipsoid, indurate, becoming mottled with dark spots when mature, the lemma with margins overlapping and partially concealing the palea, glabrous on the back, the margins, especially near the base, with a dense, matted beard of crispy hairs; stigmas 2, plumose. Staminate spikelets lanceolate, membranous, lacking glumes, the lemma short-aristate, 3–5-veined, slightly exceeding the palea; stamens 3. Costa Rica, Panama, Colombia, Venezuela, Suriname, French Guiana, Amazonian Brazil; 7 species, 2 in Venezuela, both in the flora area. Key to the Species of Arberella 1. 1.

Leaf blades 5–6 × 1.7–2.3 cm ................................................ A. aff. bahiensis Leaf blades 8–10.5 × 2.3–3.5 cm ............................................... A. venezuelae

Arberella aff. bahiensis Soderstr. & Zuloaga, Brittonia 37: 23. 1985. Cespitose perennial 30–50 cm tall; pistillate spikelets (8–)10–12 mm long, the floret 5–5.5 mm long. Forests, ca. 100 m; Amazonas (Terecay). Brazil (Bahia). Material from the flora area appears to be best referred to this Brazilian species.

Arberella venezuelae Judz. & Davidse, Novon 1: 76. 1991. Cespitose perennial 40–60 cm tall; pistillate spikelets (9–)11–12 mm long, the floret 5–6 mm long. Forests, ca. 100 m; Amazonas (Río Orinoco, Río Ventuari). Costa Rica, Panama, Colombia, Suriname, French Guiana, Amazonian Brazil. ◆Fig. 11.

6. ARISTIDA L., Sp. Pl. 82. 1753. [Subfamily Arundinoideae, Tribe Aristideae] Streptachne R. Br., Prodr. 174. 1810. Arthratherum P. Beauv., Ess. Agrostogr. 32, 152. 1812. Chaetaria P. Beauv., Ess. Agrostogr. 30, 158. 1812. Aristopsis Catasus, Folia Geobot. Phytotax. 16(4): 439. 1981. by Gerrit Davidse Annuals or mostly perennials, cespitose. Culms cylindrical or compressed, usually solid. Ligule a minute, ciliolate membrane; leaf blades linear, flat, folded, or involute. Inflorescence a solitary, terminal panicle. Spikelets terete, pedicellate, with 1 floret; disarticulation above the glumes; glumes often as long as the lemma, narrow, keeled, membranous, equal or unequal, 1–3(–5)-veined, entire or emarginate, muticous or shortly awned, the lower usually disarticulating much earlier than the upper; lemma terete, convolute or involute, rigid, sometimes terminating in a

Arberella 29

Fig. 11. Arberella venezuelae

30

P OACEAE

narrow, straight or twisted column, the column articulated or not, the margins overlapping the palea; callus acuminate, pilose; awns usually 3, terminal, equal or the lateral shorter, rarely the 2 lateral ones rudimentary or absent; palea much smaller than the lemma, hyaline or membranous; lodicules 2; stamens 1 or 3; styles 2. Caryopsis cylindrical or sulcate; hilum linear; embryo 1/3–1/2 as long as the caryopsis. Cosmopolitan in temperate and tropical climates; ca. 250 species, ca. 15 in Venezuela, 10 of these in the flora area. Key to the Species of Aristida 1. 1. 2(1). 2. 3(2). 3. 4(3). 4. 5(4). 5. 6(5). 6.

7(4). 7. 8(7). 8. 9(8). 9.

Base of callus bifid ........................................................................... A. riparia Base of callus a single point ...................................................................... 2 Ventral side of lemma grooved ....................................................... A. gibbosa Ventral side of lemma rounded ................................................................. 3 Lateral awns minute or rudimentary ............................................ A. ternipes Lateral awns sometimes shorter than the central awn but clearly evident ................................................................................................................ 4 Column not evident .................................................................................... 5 Column clearly evident between the apex of the lemma and the base of the three awns ....................................................................................... 7 Annuals ................................................................................... A. adscensionis Perennials .................................................................................................. 6 Panicle diffuse, the branches widely spreading; central and lateral awns all ± straight or slightly divergent .......................................... A. longifolia Panicle compact, the branches narrowly ascending to slightly divergent; central awn curved, at maturity nearly curved into a semicircle, the lateral awns ± straight or slightly divergent ................................. A. torta Apex of the column with a point of disarticulation, the 3 awns deciduous at maturity .................................................................................. A. setifolia Apex of the column without a point of disarticulation, the 3 awns never deciduous ................................................................................................ 8 Lower glume 1.8–2.5 mm long; plants 10–25 cm tall; awns 5–8 mm long ................................................................................................. A. capillacea Lower glume > 3.5 mm long; plants 30–85 cm tall; awns 9–16 mm long ................................................................................................................ 9 Awns straight at the base, sometimes with a single twist at spikelet maturity; leaf blades not long-persistent, mostly stiffly erect ....... A. moritzii Awns contorted at the base; leaf blades persisting, curling up in age .................................................................................................. A. recurvata

Aristida adscensionis L., Sp. Pl. 82. 1753. —Chaetaria adscensionis (L.) P. Beauv., Ess. Agrostogr. 30, 151, 158. 1812. Aristida bromoides Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 122. 1815 [1816]. —Chaetaria bromoides (Kunth) Roem. & Schult., Syst. Veg. 2: 396. 1817. —Aristida adscensionis var. bromoides (Kunth) Henrard, Meded. Rijks-Herb. 54: 62.

1926. —Aristida adscensionis subsp. bromoides (Kunth) Henrard, Meded. Rijks-Herb. 58(A): 322, t. 159. 1932. Aristida coarctata Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 122. 1815 [1816]. —Chaetaria coarctata (Kunth) Roem. & Schult., Syst. Veg. 2: 396. 1817. —Aristida adscensionis var. coarctata (Kunth) Kuntze, Revis. Gen. Pl. 33: 340. 1898.

Aristida 31

Aristida humilis Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 121. 1815 [1816]. —Chaetaria humilis (Kunth) Roem. & Schult., Syst. Veg. 2: 396. 1817. —Aristida dispersa var. humilis (Kunth) Trin. & Rupr., Sp. Gram. Stipac. 129. 1842. Weedy annual 10–80 cm tall. Roadsides and other disturbed areas in dry, herbaceous vegetation, 50–1300 m; Bolívar (Ciudad Bolívar, Upata), Amazonas (La Esmeralda). Anzoátegui, Aragua, Carabobo, Distrito Federal, Falcón, Lara, Mérida, Táchira, Trujillo, Zulia; U.S.A., Mexico, Guatemala, Honduras, Nicaragua, Caribbean, Colombia, Ecuador, Peru, Brazil, Bolivia, Chile, Paraguay, Argentina. Aristida capillacea Lam., Tabl. Encycl. 1: 156. 1791. —Chaetaria capillacea (Lam.) P. Beauv., Ess. Agrostogr. 30, 158, t. 8, fig. 5. 1812. Aristida elegans Rudge, Pl. Guian. 22, t. 30. 1805. Delicate annual to 25 cm tall with spreading inflorescence branches. Disturbed areas in savannas, 50–1200 m; widespread in Bolívar and Amazonas. Anzoátegui, Apure, Aragua, Cojedes, Guárico, Monagas, Portuguesa, Sucre, Táchira; Mexico, Guatemala, Belize, Honduras, Nicaragua, Costa Rica, Panama, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Argentina. ◆Fig. 14. Aristida gibbosa (Nees) Kunth, Enum. Pl. 1: 189. 1833. —Chaetaria gibbosa Nees in Mart. et al., Fl. Bras. Enum. Pl. 2: 383. 1829. Aristida marginalis Ekman, Ark. Bot. 10(17): 23, t. 3, fig. 2, t. 6, fig. 12. 1911. Aristida orizabensis E. Fourn., Mexic. Pl. 2: 78. 1886. Perennial 50–95 cm tall; inflorescence narrow but slightly open. Savannas, 50–1200 m; Bolívar (Auyán-tepui, between Caicara and Ciudad Bolívar, Roraima-tepui). Anzoátegui, Aragua, Sucre; Mexico, Honduras, Costa Rica, Panama, Caribbean, Colombia, Guyana, Brazil, Bolivia. Aristida longifolia Trin., Mém. Acad. Imp. Sci. St. Pétersbourg, Sér. 6, Sci. Math. 1(1): 84. 1830.

Cespitose perennial to 100 cm; inflorescence branches spreading; culm bases often with a distinctive purple coloration. Savannas, 50–1000 m; widespread in Bolívar and Amazonas. Anzoátegui, Apure, Carabobo, Guárico, Lara, Zulia; Mexico, Belize, Nicaragua, Colombia, Guyana, Suriname, French Guiana, Brazil, Bolivia. ◆Fig. 18. Aristida moritzii Henrard, Meded. RijksHerb. 54: 133. 1926. Aristida pittieri Henrard, Meded. RijksHerb. 54(A): 447, fig. 1927, syn. nov. Weak perennial to 60 cm tall. Weedy areas and roadsides in savannas and open areas in shrubby, secondary growth, 50–500(–2000) m; common in Bolívar. Colombia. ◆Fig. 17. Aristida recurvata Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 123. 1815 [1816]. —Chaetaria recurvata (Kunth) Roem. & Schult., Syst. Veg. 2: 397. 1817. Aristida neesiana Trin. & Rupr., Sp. Gram. Stipac. 113. 1842. Aristida riedeliana Trin. & Rupr., Sp. Gram. Stipac. 113. 1842. Densely tufted perennial to 85 cm; older leaf blades curling. Savannas, 400–1400 m; widespread in Delta Amacuro, Bolívar, and Amazonas. Anzoátegui, Aragua, Distrito Federal, Falcón, Guárico, Lara, Miranda, Monagas; Belize, Honduras, Costa Rica, Panama, Colombia, Guyana, Suriname, Peru, Brazil, Bolivia. ◆Fig. 15. Aristida riparia Trin., Mém. Acad. Imp. Sci. St.-Pétersbourg, Sér. 6, Sci. Math., Seconde Pt. Sci. Nat. 4, 2(1): 48. 1836. Aristida implexa var. aequa Trin. & Rupr., Sp. Gram. Stipac. 124. 1842. Aristida planifolia Swallen, Ann. Missouri Bot. Gard. 30: 145. 1943. Perennial to 85 cm tall; inflorescences large, spike-like, with appressed branches. Savannas, near sea level to 1200 m, widespread in Delta Amacuro, Bolívar, and Amazonas. Anzoátegui, Guárico, Monagas; Panama, Guyana, Suriname, French Guiana, Brazil, Bolivia, Paraguay, Argentina. ◆Fig. 12. Aristida setifolia Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 122. 1815 [1816].

32

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Fig. 12. Aristida riparia

Fig. 14. Aristida capillacea

Fig. 13. Aristida torta

Aristida 33

Fig. 15. Aristida recurvata

Fig. 16. Aristida setifolia

Fig. 17. Aristida moritzii

34

P OACEAE

Fig. 18. Aristida longifolia

Arthrostylidium 35

—Chaetaria setifolia (Kunth) Nees in Mart. et al., Fl. Bras. Enum. Pl. 2: 381. 1829. Aristida arenaria Trin., Gram. Panic. 25. 1826. —Aristida setifolia var. arenaria (Trin.) Trin. & Rupr., Sp. Gram. Stipac. 126. 1842. Aristida setifolia var. grandiflora Trin. & Rupr., Sp. Gram. Stipac. 127. 1842. Aristida setifolia var. intermedia Trin. & Rupr., Sp. Gram. Stipac. 127. 1842. Aristida tarapotana Mez, Repert. Spec. Nov. Regni Veg. 17(8–12): 151. 1921. Aristida doelliana Henrard, Meded. RijksHerb. 54: 154, 161, fig. s.n. 1926. Annual to 70 cm tall; leaves stiff, involute; awns deciduous. Roadsides and other disturbed areas in savannas, 50–1000 m; very common in Bolívar. Anzoátegui, Aragua, Distrito Federal, Guárico, Lara, Miranda, Sucre, Táchira, Yaracuy; Mexico, Belize, Honduras, Colombia, Guyana, Suriname, Peru, Brazil. ◆Fig. 16. Aristida ternipes Cav., Icon. 5: 46. 1799. Streptachne scabra Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 124, t. 40. 1815 [1816]. —Aristida scabra (Kunth)

Kunth, Révis. Gramin. 1: 62. 1829. Streptachne tenuis Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 124–125. 1815 [1816]. —Aristida tenuis (Kunth) Kunth, Révis. Gramin. 1: 62. 1829. Erect perennials. Disturbed areas in savannas, 50–1000 m; apparently very rare in Bolívar (Gran Sabana). Guárico, Zulia; U.S.A., Mexico, Central America, Caribbean, Colombia, Ecuador, Bolivia. Aristida ternipes is known only from a single specimen from the flora area, and it has not been verified. Aristida torta (Nees) Kunth, Enum. Pl. 1: 190. 1833. —Chaetaria torta Nees in Mart. et al., Fl. Bras. Enum. Pl. 2: 386. 1829. Aristida tincta Trin. & Rupr., Sp. Gram. Stipac. 3: 111. 1842. Perennial 50–100 cm tall; leaves stiff; panicle branches loosely erect. Moist areas in savannas, 50–1300 m; very common in Bolívar and Amazonas. Apure, Guárico, Lara; Mexico, Belize, Honduras, Nicaragua, Costa Rica, Panama, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 13.

7. ARTHROSTYLIDIUM Rupr., Bambuseae 27. 1839 [preprint]; Mém. Acad. Imp. Sci. St.-Pétersbourg, Sér. 6, Sci. Math., Seconde Pt. Sci. Nat. 5(2): 117. 1840. [Subfamily Bambusoideae, Tribe Bambuseae] by Emmet J. Judziewicz Small to medium-sized, cespitose, unarmed, woody bamboos, generally scandent or clambering. Rhizomes short, determinate. Culms cylindrical, hollow with a small to large lumen, the midculm nodes with a single primary branch bud, this when developing producing 3–many branches at the summit of a promontory. Culm leaves with blades erect; foliage leaves with both inner and outer ligules present; oral setae inconspicuous to prominent; blades linear to lanceolate or less commonly ovate or elliptic, the midvein not prominent. Inflorescences typically determinate, spicate racemes lacking bracts, the rachis straight or zigzag. Spikelets subsessile, bisexual; glumes 1 or 2, about half as long as the spikelet, persistent, papery; lowermost floret usually sterile; functional florets 1–several, at maturity disarticulating and falling attached to the prominent rachilla internode, the spikelets terminating in 1–several successively reduced and sterile florets; lemma and palea subequal, the margins of the lemma clasping the rachilla internode; stamens 3; stigmas 2. Southern Mexico, Central America, West Indies, tropical South America (not eastern Brazil); ca. 30 species, ca. 12 in Venezuela, 8 of these in the flora area.

36

P OACEAE

The four unidentified species (A–D) are based on collections that are too fragmentary to either refer to previously described species or describe as new species. Key to the Species of Arthrostylidium 1.

1.

2(1). 2. 3(2). 3. 4(3). 4. 5(4). 5. 6(5). 6. 7(5). 7.

Basal internode of culm 2–4 m long, much longer than the following internode which is only 1–2 cm long; foliage leaf blades 17–20 times as long as wide; spikelets 40–70 mm long, the lemmas with awns 3– 10 mm long; Amazonas (Cerro Marahuaka, Sierra de Maigualida) ........................................................................................... A. schomburgkii Basal internode of culm < 1 m long, about as long as succeeding internodes; foliage leaf blades 4–13 times as long as wide; spikelets 10– 30 mm long, the lemmas awnless or mucronate .................................. 2 Midculm nodes encircled by prominent, corky, reflexed, skirt-like outgrowths ...................................................................................... A. scandens Midculm nodes lacking skirt-like outgrowths .......................................... 3 Foliage leaf sheaths densely papillose-hispid on back, their summits with numerous, prominent oral setae ..................................................... A. sp. A Foliage leaf sheaths glabrous on back, their summits with prominent oral setae or not ............................................................................................. 4 Foliage leaf blades ca. 20 × 2–2.5 cm, cuneate basally; Amazonas (Sierra Parima)............................................................................................. A. sp. B Foliage leaf blades 3.5–16 × 0.5–2.2 cm, rounded basally ....................... 5 Culms smooth; inflorescence flexuous to strongly zigzag ........................ 6 Culms scabrous, at least below the nodes; inflorescence (as far as known) straight ................................................................................................... 7 Foliage leaf blades 80–140 × 7–15 mm; inflorescence strongly zigzag; 1000–1900 m .......................................................................... A. venezuelae Foliage leaf blades 35–45 × 5–6 mm; inflorescence merely flexuous; 100– 300 m ................................................................................................ A. sp. C Branches at midculm nodes 10–40; eastern Bolívar, 600–1500 m ............ ................................................................................................. A. pubescens Branches at midculm nodes 2–4; western Amazonas, ca. 100 m ...... A. sp. D

Arthrostylidium pubescens Rupr., Bambuseae. 29. 1839 [preprint]; Mém. Acad. Imp. Sci. St.-Pétersbourg, Sér. 6, Sci. Math., Secunde Pt. Sci. Nat. 5(2): 119, pl. 4, fig. 14. 1840. Scandent bamboo to 5 m long, 1 cm diameter; foliage leaf blades 8–16 × 1–2.2 cm; inflorescence 10–25 cm long; spikelets 1.8–2.5 mm long, the florets 5–9 mm long, awnless. Montane forests, 600–1500 m; Bolívar (Altiplanicie de Nuria, Amaruay-tepui, Gran Sabana, Ptari-tepui, headwaters of Río Apongao). Aragua, Distrito Federal, Falcón, Mérida, Miranda, Táchira, Trujillo; Costa Rica, Panama, Colombia, Trinidad-Tobago.

1964. —De-beu-ni. Arthrostylidium cacuminis McClure, Mem. New York Bot. Gard. 10(5): 3. 1964. Thicket-forming, scandent bamboo; culms to 5 m long, weak; midculm nodes bearing 2 or 3 main branches; foliage leaf blades 9–20 × 1.7–4.5 cm; inflorescence 15–30 cm long; spikelets 20–25 mm long, 3–5-flowered, the lemmas 7–11 mm long, awnless. Steep forested slopes, elfin forests, stream banks, near waterfalls, 500–1600 m; Bolívar (Gran Sabana, Sierra Ichún), Amazonas (Cerro Aracamuni, Cerro Huachamacari, Cerro Marahuaka, Sierra de la Neblina). Guyana, Suriname.

Arthrostylidium scandens McClure, Mem. New York Bot. Gard. 10(5): 4.

Arthrostylidium schomburgkii (Benn.) Munro, Trans. Linn. Soc. London 26: 41.

Arthrostylidium 37

Fig. 19. Arthrostylidium schomburgkii

38

P OACEAE

1868. —Arundinaria schomburgkii Benn., Trans. Linn. Soc. London 18: 562. 1841. —Cura, Curata. Loosely cespitose bamboo 12–15 m long with straight, smooth, weak, hollow culms 1– 3.5 cm diameter; branches ca. 50–100 at each node; foliage leaf blades 17–22 × 1–1.2 cm; spikelets stout, 7–9-flowered. Cloud forests on talus slopes of tepuis, 1500–1700 m; Amazonas (Cerro Marahuaka, Sierra de Maigualida). Endemic. ◆Fig. 19. The very long lowest culm internodes of Arthrostylidium schomburgkii are fashioned into blowguns called “cerbatanas” by Hoti and Yekwana Amerindians. Arthrostylidium venezuelae (Steud.) McClure, J. Wash. Acad. Sci. 32: 172. 1942. —Chusquea venezuelae Steud., Syn. Pl. Glumac. 1: 337. 1855 [1854]. —Luñek, Luse. Scrambling, scandent, thicket-forming bamboo with culms 1–4 m long; branches 7– 15 at each node; inflorescence 4–12 cm long; spikelets 10–20 mm long, 5–7-flowered. Montane forests, 1000–1900 m; Bolívar (Cerro Guaiquinima, Cerro Venamo, Macizo del Chimantá, mesa between Ptari-tepui and Sororopán-tepui, eastern base Uaipántepui), Amazonas (Cerro Marahuaka, western Cerro Yutajé, Sierra de la Neblina). Venezuelan Coastal cordillera, Mérida; Central America, Lesser Antilles, Colombia, Guyana, Ecuador, Peru, northwestern Brazil. Arthrostylidium sp. A Climbing bamboo; culms slender, smooth;

branches 2–4 at each node; foliage leaf blades 8–12 × 1.4–1.6 cm; single collection sterile. Forests, ca. 100 m; Amazonas (Río Coromoto). Arthrostylidium sp. B Erect bamboo ca. 1 m tall; foliage leaves glabrous; single collection sterile. Montane forests, ca. 1300 m; Amazonas (Sierra Parima). Of doubtful affinity, and only tentatively referred to this genus. Arthrostylidium sp. C Scandent bamboo climbing to 15 m, mostly leafless when in flower; branches at midculm nodes ca. 50; inflorescences 6–8 cm long; spikelets 12–20 mm long, 3–6-flowered, awnless. Forested slopes, 100–300 m; Bolívar (Piedra Mapollos near Río Parguaza). Arthrostylidium sp. D. —Jibaju. Shrubby bamboo 1–3 m tall, forming dense colonies; foliage leaves with oral setae few, the blades 12–16 × 1.2–2 cm, often reflexed, rounded basally above the pseudopetioles; inflorescence 5–8 cm long, few-flowered, the rachis straight; spikelets all disarticulated in the single flowering specimen, the florets 8–10 mm long, awnless. Locally dominant in understory of seasonally flooded scrub forests on white sand, granitic outcrops, 100–200 m; Amazonas (Caño Pimichín, Cerro Cucurito, Río Atabapo, Río Atacavi, Río Baría, Río Pacimoni, San Pedro de Cataniapo). Colombia (Meta). The single flowering collection is over-mature, so description of this distinctive species must await collection of more material.

8. ARUNDINELLA Raddi, Agrostogr. Bras. 36, t. 1, fig. 3. 1823. [Subfamily Panicoideae, Tribe Arundinelleae] by Emmet J. Judziewicz Mostly coarse, cespitose perennials. Leaves with ligules membranous; blades linear. Inflorescence a terminal, contracted, bushy panicle, the spikelets usually paired, disarticulating from pedicel at maturity, or the glumes and lower floret persistent. Spikelets laterally compressed, lanceolate-ovate, gaping at maturity, 2flowered; glumes unequal, slenderly lanceolate, keeled, few-veined; lower glume shorter than the lower floret; upper glume as long as the spikelet, much exceeding the florets; lower floret sterile or staminate, lanceolate, persistent, awnless, membranous, few-veined; upper floret bisexual, much shorter than and concealed by the lower floret, at maturity disarticulating from the rachilla internode; lemma with

Arundinella 39

Fig. 20. Arundinella hispida

40

P OACEAE

callus bearded, long-awned from the tip or from between 2 apical teeth, the awn geniculate; palea enclosed by the margins of the lemma. Pantropics; ca. 50 species, 2 in Venezuela, 1 of these in the flora area. Arundinella hispida (Humb. & Bonpl. ex Willd.) Kuntze, Revis. Gen. Pl. 2: 761. 1891. —Andropogon hispidus Humb. & Bonp. ex Willd., Sp. Pl. 4: 908. 1806. Piptatherum confine Schult., Mantissa 2: 184. 1824. —Arundinella confinis (Schult.) Hitchc. & Chase, Contr. U.S. Natl. Herb. 18: 290. 1917. Reed-like perennial 1–2 m tall; leaves 15– 30 × 0.6–1.2 cm, coarse; inflorescence 15–30 cm long, narrowly ellipsoid, bushy; spikelets

3–4.5 mm long (excluding kinked, brown awns 2–5 mm long), strongly veined, the glumes slightly recurved. Savannas, forest edges, edges of Mauritia palm swamps, 50– 2200 m; Bolívar (near Ciudad Piar, 29 km east northeast of Icabarú, Roraima-tepui, Sororopán-tepui, Upuigma-tepui), Amazonas (Río Orinoco, Sierra Parima). Widespread in the Venezuelan Coastal Cordillera; Neotropics. ◆Fig. 20.

9. ATRACTANTHA McClure, Smithsonian Contr. Bot. 9: 42. 1973. [Subfamily Bambusoideae, Tribe Bambuseae] by Emmet J. Judziewicz Small to medium-sized, scandent, cespitose bamboos. Rhizomes short, determinate. Culms slender, hollow to solid, the midculm nodes with a branch bud at the summit of a promontory, this producing 3 primary branches, which may soon rebranch to form numerous subsidiary branches. Culm leaves with erect blades; foliage leaves with both inner and outer ligules present; oral setae usually numerous; blades linear to lanceolate-ovate. Inflorescences capitate heads to spicate racemes, indeterminate (forming pseudospikelets) or determinate (forming true spikelets); if indeterminate, then pseudospikelets with branching axes subtended by a glumelike bract and bearing a prophyllate bract as the first lateral appendage. Spikelets with one functional floret and occasionally a second, tiny, sterile rudiment on an elongate prolongation of the rachilla; floret lanceolate, falcate, attenuate, indurate, brownish, smooth, obscurely veined; lemma 7–11-veined, mucronate to shortawned, concealing the palea; lodicules (0)3; stamens 3; stigmas 2. Colombia, Venezuela, Brazil (Amazonas, Bahia, Espírito Santo); 5 species, 1 in Venezuela. Atractantha amazonica Judz. & L.G. Clark, Novon 1: 78. 1991. Cespitose woody bamboo; culms scandent, to 6 m long, later pendent in trees; foliage leaf blades 10–17 × 1–1.4 cm; inflorescences of 1(2) terminal, spicate racemes 7–15 cm

long, bearing true spikelets; spikelets 20–28 × 1.5–2 mm; glumes 6–10 mm long, shortawned, the florets spindle-shaped, slightly falcate, pungent. Lowland forests, ca. 100 m; Amazonas (Río Baría). Colombia (Vaupés), Brazil (Amazonas: Río Marié). ◆Fig. 21.

10. AULONEMIA Goudot, Ann. Sci. Nat. Bot. sér. 3, 5: 76. 1846. [Subfamily Bambusoideae, Tribe Bambuseae] by Emmet J. Judziewicz Small to medium-sized, cespitose bamboos. Culms erect or scandent, hollow; midculm nodes each bearing a single, dominant branch, this strongly divergent and sometimes nearly as large as the culm; inner and outer ligules present; oral setae

Atractantha

Fig. 21. Atractantha amazonica

41

42

P OACEAE

present, typically prominent. Leaf blades linear or lanceolate to more commonly ovate, often reflexed, deciduous. Inflorescences terminal, open panicles of determinate, few- to many-flowered spikelets. Spikelets with florets disarticulating above the persistent glumes, the lower glume 1–3-veined, conspicuously shorter than the 5–7-veined upper glume; lowermost and uppermost florets often sterile; functional florets with lemmas 7–9-veined, obtuse, mucronate, or aristate; stamens 3; stigmas 2. Southern Mexico, Central America, Colombia, Venezuela, Guyana, Ecuador, Peru, Brazil, Bolivia; 40–50 species, ca. 12 in Venezuela, 7 of these in the flora area. Key to the Species of Aulonemia 1. 1. 2(1). 2. 3(2). 3. 4(3). 4. 5(4). 5. 6(5).

6.

Foliage leaf blades with midrib very excentric, placed only 5–7 mm from the edge of a blade 22–25 mm wide ................................................ A. sp. A Foliage leaf blades with midrib not noticeably excentric ........................ 2 Foliage leaf sheaths with numerous oral setae all along their margins ...................................................................................... A. aff. subpectinata Foliage leaf sheaths with few or no marginal oral setae ......................... 3 Foliage leaf blades densely puberulent on lower surface .................. A. sp. B Foliage leaf blades glabrous on lower surface .......................................... 4 Spikelets 12–20 mm long ................................................................. A. deflexa Spikelets (at least the largest in a given inflorescence) 22–70 mm long ................................................................................................................ 5 Dwarf plants ca. 0.5 m tall; foliage leaf blades ca. 4 × 0.7 cm; spikelets purple throughout ............................................................................ A. sp. C Plants 1–3(–10) m tall or long; foliage leaf blades 11–20 × 1.8–4.5 cm; spikelets with glumes purple, the florets greenish .............................. 6 Spikelets 2.2–4 cm long; lemmas 8–10 mm long, obtuse, slightly tridentate, glabrous to sparsely puberulent on the back, the margins shortciliate ................................................................................ A. chimantaensis Spikelets 4–7 cm long; lemmas 10–13 mm long, acute, densely puberulent throughout with prickle-like hairs .................................. A. jauaensis

Aulonemia chimantaensis Judz. & Davidse, Novon 1: 81. 1991. Shrub 1–2 m; foliage leaf blades 11–25 × 1.3–2.5 cm; spikelets to 15-flowered, the lemmas strongly veined. Thickets, stream sides and savannas on tepui summits, 2100–2200 m; Bolívar (Macizo del Chimantá). Endemic. ◆Fig. 23.

1.7–4.5 cm; spikelets 12–20 mm long, loosely to densely 4- or 5-flowered, purple; lemmas 7–9 mm long, glabrous, shiny or not, weakly veined. Forming thickets in open places, bases of cliffs, stream sides, bogs, rocky forests of tepuis, 2100–2800 m; Bolívar (Ilútepui, Ptari-tepui, Roraima-tepui). Adjacent Guyana. ◆Fig. 24.

Aulonemia deflexa (N.E. Br.) McClure, Smithsonian Contr. Bot. 9: 56. 1973. —Arundinaria deflexa N.E. Br., Trans. Linn. Soc. London, Bot. 6: 75. 1901. Arthrostylidium steyermarkii McClure, Fieldiana, Bot. 28: 31. 1951. —Aulonemia steyermarkii (McClure) McClure, Smithsonian Contr. Bot. 9: 61. 1973. Shrub 1–2 m; foliage leaf blades 10–22 ×

Aulonemia jauaensis Judz. & Davidse, Novon 1: 83. 1991. Scandent bamboo with culms to 10 m long; foliage leaf blades 15–20 × 3–4.5 cm; spikelets to 23-flowered, the lemmas weakly veined. Wet forests at bases of waterfalls on tepui, 1900–2100 m; Bolívar (Cerro Jaua). Endemic. ◆Fig. 22.

Aulonemia 43

Fig. 22. Aulonemia jauaensis

Fig. 23. Aulonemia chimantaensis

44

P OACEAE

Fig. 24. Aulonemia deflexa

Axonopus 45

Aulonemia aff. subpectinata (Kuntze) McClure, Smithsonian Contr. Bot. 9: 61. 1973. —Arthrostylidium subpectinatum Kuntze, Revis. Gen. Pl. 2: 760. 1891. Thicket-forming bamboo to 2 m tall; foliage leaf blades 15–25 × 3–6 cm; both collections sterile. Shrubby forests in ravines on tepuis, ca. 2200 m; Bolívar (Macizo del Chimantá), Amazonas (Cerro Coro Coro). Anzoátegui, Distrito Federal, Mérida, Miranda, Táchira. Aulonemia sp. A Shrub ca. 1.5 m; foliage leaves with sheath summits and upper margins with flattened, partially confluent orangish brown oral setae 1.5–3.5 cm long; blades 15–25 × 2.2–2.5 cm, cuneate basally; one sterile collection. Tepui slope forests, 600–700 m; Amazonas (eastern slope of Cerro Huachamacari). Endemic. Aulonemia sp. A is perhaps related to A.

subpectinatum or A. purpuratum (McClure) McClure of the Coastal Cordillera, the latter with smaller, less excentric blades only 8–12 × 0.7–1.7 cm. Aulonemia sp. B Shrub ca. 1.5 m; foliage leaf sheath summit auriculate; oral setae 1 cm long, prominent; blades 20–25 × 4–5.5 cm; one sterile collection, tepui summit, 1000–2000 m; Amazonas (Cerro Duida). Endemic. Aulonemia sp. C Shrub ca. 0.5 m; leaf blades ca. 4 × 0.7 cm; inflorescence 10–15 cm long, with few, spreading branches; spikelets 2–2.7 cm long, 5–7-flowered, purple; lemmas 9–11 mm long, acute, harshly puberulent throughout. Rocky summit plains of tepuis, ca. 2600 m; locally common in Amazonas (Cerro Marahuaka). Endemic. Aulonemia sp. C is perhaps a dwarf-flowered individual.

11. AXONOPUS P. Beauv., Ess. Agrostogr. 12, 154. 1812. [Subfamily Panicoideae, Tribe Paniceae] Cabrera Lag., Gen. Sp. Pl. 5. 1816. —Panicum sect. Cabrera (Lag.) Trin., Mém. Acad. Imp. Sci. St.-Pétersbourg, Sér. 6, Sci. Math., Seconde Pt. Sci. Nat. 3, 1(2–3): 193, 195. 1834. —Paspalum sect. Cabrera (Lag.) Döll in Mart., Fl. Bras. 2(2): 113. 1877. —Axonopus sect. Cabrera (Lag.) Chase, Proc. Biol. Soc. Wash. 24: 132, 134. 1911. Anastrophus Schltdl., Bot. Zeitung (Berlin) 8: 681. 1850. —Paspalum sect. Anastrophus (Schltdl.) Benth. & Hook., Gen. Pl. 3: 1098. 1883. —Axonopus sect. Anastrophus (Schltdl.) Pilg. in Engl. & Prantl, Nat. Pflanzenfam. ed. 2, 14e: 53. 1940. Lappagopsis Steud., Syn. Pl. Glumac. 1: 112. 1855 [1854]. by Gerrit Davidse Perennials or annuals, cespitose, stoloniferous, or rhizomatous. Sheaths keeled; ligule a membrane; blades linear to linear-lanceolate, flat, folded or involute. Inflorescences terminal and axillary, of several to many digitate or paniculate, slender racemes, 1–several from the upper or terminal nodes, the spikelets borne in 2 alternating rows on the lower sides of a nearly flattened or triquetrous rachis, overlapping sequentially, rarely sunken in the rachis, the racemes simple, rarely branched, usually terminating in a spikelet, the tip rarely sterile. Spikelets subsessile or with very short pedicels, dorsally compressed, oriented with the upper glume and back of the upper lemma away from the rachis; lower glume, usually absent; upper glume and lower lemma membranous, as large as the upper floret, sometimes longer, equal or subequal, 2–7-veined; lower floret sterile; lower palea absent; upper lemma rigid, its margins slightly involute; upper floret bisexual; upper palea similar to the upper lemma; lodicules 2, stamens 3; stigmas 2; embryo 1/3–1/2 as long as the caryopsis; hilum elliptic or shortly linear.

46

P OACEAE

Tropics and subtropics of the New World, Africa; ca. 90 species, 38 in Venezuela, 34 of these in the flora area. One species of Axonopus is native to Africa, and several species have been widely introduced into the Old World. Key to the Species of Axonopus 1. 1. 2(1).

Rachis prominently pilose with papillose-based, bristle-like hairs ......... 2 Rachis glabrous or inconspicuously pubescent ......................................... 5 Spikelets fitting tightly into pockets sunken into the rachis; rachis 1– 1.5 mm wide, extending beyond the ultimate spikelet as a flattened, sterile point 2–3 mm long ...................................................................... 3 2. Spikelets borne on the triquetrous rachis, not in sunken pockets; rachis 0.3–0.7 mm wide, terminating in a spikelet ......................................... 4 3(2). Annuals; second lowest raceme in the primary inflorescence 2–5.5 cm long; rachis margins inconspicuously cartilaginous .............. A. excavatus 3. Perennials; second lowest raceme in the primary inflorescence 6–12 cm long; rachis margins distinctly, broadly cartilaginous ........................... ........................................................................................ A. chrysoblepharis 4(2). Blades clearly differentiated from the sheaths on the abaxial side, the collar distinctly evident ................................................................ A. aureus 4. Blades not clearly differentiated from the sheaths on the abaxial side, the collar not clearly distinct ......................................................... A. canescens 5(1). Annuals; spikelets 0.9–1.5 mm long ........................................... A. capillaris 5. Perennials; spikelets 1.5–4.5 mm long ..................................................... 6 6(5). Mature upper florets brown or purple ...................................................... 7 6. Mature upper florets straw-colored or whitish ....................................... 13 7(6). Upper floret 0.5–1.2 mm shorter than the spikelet .............. A. leptostachyus 7. Upper floret about as long as the spikelet ................................................ 8 8(7). Leaves primarily cauline, mostly borne in the upper 2/3 of the culm, dimorphic, those of the primary culm approximately twice as long as those of the numerous branches; margins of the leaf blades with wellspaced, papillose-based cilia near the base .............................. A. ramosus 8. Leaves primarily basal or at least mostly borne in the lower 1/2 of the culm, monomorphic, the culms unbranched except at the base; margins of the leaf blades with or without papillose-based cilia ............... 9 9(8). Spikelets 2.9–3 mm long ................................................................. A. gracilis 9. Spikelets 1.5–2.5 mm long ....................................................................... 10 10(9). Spikelets 2.3–2.5 mm long; tips of leaf blades obtuse as folded .......................................................................................... A. magallanesiae 10. Spikelets 1.5–2 mm long; tips of leaf blades boat-shaped and acute as folded .................................................................................................... 11 11(10). Sheaths strongly equitant; elongated internodes at the base of the plant 4 or more, these usually completely covered by the strongly distichous sheaths; collar not evident or inconspicuous .................... A. flabelliformis 11. Sheaths keeled but not strongly equitant; elongated internodes usually < 4 and at least partially exposed; collar well developed .................. 12 12(11). Spikelets glabrous .................................................................. A. cuatrecasasii 12. Spikelets sparsely pubescent........................................................ A. pennellii

Axonopus 47

13(6). Midvein of the upper glume suppressed or not well developed, at least in the majority of the spikelets ............................................................... 14 13. Midvein of the upper glume usually well developed in the majority of the spikelets ............................................................................................... 20 14(13). Spikelets densely hairy, their veins often obscured by the brown or white hairs ........................................................................................... A. purpusii 14. Spikelets glabrous or sparsely hairy, when hairy their veins not obscured by the hairs .......................................................................................... 15 15(14). Floret about as long as the spikelet, or if slightly shorter, then only 0.1– 0.3 mm shorter ..................................................................................... 16 15. Floret 0.4 mm or more shorter than the spikelet ................................... 19 16(15). Upper floret pale brown .................................................................. A. gracilis 16. Upper floret straw-colored ....................................................................... 17 17(16). Culms prostrate in the lower half, all basal internodes elongated; leaves cauline, the functional green leaves mostly borne in the upper 1/3 of flowering culms; spikelets 2.1–2.8 mm long ................................... A. sp. A 17. Culms primarily erect, the basal internodes elongated or not; leaves primarily basal, if also cauline, then the functional green leaves present to the base of flowering culms; spikelets 1.5–2.3 mm long ................ 18 18(17). Plants mostly < 50 cm tall; racemes 2–4, 3–5 cm long; broadest leaf blades 3–4 mm wide ................................................................ A. fissifolius 18. Plants mostly 75–150 cm tall; racemes 10–24, mostly > 12 cm long; broadest leaf blades 12–22 mm wide .............................................. A. iridifolius 19(15). Spikelets 2–2.5 mm long; plants stoloniferous; upper floret 0.3–0.6 mm shorter than the spikelet ...................................................... A. compressus 19. Spikelets 3–3.6 mm long; plants cespitose, never stoloniferous; upper floret 0.8–1.2 mm shorter than the spikelet ........................ A. longispicus 20(13). Culms with numerous elongated internodes; developed leaf blades 3– 7 cm long, borne primarily in the upper 1/2 of the culm ........ A. caulescens 20. Culms usually with few elongated internodes; developed leaf blades usually > 10 cm long, borne primarily near the base or in the lower 1/3 of the culm ................................................................................................ 21 21(20). Leaf blades 1–3 mm wide ........................................................................ 22 21. Leaf blades 5–30 mm wide ...................................................................... 23 22(21). Inflorescences usually noticeably shorter than the leaves, sometimes as long; upper floret 0.5–0.7 mm shorter than the spikelet ....................... ......................................................................................... A. triglochinoides 22. Inflorescences much longer than the leaves; upper floret 0.2–0.6 mm shorter than the spikelet ................................................. A. casiquiarensis 23(21). Leaf blades acuminate to acute at the apex, never splitting ................. 24 23. Leaf blades rounded or obtuse at the apex, often splitting .................... 25 24(23). Upper floret nearly as long as the spikelet; spikelets 1.7–2.1 mm long .................................................................................................... A. eminens 24. Upper floret 0.8–1.2 mm shorter than the spikelet; spikelets 3–3.6 mm long ........................................................................................ A. longispicus 25(23). Lowest internodes of the culm alternating between a long internode and 2 or more very short internodes, the long internodes easily visible, the short internodes hidden by the overlapping sheaths which form distinct fan-shaped clusters along the culms .......................................... 26

48

P OACEAE

25.

26(25). 26. 27(26). 27. 28(25). 28. 29(28). 29. 30(29).

30.

31(29). 31. 32(31). 32. 33(28). 33 34(33). 34. 35(34). 35. 36(35). 36. 37(36). 37. 38(37). 38.

Lowest internodes of the culm all ± evenly short, hidden by the overlapping sheaths, or if the lower internodes elongated and the culm partially exposed, then not alternating with several very short internodes .............................................................................................................. 28 Racemes 15–80 .................................................................................. A. anceps Racemes 1–12 ........................................................................................... 27 Sheaths 15–20 cm long; flattened leaf blades 8–25 mm wide; plants 100– 300 cm tall ......................................................................... A. arundinaceus Sheaths 6–12 cm long; flattened leaf blades 3–11 mm wide; plants 70– 100 cm tall ........................................................................................ A. sp. B Rachis pilose, at least near the base, the hairs 0.5–3 mm long ............. 29 Rachis scabrous or glabrous .................................................................... 33 Longest leaf blades 4–18 cm long; plants 30–65 cm tall ........................ 30 Longest leaf blades 20–40 cm long; plants 70–200 cm tall .................... 31 Spikelets 2.2–2.5 mm long; upper floret 0.1–0.2 mm shorter than the floret; collar glabrous or sparsely puberulent near the margins of the sheath ............................................................................................... A. sp. C Spikelets 2.2–2.5 mm long; upper floret 0.1–0.2 mm shorter than the floret; collar glabrous or sparsely puberulent near the margins of the sheath ................................................................................. A. chimantensis Spikelets 1.8–2.3 mm long .................................................... A. suffultiformis Spikelets 2.5–4 mm long .......................................................................... 32 Spikelets 2.5–2.9 mm long; sheaths entirely glabrous; collar glabrous ................................................................................................. A. yutajensis Spikelets 3.5–4 mm long; sheaths densely pilose on the back and margins or only on the margins; collar pilose .......................................... A. villosus Racemes (15–)25–80 ......................................................................... A. anceps Racemes 4–18(–25) .................................................................................. 34 Sheath apex noticeably wider than the leaf base; blades and sheaths glabrous, rarely pilose in the ligular area .............................. A. steyermarkii Sheath apex about as wide as the leaf base; blades and/or sheaths pubescent at least in part or on the margins ............................................... 35 Collar glabrous ............................................................................. A. schultesii Collar pilose or puberulent ...................................................................... 36 Plants 25–70 cm tall; racemes 2–4, 2.5–10 cm long ............. A. chimantensis Plants 75–150 cm tall; racemes 7–25, 11–25 cm long ............................ 37 Upper floret 0.5–0.7 mm shorter than the spikelet, spikelets 2.8–3.8 mm long ..................................................................................... A. surinamensis Upper floret 0.1–0.3 mm shorter than the spikelet, spikelets 2.1–2.7 mm long ....................................................................................................... 38 Leaves primarily basal; culms usually with 1 elongated internode below the flag leaf ................................................................................ A. equitans Leaves basal and cauline; culms usually with 2–5 elongated internodes below the flag leaf ................................................................... A. iridifolius

Axonopus anceps (Mez) Hitchc., Man. Grasses W. Ind. 190. 1936. —Paspalum anceps Mez, Repert. Spec. Nov. Regni Veg. 15: 61. 1917.

Paspalum scoparium var. parviflorum Döll in Mart., Fl. Bras. 2(2): 107. 1877. Axonopus pruinosus Henrard, Blumea 5: 527. 1945.

Axonopus 49

Axonopus caracarahyensis G.A. Black & Fróes, Bol. Tecn. Inst. Agron. N. 20: 34, t. 3. 1950. Axonopus erectus Swallen, Fieldiana, Bot. 28: 19. 1951. Axonopus aturensis Luces, Bol. Soc. Venez. Ci. Nat. 15: 23, fig. 16. 1953. Perennial 60–200 cm tall; sheaths strongly equitant, usually glaucous; spikelets 1.7–2.6 mm long. Savannas, frequently in the ecotone with adjoining forests, near sea level to 1400 m; widespread in Delta Amacuro, Bolívar, and Amazonas. Anzoátegui, Apure, Guárico, Miranda, Monagas, Sucre, Zulia; Trinidad, Colombia, Guyana, Suriname, French Guiana, Brazil. ◆Fig. 25. A growth form with the long internodes alternating with a cluster of very short internodes in the lower part of the culm occurs sporadically in this species, including an isotype of Axonopus erectus (Steyermark 59400, F). Axonopus arundinaceus G.A. Black, Mem. New York Bot. Gard. 9: 251. 1957. Perennial 1–3 m tall with elongated basal internodes alternating with clusters of short internodes. Montane savanna along stream, ca. 800 m; Amazonas (Cerro Moriche). Endemic. Axonopus aureus P. Beauv., Ess. Agrostogr. 12. 1812. —Paspalum aureum (P. Beauv.) Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 93, t. 27. 1815 [1816]. —Digitaria aurea (P. Beauv.) Spreng., Syst. Veg. 1: 272. 1825 [1824]. Panicum chrysites Steud., Syn. Pl. Glumac. 1: 38. 1855 [1853]. —Paspalum chrysites (Steud.) Döll in Mart., Fl. Bras. 2(2): 117. 1877. —Axonopus chrysites (Steud.) Kuhlm., Comm. Lin. Telegr., Bot. 67(Bot. 11): 88. 1922. Panicum exasperatum Nees ex Steud., Syn. Pl. Glumac. 1: 62. 1855 [1853]. —Axonopus exasperatus (Nees ex Steud.) G.A. Black, Advancing Frontiers Pl. Sci. 5: 168, fig. 8d. 1963. Paspalum carinato-vaginatum Mez, Repert. Spec. Nov. Regni Veg. 15: 31. 1917. —Axonopus carinato-vaginatus (Mez) H. Scholz, Willdenowia 8: 95. 1977. Axonopus minutus Luces, Bol. Soc. Venez. Ci. Nat. 15: 22, fig. 15. 1953.

Perennial, the triquetrous racemes bearing golden, papillose-based, bristle-like hairs, the base of the leaf blades usually spreading. Savannas, 50–900 m; widespread in Bolívar and Amazonas. Táchira, Yaracuy; Mexico, Belize, Guatemala, Honduras, El Salvador, Nicaragua, Costa Rica, Panama, Caribbean, Colombia, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia. ◆Fig. 26. Axonopus canescens (Nees ex Trin.) Pilg., Nat. Pflanzenfam. ed. 2, 14E: 55. 1940. —Paspalum pulchrum Nees in Mart. et al., Fl. Bras. Enum. Pl. 2: 79. 1829. —Axonopus pulcher (Nees) Kuhlm., Comm. Lin. Telegr., Bot. 67(Bot. 11): 88. 1922. Panicum chrysodactylon Trin., Mém. Acad. Imp. Sci. St.-Pétersbourg, Sér. 6, Sci. Math., Seconde Pt. Sci. Nat. 3, 1(2– 3): 197. 1834. —Paspalum chrysodactylon (Trin.) Döll in Mart., Fl. Bras. 2(2): 118. 1877. —Axonopus chrysodactylus (Trin.) Kuhlm., Comm. Lin. Telegr., Bot. 67(Bot. 11): 48, 87. 1922. Paspalum chrysodactylon var. psilachne Döll in Mart., Fl. Bras. 2(2): 118. 1877. —Axonopus canescens var. psilachne (Döll) G.A. Black, Advancing Frontiers Pl. Sci. 5: 167. 1963. Paspalum chrysodactylon var. glabratum Döll in Mart., Fl. Bras. 2(2): 118. 1877. —Axonopus sprucei var. glabratus (Döll) G.A. Black, Advancing Frontiers Pl. Sci. 5: 173. 1963. Paspalum chrysodactylon var. villosum Döll in Mart., Fl. Bras. 2(2): 118. 1877. Axonopus sprucei G.A. Black, Advancing Frontiers Pl. Sci. 5: 172. 1963. —Axonopus carinato-vaginatus var. sprucei (G.A. Black) H. Scholz, Willdenowia 8: 95. 1977. Axonopus paucisetosus G.A. Black, Advancing Frontiers Pl. Sci. 5: 170. 1963. Perennial, the triquetrous racemes bearing golden, papillose-based bristle-like hairs, the base of the leaf blades usually erect. Savannas, 50–1100 m; widespread in Bolívar and Amazonas. Falcón, Guárico, Monagas, Táchira, Yaracuy, Zulia; Colombia, Guyana, Suriname, French Guiana. Brazil, Bolivia. Axonopus capillaris (Lam.) Chase, Proc. Biol. Soc. Wash. 24: 133. 1911. —Paspalum capillare Lam., Tabl. Encycl. 1:

50

P OACEAE

176. 1791. —Anastrophus capillaris (Lam.) Nash, N. Amer. Fl. 17(2): 161. 1912. —Chinchorro de culebra. Paspalum extenuatum Nees in Mart. et al., Fl. Bras. Enum. Pl. 2: 25. 1829. —Axonopus extenuatus (Nees) Kuhlm., Comm. Lin. Telegr., Bot. 67 (Bot. 11) 87. 1922. Paspalum minutum Trin., Linnaea 10(3): 293. 1836. Axonopus laxus Luces, Bol. Soc. Venez. Ci. Nat. 15: 20, fig. 13. 1953. Delicate annual 10–60 cm tall; racemes 2; spikelets 0.9–1.5 mm long. Disturbed areas in savannas, cultivated areas, roadsides, clearings, 50–600 m; Bolívar (Ciudad Bolívar-Caicara del Orinoco, near La Paragua, Maripa-Aripao), Amazonas (San Carlos de Río Negro). Anzoátegui, Falcón, Guárico, Portuguesa, Sucre; Guatemala, Honduras, El Salvador, Costa Rica, Panama, Caribbean, Colombia, Guyana, French Guiana, Suriname, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 29. Axonopus casiquiarensis Davidse, Ann. Missouri Bot. Gard. 74: 419, fig. 4. 1987. Stoloniferous perennial to 90 cm tall; leaves narrow; racemes usually 2 or 3, rarely 5. Colombia, Venezuela, Brazil; 2 varieties, both in the flora area. Key to the Varieties of A. casiquiarensis 1 Spikelets 2.7–3.7 mm long ........................ .............................. var. casiquiarensis 1. Spikelets 2.1–2.5 mm long .............. var. A A. casiquiarensis var. casiquiarensis White-sand savannas, river banks, sandy pockets on lajas, 50–1100 m; rare in Bolívar (Gran Sabana), common in Amazonas (Río Atabapo, Río Casiquiare, Río Pasimoni). Colombia, Brazil. A. casiquiarensis var. A Moist savannas, white-sand savannas, 200–500 m; rare in Amazonas (base of Cerro Duida, Cerro Vinilla, La Esmeralda). Endemic. Axonopus caulescens (Mez) Henrard, Blumea 4: 510. 1941. —Paspalum caulescens Mez, Bot. Jahrb. Syst. 56(Beibl. 125): 10. 1921. Perennial 30–70 cm tall, with numerous elongated internodes and narrow (2–3 mm),

acute, typically widely spreading, arcuate leaf blades, the leaves borne toward the top of the culms. Crevices in rocks, sandy pockets in rocky places on bluffs and river banks, 700–1700 m; widespread in Bolívar. Guyana. ◆Fig. 30. Axonopus chimantensis Davidse, Ann. Missouri Bot. Gard. 74: 418, fig. 3. 1987. Perennial 25–70 cm tall; sheaths strongly equitant with pilose collars. Tepui meadows, open, moist sandy areas along streams and at base of rocks, 1900–2200 m; Bolívar (Macizo del Chimantá). Endemic. Axonopus chrysoblepharis (Lag.) Chase, Proc. Biol. Soc. Wash. 24: 134. 1911. —Cabrera chrysoblepharis Lag., Gen. Sp. Pl. 5. 1816. —Panicum chrysoblephare (Lag.) Steud., Syn. Pl. Glumac. 1: 38. 1855 [1853]. —Paspalum chrysoblephare (Lag.) Döll in Mart., Fl. Bras. 2(2): 119. 1877. Paspalum immersum Nees in Mart. et al., Fl. Bras. Enum. Pl. 2: 82. 1829. —Panicum immersum (Nees) Trin., Mém. Acad. Imp. Sci. St.-Pétersbourg, Sér. 6, Sci. Math., Seconde Pt. Sci. Nat. 3, 1(2–3): 197. 1834. —Axonopus immersus (Nees) Kuhlm., Comm. Lin. Telegr., Bot. 67(Bot. 11): 87. 1922. Paspalum gnaphalioideum Müll. Hal., Bot. Zeitung (Berlin) 19(45): 332. 1861. Paspalum immersum var. pilosum Döll in Mart., Fl. Bras. 2(2): 114. 1877. —Axonopus aureus var. pilosus (Döll) Henrard, Blumea 4: 510. 1941. Panicum savannarum Schltdl. ex G.A. Black, Advancing Frontiers Pl. Sci. 5: 161. 1963. Perennial with sunken spikelets and golden, papillose-based, bristle-like hairs surrounding the raceme cavities. Savannas, 50–500 m; expected in Bolívar. Aragua, Barinas, Cojedes, Guárico, Lara, Portuguesa; Bolivia, Brazil, Caribbean, Colombia, Costa Rica, French Guiana, Guatemala, Mexico, Panama, Paraguay, Peru. Axonopus chrysoblepharis is not yet definitely known from the Venezuelan Guayana, but is to be expected from the lowland savannas of Bolívar. Axonopus compressus (Sw.) P. Beauv., Ess. Agrostogr. 12, 154. 1812. —Milium compressum Sw., Prodr. 24. 1788.

Axonopus 51

—Agrostis compressa (Sw.) Poir. in Lam., Encycl. suppl. 1: 259. 1810. —Paspalum compressum (Sw.) Raspail, Ann. Sci. Nat. (Paris) 5: 301. 1825. Paspalum tristachyon Lam., Tabl. Encycl. 1: 176. 1791. Paspalum platycaulon Poir. in Lam., Encycl. 5: 34. 1804. —Digitaria platicaulis (Poir.) Desv., Mém. Soc. Agric. Angers 1: 166. 1831. —Panicum platycaulon (Poir.) Kuntze, Revis. Gen. Pl. 3(2): 363. 1898. —Anastrophus platycaulis (Poir.) Nash ex Small, Fl. S.E. U.S. 79. 1903. Paspalum laticulmum Spreng., Syst. Veg. 1: 245. 1825 [1824]. Digitaria uniflora Salzm. ex Steud., Nomencl. Bot. ed. 2, 1: 508. 1841 [1840]. Paspalum macropodium Steud., Syn. Pl. Glumac. 1: 19. 1855 [1853]. —Axonopus compressus var. macropodius (Steud.) G.A. Black, Advancing Frontiers Pl. Sci. 5: 82. 1963. Paspalum depressum Steud., Syn. Pl. Glumac. 1: 20. 1855 [1853]. Paspalum filostachyum A. Rich. ex Steud., Syn. Pl. Glumac. 1: 20. 1855 [1853]. Paspalum guadaloupense Steud., Syn. Pl. Glumac. 1: 18. 1855 [1853]. Paspalum conjugatum var. subcordatum Griseb., Abh. Königl. Ges. Wiss. Göttingen 7: 262. 1857. Paspalum furcatum var. fissum Döll in Mart., Fl. Bras. 2(2): 104. 1877. Paspalum furcatum var. parviflorum Döll in Mart., Fl. Bras. 2(2): 104. 1877. Paspalum raunkiaerii Mez, Repert. Spec. Nov. Regni Veg. 15: 60. 1917. Axonopus multipes Swallen, J. Wash. Acad. Sci. 23: 459. 1933. Axonopus compressus var. australis G.A. Black, Advancing Frontiers Pl. Sci. 5: 81. 1963. Stoloniferous perennial 20–85 cm tall; floret 0.3–0.6 mm shorter than the spikelet. Open secondary vegetation, roadsides, lawns, disturbed forest margins, near sea level to 1500 m; frequent in Delta Amacuro, Bolívar, and Amazonas. Anzoátegui, Apure, Aragua, Barinas, Cojedes, Distrito Federal, Falcón, Guárico, Lara, Mérida, Miranda, Monagas, Nueva Esparta, Portuguesa, Sucre, Táchira, Zulia; U.S.A., Mexico, Guatemala, Belize, Honduras, El Salvador, Nicaragua, Costa Rica, Panama, Caribbean, Colom-

bia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina, Uruguay, widely introduced in the Old World tropics. ◆Fig. 32. Axonopus cuatrecasasii G.A. Black, Advancing Frontiers Pl. Sci. 5: 147. 1963. Perennial ca. 60–80 cm tall, usually strongly stoloniferous; upper floret brown. Sandy areas near rivers, ca. 100 m; rare in Amazonas (near San Fernando de Atabapo). Anzoátegui, Monagas, Yaracuy; Colombia, Bolivia. Axonopus cuatrecasasii is tentatively recognized for the flora from a single collection (Curran s.n., NY) that is atypical because it lacks the usually prominent stolons that characterize this species. However, stolons are often neglected by collectors. Axonopus eminens (Nees) G.A. Black, Advancing Frontiers Pl. Sci. 5: 92. 1963. —Paspalum eminens Nees in Mart. et al., Fl. Bras. Enum. Pl. 2: 30. 1829. Axonopus gentilis Henrard, Blumea 5: 276, fig. s.n. 1942. Cespitose perennial 1–2 m tall with conspicuously narrowed leaf bases and numerous racemes. Savannas, 800–900 m; rare in Bolívar (near Santa Elena de Uairén). Bolivia, Guyana, Suriname, Brazil. Axonopus equitans Hitchc. & Chase, Contr. U.S. Natl. Herb. 18: 301. 1917. Perennial 40–100 cm tall. Herbaceous vegetation on granitic outcrops, 100–1100 m; Amazonas (Río Casiquiare, Río Coro Coro). Caribbean (Trinidad), Guyana, Suriname, French Guiana, Brazil. Axonopus equitans is not very clearly differentiated from A. iridifolius. As here recognized, A. equitans is interpreted as generally shorter, with primarily basal foliage, fewer racemes, and a tendency toward narrower blades compared to A. iridifolius. Axonopus excavatus (Nees ex Trin.) Henrard, Blumea 4: 509. 1941. —Paspalum excavatum Nees ex Trin., Gram. Panic. 88. 1826. Paspalum appendiculatum J. Presl, Reliq. Haenk. 1: 211. 1830. —Axonopus appendiculatus (J. Presl) Hitchc. & Chase, Contr. U.S. Natl. Herb. 18: 300. 1917.

52

P OACEAE

Annual with sunken spikelets and golden, papillose-based, bristle-like hairs surrounding the raceme cavities. Savannas, 50–200 m; Bolívar (Cerro Gavilán), Amazonas (near Puerto Ayacucho). Sucre; Panama, French Guiana, Brazil, Bolivia. ◆Fig. 27. Axonopus fissifolius (Raddi) Kuhlm., Comm. Lin. Telegr., Bot. 67(Bot. 11): 87. 1922. —Paspalum fissifolium Raddi, Agrostogr. Bras. 26. 1823. Paspalum compressum var. arenarium Bertoni, Anales Ci. Parag. 2: 153. 1918. Axonopus stragulus Chase, Contr. U.S. Natl. Herb. 22: 472, fig. 80. 1922. Axonopus ater Chase, J. Wash. Acad. Sci. 17: 143. 1927. Axonopus affinis Chase, J. Wash. Acad. Sci. 28: 180, fig. 2. 1938. —Axonopus compressus var. affinis (Chase) M.R. Hend., Malay. Wild Fls., Monocots. 339. 1954. Axonopus hirsutus G.A. Black, Advancing Frontiers Pl. Sci. 5: 55. 1963. Axonopus fissifolius var. coronatus G.A. Black, Advancing Frontiers Pl. Sci. 5: 58. 1963. Stoloniferous perennial 20–60 cm tall; floret as long as the spikelet or 0.1–0.2 mm shorter. River banks and open, disturbed, temporarily moist areas in savannas, pastures, sand bars in rivers, roadsides, dirt airstrips, 50–1100 m; widespread in Bolívar and Amazonas. Anzoátegui, Apure, Aragua, Distrito Federal, Falcón, Guárico, Mérida, Miranda, Táchira, Zulia; U.S.A. Mexico, Guatemala, Belize, Honduras, El Salvador, Nicaragua, Costa Rica, Panama, Caribbean, Colombia, Guyana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina, Uruguay, introduced into the Old World tropics. ◆Fig. 33. Axonopus flabelliformis Swallen, Bull. Torrey Bot. Club 75: 82. 1948. Axonopus kaietukensis Swallen, Bull. Torrey Bot. Club 75: 83. 1948. Axonopus purpurellus Swallen, Bull. Torrey Bot. Club 75: 83. 1948. Axonopus tamayonis Luces, Bol. Soc. Venez. Ci. Nat. 15: 21, fig. 14. 1953. Axonopus flabelliformis var. camporum G.A. Black, Advancing Frontiers Pl. Sci. 5, t. 11: 145. 1963. Axonopus flabelliformis var. decipiens G.A. Black, Advancing Frontiers Pl. Sci. 5: 146. 1963.

Perennial 30–100 cm tall, many-noded; sheaths strongly distichous and equitant, without a collar; blade apex acute to obtuse; upper floret brown. Savannas, 50–1400 m; widespread in Bolívar and Amazonas. Guyana, Suriname, French Guiana, Brazil. ◆Fig. 31. Axonopus flabelliformis is closely related to the Brazilian A. polydactylus (Steud.) Dedecca, but it differs from it primarily by its more numerous elongated internodes and more strongly distichous and equitant leaves. Axonopus gracilis G.A. Black, Mem. New York Bot. Gard. 9: 254. 1957. Cespitose perennials ca. 1 m tall; spikelets 2.6–3 mm long; upper floret pale brown. Moist areas in savannas, 1700–1800 m; Amazonas (Cerro Sipapo, Cerro Yutajé). Endemic. Axonopus iridifolius (Poepp.) G.A. Black, Advancing Frontiers Pl. Sci. 5: 125. 1963. —Paspalum iridifolium Poepp., Reise Chile 2: 324. 1836. Axonopus maguirei G.A. Black, Mem. New York Bot. Gard. 9: 252, fig. 5a. 1957, syn. nov. Axonopus piccae Giraldo-Cañas, Caldasia 21: 133, fig. 1–3. 1999, syn. nov. Cespitose perennial 55–150 cm tall. Savannas, waterfall basins, gravel bars in rivers, 100–1200 m; Bolívar (Gran Sabana), Amazonas (Cerro Huachamacari, Cerro Yutajé, Sierra de la Neblina). Guárico, Miranda; Colombia, Guyana, Peru, Brazil (Amazonas), Bolivia. Axonopus iridifolius is interpreted in a broad sense, but it is still not well understood. As interpreted here, the midvein of the lower lemma is variably developed and is clearly absent in some spikelets in some plants. The number of racemes and leaf blade length are also variable. It seems to be closely related to A. pubivaginatus Henrard, a species from Guyana, Suriname, French Guiana, and Brazil, which is differentiated primarily by the acute apex of its leaf blades. Until this complex is better understood, it seems best to treat it under the oldest available name. See also the note under A. equitans. Axonopus leptostachyus (Flüggé) Hitchc., Contr. U.S. Natl. Herb. 22: 471. 1922.

Axonopus 53

—Paspalum leptostachyum Flüggé, Gram. Monogr., Paspalum 122. 1810. Axonopus macrostachyus Hitchc. & Chase, Contr. U.S. Natl. Herb. 18: 301. 1917. Axonopus paranaensis Parodi, Revista Argent. Agron. 28: 111, fig. 3. 1961 [1962]. Perennial 75–150 cm tall, leaf tips obtuse; upper florets light brown, much shorter than the spikelet. Moist pockets in savannas, savanna ecotones with forests, open river banks, secondary vegetation around villages, margins of Mauritia palm swamps, 50–900 m; widespread in Bolívar and Amazonas. Apure, Monagas; Caribbean, Colombia, Guyana, Suriname, French Guiana, Brazil, Bolivia, Argentina. ◆Fig. 35. A number of Venezuelan specimens have been misidentified as Axonopus centralis Chase. Axonopus longispicus (Döll) Kuhlm., Comm. Lin. Telegr., Bot. 67(Bot. 11): 87. 1922. —Paspalum longispicum Döll in Mart., Fl. Bras. 2(2): 105. 1877. Axonopus rivularis G.A. Black, Mem. New York Bot. Gard. 9: 250. 1957. Axonopus hitchcockii G.A. Black, Advancing Frontiers Pl. Sci. 5: 106. 1963. Cespitose perennial with the pale upper floret conspicuously shorter than the spikelets. Rocky stream banks, often in poor sandy soils, 50–1200 m; Bolívar (Gran Sabana, Río Acanán), Amazonas (near Capuana, Río Cuao, Río Cunucunuma). Colombia, Guyana, French Guiana, Brazil. Axonopus magallanesiae Giraldo-Cañas, Caldasia 22: 237, fig. 1. 2000 [2001]. Cespitose perennial, leaves basal, strongly equitant; upper floret brown. Depressions in rocks, 1700–1800 m; Bolívar (Cerro Jaua). Endemic. Axonopus magallanesiae is only known from the type collection. Axonopus pennellii G.A. Black, Advancing Frontiers Pl. Sci. 5: 142, fig. 4g. 1963. Perennial ca. 90 cm tall; upper florets; brown. Savannas, 50–500 m; Bolívar (base of Auyán-tepui, near Maripa). Anzoátegui, Barinas, Monagas; Colombia, Brazil. This determination is tentative because the Venezuelan specimens have narrower leaf blades and less pubescent spikelets than those from Colombia.

Axonopus purpusii (Mez) Chase, J. Wash. Acad. Sci. 17: 144. 1927. —Paspalum purpusii Mez, Bot. Jahrb. Syst. 56(Beibl. 125): 10. 1921. Paspalum platycaulon var. gracilius Döll in Mart., Fl. Bras. 2(2): 102. 1877. Paspalum platycaulon var. parviflorum Döll in Mart., Fl. Bras. 2(2): 102. 1877. Paspalum flexile Mez, Bot. Jahrb. Syst. 56(Beibl. 125): 9. 1921. —Axonopus flexilis (Mez) Henrard, Blumea 4: 510. 1941. Axonopus anomalus Swallen, Contr. U.S. Natl. Herb. 29: 268. 1948 [1949]. Cespitose perennial. Savannas, 50–400 m; common in Bolívar and Amazonas; Anzoátegui, Apure, Barinas, Guárico, Mérida, Portuguesa, Táchira, Yaracuy, Zulia; Mexico, Guatemala, Honduras, Nicaragua, Belize, Costa Rica, Panama, Colombia, Guyana, Suriname, French Guiana, Brazil, Bolivia, Paraguay. Axonopus ramosus Swallen, Contr. U.S. Natl. Herb. 29: 413. 1950. Slender perennial 15–18 cm tall with numerous elongated internodes and branching abundantly at the upper nodes; leaves dimorphic, the blades spreading and flat; upper floret brown. Savannas, ca. 300 m; Amazonas (Río Ocamo). Suriname, French Guiana, expected in Guyana and adjacent Brazil. Axonopus ramosus is known in the flora area from only one sterile collection (A. Fernández 6701, MO). However, it is to be expected in Bolívar. It is very similar in its unusual growth form to a miniature version of the typical Myriocladus habit. Vegetatively it is best distinguished from the vegetatively somewhat similar A. caulescens by its flat (not folded) leaf blades and the shorter cilia on the ligules. The shorter spikelets (1.2–1.6 mm) with brown upper florets also immediately distinguish it from A. caulescens, which has larger spikelets (2.2–2.8 mm) with straw-colored upper florets. Axonopus schultesii G.A. Black, Advancing Frontiers Pl. Sci. 5: 123. 1963. Cespitose perennial 30–60 cm tall; leaves strongly equitant with very blunt leaf tips. White-sand savannas, 50–400 m; widespread in Amazonas. Colombia, Brazil (Amazonas). ◆Fig. 34.

54

P OACEAE

Axonopus steyermarkii Swallen, Fieldiana, Bot. 28: 20. 1951. Perennial 40–75 cm tall, the base of the leaf blades approximately 1/2–3/4 as wide as the sheath apex, pseudopetiolate. Tepui bluffs, river banks, 1000–1500 m; Amazonas (Cerro Duida, Cerro Marahuaka). Endemic. ◆Fig. 37. Axonopus suffultiformis G.A. Black, Mem. New York Bot. Gard. 9: 253, fig. 5b. 1957. Perennial 75–110 cm tall; sheaths strongly equitant; spikelets 2–2.5 mm long. Guyana, Venezuela; 2 varieties, both in the flora area. Key to the Varieties of A. suffultiformis 1. Collar well differentiated .......................... ............................... var. suffultiformis 1. Collar not differentiated .................. var. A A. suffultiformis var. suffultiformis Savannas, shrub islands, rocky open places, 200–1700 m; Bolívar (Cerro Guaiquinima, Cerro Guanay), Amazonas (Caño Picure, Cerro Aracamuni, Cerro Parú). Guyana. ◆Fig. 36. A. suffultiformis var. A Open herbaceous vegetation, 500–600 m; Amazonas (Sierra Unturán). Endemic. Axonopus surinamensis (Hochst. ex Steud.) Henrard, Blumea 5: 275. 1942. —Panicum surinamense Hochst. ex Steud., Syn. Pl. Glumac. 1: 42. 1855 [1853]. Axonopus fockei (Mez) Henrard, Blumea 4: 510. 1941. —Paspalum fockei Mez, Repert. Spec. Nov. Regni Veg. 15: 62. 1917. Cespitose perennial to 150 cm tall. Savannas, ca. 1200 m; Bolívar (Gran Sabana). Guyana, Suriname, French Guiana, Brazil. The report of Axonopus surinamensis for the flora area is tentative because it is based on a single, unverified collection, Trujillo 11583 (MY) from La Ciudadela in the Gran Sabana. Axonopus triglochinoides (Mez) Dedecca,

Bragantia 15: 280, fig. 22. 1956. —Paspalum triglochinoides Mez, Repert. Spec. Nov. Regni Veg. 15: 61. 1917. Cespitose perennial with narrow, erect leaves longer than to as long as the inflorescences. Sandy accumulation in cracks of granitic outcrops, sand bars, 100–200 m; Amazonas (Río Atabapo, Río Guayapo, Río Negro). Colombia, Brazil. ◆Fig. 28. Axonopus villosus Swallen, Fieldiana, Bot. 28: 21. 1951. Perennial 70–150 cm tall; sheaths strongly equitant, ciliate, the sides densely pilose or glabrous; collar pilose. Open herbaceous formations among rocks, 1000–1900 m; Amazonas (Cerro Duida, Cerro Guanay, Cerro Huachamacari, Cerro Yapacana). Endemic. Axonopus yutajensis G.A. Black, Mem. New York Bot. Gard. 9: 251. 1957. Perennial 75–100 cm tall; sheaths strongly equitant, glabrous; collars glabrous. Rocky savannas, 600–1400 m; Amazonas (Cerro Camani, Cerro Yutajé). Endemic. Axonopus sp. A Perennial 60–150 cm long, elongated internodes 5–20; nodes glabrate in the lower half of the culm, pilose in the upper part of the culm. Leaves cauline; sheaths keeled, glabrous to pilose; ligule a ciliolate membrane, the membrane 0.1–0.3 mm long, the cilia 0.2–0.3 mm long; collar scarcely differentiated, marked by a small cluster of pilose hairs at the margins or by a prominent line of long-pilose hairs from the margins to the keel; blades 9–22 cm long, 4–5 mm wide, folded or flattened, glabrous except long-ciliate in the lower 1/5–4/5 of the margins, the apex boat-shaped, shortly acute to obtuse. Inflorescences terminal and axillary; common axis 1.5–2.5 cm long; racemes 3–6, 6.5– 9.5 cm long; rachis 0.5–0.6 mm wide, scaberulous, without any cilia or sparsely ciliate with hairs 1–1.5 mm long, spikelets 14 or 15 per 25 mm; pedicels 0.2–0.3 mm long; spikelets 2.1–2.8 mm long, 0.9–1.1 mm wide, elliptic, obtuse, the upper glume and lower lemma equal or the upper glume to 0.3 mm shorter than the lower lemma; upper glume 4- or 5-veined, glabrous, the midvein usually

Axonopus 55

not evident, occasionally weakly developed; lower lemma 5-veined, glabrous, the midvein usually weakly developed; floret as long as the spikelet, pale brown, the apex essentially glabrous, obtuse; anthers 1.4–1.7 mm long. Herbaceous vegetation on rocky tepui summits, 1700–2200 m; Bolívar and Amazonas (Cerro Coro Coro, Cerro Guanay). Endemic. Axonopus sp. A is based on Huber 11039 (MYF, NY) and Huber 12339 (MO, MYF). It is remarkable for its long, slender culms, with numerous elongated internodes at the base, which according to the collector’s notes are creeping. For the most part they do not root at the nodes. In flowering culms the functional, green leaves are borne in the upper third of the culms. Axonopus sp. B Perennial ca. 100 cm long, elongated internodes at the base of the culms 9, in the middle part of the culm alternating with clusters of very short internodes, the clusters sometimes with short branches bearing narrower leaves; nodes sparsely pilose. Leaves cauline; sheaths keeled, pilose, especially toward the apex, winged; ligule a ciliolate membrane, the membrane 0.1–0.2 mm long, the cilia ca. 0.1 mm long; collar slightly differentiated, pilose; blades 9–26 cm long, 4–5 mm wide as folded, pilose, the hairs less dense toward the apex, long-ciliate along the margins, the apex boat-shaped, obtuse; inflorescences terminal and axillary; common axis 1.5–3 cm long; racemes 5 or 6, 4–12 cm long; rachis 0.5–0.7 mm wide, scaberulous and sparsely ciliate with hairs 1–1.5 mm long, spikelets 12 per 25 mm; pedicels 0.3– 0.6 mm long; spikelets 2.5–2.6 mm long, 0.9– 1.1 mm wide, elliptic, obtuse, the upper glume and lower lemma equal; upper glume 4- or 5-veined, glabrous, the midvein sometimes not evident, usually weakly developed; lower lemma 5-veined, glabrous, the midvein usually weakly developed; floret as long as the spikelet, straw-colored, the apex essentially glabrous, obtuse. Herbaceous vegetation on granitic mountain summits, 1800–1900 m; Amazonas (Serranía Uasadi). Endemic. Axonopus sp. B is based on Huber 12883 (MO, MYF). It is characterized, as far as known, by the numerous elongated internodes alternating with clusters of short in-

ternodes. This is also the growth habit of A. arundinaceus, which, however, is a much larger plant with longer and broader leaves and more numerous racemes. The entire group of related species (A. yutajensis, A. villosus, A. arundinaceus) is only known from a few collections, and the variation and limits of these species remain poorly understood. The growth habit also occurs in A. anceps, but only sporadically. In this species one or two of the lower internodes may be elongated, and it is possible that this is due to environmental conditions, fungal infections or some other factor, although these possibilities remain to be investigated. Axonopus sp. C Perennial 30–42 cm tall, without elongated internodes at the base of the culms; nodes pilose. Leaves basal; sheaths 2–5.5 cm long, keeled, strongly equitant, the keel winged, ciliate in the upper half; ligule a ciliolate membrane, the membrane 0.1–0.2 mm long, the cilia 0.4–0.6 mm long; collar distinct, well differentiated, horizontal, glabrous to pilose; blades 7–17 cm long, 2–4.2 mm wide as folded, stiffly erect, strongly folded from the base to the apex, glabrous on the sides, ciliate on the keel and the margins in the lower 1/3, the apex boat-shaped, obtuse. Inflorescences terminal and axillary; common axis 1.5–2.5 cm long; racemes 5–7, 3–8 cm long; rachis ca. 0.4 mm wide, scaberulous, spikelets 10–18 per 25 mm; pedicels 0.4–0.8 mm long; spikelets 2.2–2.4 mm long, 0.8–1 mm wide, elliptic, broadly acute, the upper glume and lower lemma equal, as long as the spikelet, 5–7-veined, sulcate between the veins, the interveins inconspicuously appressed-pubescent, the two central interveins wider than the lateral interveins; upper floret about as long as the spikelet, straw-colored, the apex puberulent, obtuse; anthers 3, ca. 1.9 mm long. Gravelly, open savannas, 1200–1400 m; Bolívar (Gran Sabana near Salto Aponguao and Parupa). Endemic. Axonopus sp. C is based on Huber 7199 (MO, MYF) and Huber 10491 (MO, MYF, NY). It is closely related to A. anceps but is much smaller in all respects, has fewer racemes in its inflorescences, and has short, distinctly erect leaf blades.

56

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Fig. 25. Axonopus anceps

Axonopus 57

Fig. 26. Axonopus aureus

Fig. 27. Axonopus excavatus

58

P OACEAE

Fig. 28. Axonopus triglochinoides

Fig. 29. Axonopus capillaris

Axonopus 59

Fig. 30. Axonopus caulescens

Fig. 31. Axonopus flabelliformis

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Fig. 32. Axonopus compressus

Fig. 33. Axonopus fissifolius

Axonopus 61

Fig. 34. Axonopus schultesii

Fig. 35. Axonopus leptostachyus

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P OACEAE

Fig. 36. Axonopus suffultiformis var. suffultiformis

Fig. 37. Axonopus steyermarkii

Bouteloua 63

12. BOUTELOUA Lag., Varied. Ci 2(4): 134. 1805. [Subfamily Chloridoideae, Tribe Cynodonteae] by Emmet J. Judziewicz Cespitose to rhizomatous annuals or perennials. Leaves often basal, the ligules ciliate, the blades linear. Inflorescence a terminal panicle; branches 1–many, borne singly at the nodes, sessile to short-pedicellate, sparsely to densely flowered, triquetrous, often raceme-like, sometimes deciduous from the rachis. Spikelets borne in 2 rows along the branches, several-flowered, disarticulating above the glumes, the lowest floret bisexual, the upper florets staminate or sterile, often rudimentary; glumes 1-veined, equal to unequal, longer or shorter than the lowest floret; lowest floret with lemma narrow, 3-veined, often trilobate with the midvein prolonged into an awn, the palea membranous; upper florets often reduced to aristate or triaristate rudiments. North America, Canada, U.S.A., Mexico, Central America, West Indies, Colombia, Guyana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina, Uruguay; ca. 40 species, 4 in Venezuela, 1 of these in the flora area. Bouteloua americana (L.) Scribn., Proc. Acad. Nat. Sci. Philadelphia 43: 306. 1891. —Aristida americana L., Syst. Nat. ed. 10, 879. 1759. Plant perennial, decumbent, rooting at nodes, erect portions 25–75 cm tall; inflorescence 10–20 cm long, with 5–12 racemose branches 2–5 cm long; spikelets 10–12 mm long. Disturbed places, near sea level to 100 m; Bolívar (Ciudad Bolívar). Aragua, Falcón, Guárico, Lara, Miranda, Sucre, Táchira, Zulia; southern Mexico, Central America, West Indies, Colombia, Guyana, Ecuador, northeastern Brazil. ◆Fig. 38.

Fig. 38. Bouteloua americana

64

P OACEAE

13. BRACHIARIA (Trin.) Griseb. in Ledeb., Fl. Ross. 4: 469. 1853. —Panicum sect. Brachiaria Trin., Mém. Acad. Imp. Sci. St.-Pétersbourg, Sér. 6, Sci. Math., Seconde Pt. Sci. Nat. 3(2): 233. 1834. [Subfamily Panicoideae, Tribe Paniceae] by Emmet J. Judziewicz Annuals or perennials, often decumbent and rooting and forming large colonies. Leaves with ligules short-membranous, often ciliate; blades linear. Inflorescence either of unbranched racemes or with primary branches that rebranch sparingly, the spikelets borne singly, in pairs, or in small fascicles in 2 rows. Spikelets ovate to obovate, disarticulating below the glumes, nearly terete to dorsally compressed, 2-flowered, the lower glume and lower lemma turned toward the rachis; lower glume 1/4 to nearly as long as spikelet, broad and slightly inflated, 1–manyveined; upper glume and lower lemma about as long as spikelet, 5–9-veined, membranous; lower floret sterile or staminate, the palea well developed; upper floret nearly as long as to more commonly distinctly shorter than the spikelet, cartilaginous, elliptic, acute, papillose to rugulose, straw-colored, the margins of the lemma only loosely enfolding the edges of the palea. Tropical and warm-temperate areas worldwide; ca. 100 species, 8 in Venezuela, 7 of these in the flora area. Key to the Species of Brachiaria 1. 1. 2(1). 2. 3(2). 3. 4(1). 4. 5(4). 5. 6(4). 6.

Inflorescence paniculate, at least the lower branches rebranching ........ 2 Inflorescence of unbranched racemes ....................................................... 4 Spikelets 2.2–2.6 mm long, usually strongly cross-veined ...... B. fasciculata Spikelets 3–3.6 mm long, strongly cross-veined or not ............................ 3 Plants 30–45 cm tall; spikelet pedicels pilose; spikelets finely pubescent, strongly cross-veined, the lower glume 1.5–1.8 mm long ............ B. mollis Plants 100–300 cm tall; spikelet pedicels glabrous; spikelets glabrous, not cross-veined, the lower glume 0.8–1.2 mm long .................. B. mutica Spikelets glabrous ...................................................................................... 5 Spikelets pubescent, at least in upper portion ......................................... 6 Spikelets (3–)3.5–4 mm long; upper glume 5–7-veined ................. B. arrecta Spikelets 4.3–4.7 mm long; upper glume 9-veined ................. B. plantaginea Lower glume 3/4 to as long as spikelet, broadly ovate ............. B. humidicola Lower glume 1/4–3/5 as long as spikelet, ovate-triangular ........ B. decumbens

Brachiaria arrecta (Hack. ex T. Durand & Schinz) Stent, Bothalia 1: 263. 1924. —Panicum arrectum Hack. ex T. Durand & Schinz, Consp. Fl. Afric. 5: 741. 1894. Urochloa arrecta (Hack. ex T. Durand & Schwinze) Morrone & Zuloaga, Darwiniana 31. 69. 1992. Sprawling, decumbent, and rooting plant, the culms erect, to 1.5 m tall; inflorescence of 6–8 racemes. Cultivated for forage and sparingly naturalized, 500–1400 m; Bolívar (near La Escalera and Cerro Venamo). Falcón, Lara, Monagas; Honduras, Costa Rica, Colombia, Trinidad-Tobago, Guyana, Suriname, French Guiana, Peru, Brazil, Argentina, na-

tive of tropical Africa. Brachiaria decumbens Stapf in Prain, Fl. Trop. Afr. 9: 528. 1919. —Urochloa decumbens (Stapf) R.D. Webster, Australian Paniceae 234. 1987. Sprawling, decumbent plants, erect culms ca. 0.3 m tall; inflorescence of 4–7 racemes; spikelets 4–6 mm long, narrowly elliptic. Cultivated for forage and sparingly naturalized, ca. 100 m; Amazonas (San Carlos de Río Negro). Apure, Distrito Federal; Honduras, Nicaragua, Costa Rica, Panama, Guyana, Suriname, Peru, Brazil, Bolivia, Paraguay, Argentina, native of tropical Africa.

Brachiaria 65

Fig. 39. Brachiaria fasciculata

Fig. 40. Brachiaria mutica

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P OACEAE

Fig. 41. Brachiaria plantaginea

Brachiaria fasciculata (Sw.) Parodi, Darwiniana, 15: 96. 1969. —Panicum fasciculatum Sw., Prodr. 22. 1788.

—Urochloa fasciculata (Sw.) R.D. Webster, Australian Paniceae 235. 1987. Panicum fuscum Sw., Prodr. 23. 1788.

Briza 67

—Urochloa fusca (Sw.) B.F. Hansen & Wunderlin, Novon 11(3): 368. 2001. Tufted annual, culms decumbent and rooting, erect portions to 50 cm tall; inflorescence an open panicle 8–13 cm long; spikelets broadly obovate, orangish brown to nearly black. Disturbed open areas, near sea level to 200 m; northern Bolívar. Widespread elsewhere in Venezuela; southeastern U.S.A., Mexico, Central America, West Indies, tropical South America. ◆Fig. 39. Brachiaria humidicola (Rendle) Schweick., Bull. Misc. Inform. 1936: 297. 1936. —Panicum humidicolum Rendle in Hiern, Cat. Afr. Pl. 2: 169. 1899. Urochloa humidicola (Rendle) Morrone & Zuloaga, Darwiniana 31: 80. 1992. Stoloniferous plants, erect culms to 50 cm tall; inflorescence of 2–4 racemes; spikelets 4–4.5 mm long, broadly elliptic. Cultivated for forage and sparingly naturalized, 500– 1400 m; Bolívar (near La Escalera and Cerro Venamo). Zulia; Panama, West Indies, Guyana, Suriname, French Guiana, Ecuador, Brazil, Bolivia, Paraguay, Argentina, native of tropical Africa. Brachiaria mollis (Sw.) Parodi, Darwiniana 15: 100. 1969. —Panicum molle Sw., Prodr. 22. 1788. Urochloa mollis (Sw.) Morrone & Zuloaga, Darwiniana 33: 85. 1992. Tufted annual, culms sprawling and rooting at nodes, erect portions to 50 cm tall; inflorescence an open panicle 6–9 cm long; spikelets obovate, puberulent, purple or dusky. Disturbed, open areas, 100–300 m; Bolívar (Altiplanicie de Nuria, Guri). Anzoátegui, Aragua, Guárico, Lara, Monagas,

Portuguesa, Zulia; Mexico, Central America, Jamaica, Colombia, Guyana, French Guiana, Ecuador, Peru, Brazil. Brachiaria mutica (Forssk.) Stapf in Prain, Fl. Trop. Afr. 9: 526. 1919. —Panicum muticum Forssk., Fl. Aegypt.-Arab. 20. 1775. —Urochloa mutica (Forssk.) T.Q. Nguyen, Novosti Sist. Vyssh. Rast. 1966: 13. 1966. —Hierba del Pará. Panicum purpurascens Raddi, Agrostogr. Bras. 47. 1823. Robust, sprawling, clone-forming perennial 1–3 m tall; inflorescence of 10–18 racemes; spikelets ovate, gaping, green flushed with purple. Cultivated for forage and widely naturalized, near sea level to 100 m; Delta Amacuro (Caño Mánamo, Isla Cocuina, Pedernales, Tucupita), Bolívar (Río Orinoco). Widespread elsewhere in Venezuela; southern U.S.A., Mexico, Central America, West Indies, tropical South America, native of Africa. ◆Fig. 40. Brachiaria plantaginea (Link) Hitchc., Contr. U.S. Natl. Herb. 12: 212. 1909. —Panicum plantagineum Link, Hort. Berol. 1: 206. 1827. —Urochloa plantaginea (Link) R.D. Webster, Syst. Bot. 13: 607. 1988. Tufted annual, culms creeping and rooting, to 40 cm tall; inflorescence of 3 or 4 racemes. Naturalized in wet, weedy places, near sea level to 100 m; Delta Amacuro (Isla de Guara). Apure, Distrito Federal, Falcón, Miranda, Monagas, Portuguesa, Táchira, Zulia; southern U.S.A., Mexico, Central America, West Indies, French Guiana, Ecuador, Brazil, Bolivia, Paraguay, Argentina, native of tropical Africa. ◆Fig. 41.

14. BRIZA L., Sp. Pl. 70. 1753. [Subfamily Pooideae, Tribe Poeae] by Gerrit Davidse Cespitose annuals. Stems hollow; ligule membranous. Inflorescence a panicle. Spikelets laterally compressed, several-flowered; glumes and lemmas very broad, blunt, inflated, placed at right angles to the rachilla; disarticulation above the glumes and between florets; glumes faintly veined, nearly circular, cucullate; lemma nearly circular, cordate at the base, faintly 5–9-veined; palea much shorter and narrower; stamens 3. Mostly Mediterranean, several species widely naturalized on all continents except Antarctica; ca. 5 species, 2 in Venezuela, 1 of these in the flora area.

68

P OACEAE

Briza minor L., Sp. Pl. 70. 1753. Delicate annual with pendant spikelets on flexuous pedicels. Roadside weed, ca. 1100 m; Bolívar (plateau of Cerro Venamo). Canada, U.S.A., Mexico, Guatemala, Honduras, Costa Rica, Panama, Caribbean, Colombia, Ecuador, Peru, Brazil, Bolivia, Chile, Argentina, Uruguay, Old World. ◆Fig. 42. Briza minor is native to Eurasia but is now widely naturalized in all temperate areas of the world and middle to upper elevations in tropical mountains. This is apparently a recent introduction.

Fig. 42. Briza minor

15. CENCHRUS L., Sp. Pl. 1049. 1753. [Subfamily Panicoideae, Tribe Paniceae] by Emmet J. Judziewicz Annuals or perennials. Leaves with ligules ciliate; blades linear. Inflorescence usually densely spicate, of deciduous, spiny burs formed by connate, flattened or spine-like sterile branches, each bur enclosing one or several persistent spikelets. Spikelets membranous, lanceolate to ovate, 2-flowered; lower glume about half as long as the spikelet, 1–3-veined; upper glume and lower lemma as long as spikelet, usually several-veined; lower floret usually staminate, with a well-developed palea; upper floret bisexual, firmly membranous, 5–7-veined, the palea only loosely embraced by the lemma.

Cenchrus 69

Fig. 43. Cenchrus echinatus

Fig. 44. Cenchrus brownii

70

P OACEAE

Fig. 45. Cenchrus ciliaris [expected in flora area]

Chloris 71

Temperate and tropical regions worldwide; ca. 20 species, ca. 6 in Venezuela, 2 of these in the flora area. Cenchrus ciliaris L., cultivated for forage and sometimes escaping, may eventually be found in the flora area. ◆Fig. 45. Key to the Species of Cenchrus 1. 1.

Spike densely flowered, the rachis concealed by the burs; at least some of the outer bristles as long as the bur .......................................... C. brownii Spike loosely flowered, the rachis visible between the burs; outer bristles at most 1/2 as long as the bur ................................................... C. echinatus

Cenchrus brownii Roem. & Schult., Syst. Veg. 2: 258. 1817. Annual, culms extensively creeping and rooting; burs 5–7 mm long. Trail sides and other disturbed places, 100–800 m; Bolívar (Cerro Akurimá along middle Río Caroní, Moitaco, El Callao, Río Ambuituir south of Auyán-tepui, Río Cuyuni, Santa María del Vapor). Widespread elsewhere in Venezuela; southeastern U.S.A., Mexico, Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru,

Brazil, Bolivia, introduced in Old World tropics. ◆Fig. 44. Cenchrus echinatus L., Sp. Pl. 1050. 1753. Annual, culms extensively creeping and rooting; burs 4–5.5 mm long. Savannas, disturbed open areas, 100–400 m; Bolívar (Las Claritas, Upata), Amazonas (Río Orinoco). Widespread elsewhere in Venezuela and in tropical and warm-temperate areas throughout the New World, introduced into Old World tropics. ◆Fig. 43.

16. CHLORIS Sw., Prodr. 25. 1788. [Subfamily Chloridoideae, Tribe Cynodonteae] by Emmet J. Judziewicz Rhizomatous, stoloniferous, or cespitose annuals or perennials. Culms solid; ligules membranous, ciliate, or occasionally absent; blades generally linear. Inflorescence a digitate or subdigitate cluster of (1–)several to many, slender, spike-like racemes; rachis triquetrous, scabrous, bearing subsessile spikelets alternately from 2 sides. Spikelets disarticulating above the glumes, weakly to strongly laterally compressed, several-flowered, often only the lowest floret functional; glumes narrow, unequal, keeled, 1-veined, hyaline, often shorter than the spikelet; lowest floret bisexual, usually as long as the spikelet; lemma laterally compressed, 3-veined, the lateral veins marginal and often ciliate, strongly keeled or not, with a bearded callus, the apex shortly bifid and bearing a subapical, scabrous awn; palea slightly shorter than the lemma, the keels submarginal, often ciliate at the apex; upper florets sterile, inconspicuous to conspicuous and inflated, awned or awnless; stamens 3; stigmas 2. Caryopsis dorsally compressed. Tropical and warm-temperate areas worldwide; ca. 55 species, 6 in Venezuela, 3 of these in the flora area. Key to the Species of Chloris 1. 1.

Sterile floret(s) narrowly lanceolate, ca. 2 mm long, the apex acute; racemes 4–7 cm long ................................................................... C. radiata Sterile floret(s) elliptic, obovate, or round, ca. 1.5 mm long, the apex acute, rounded, or truncate; racemes 5–15 cm long ............................. 2

72

2(1).

2.

P OACEAE

Sterile florets obovate to round, unequal, the lower conspicuously longer than the upper; racemes 5–8 cm long, straight; lower glume ca. 1 mm long; annual ................................................................................ C. barbata Sterile florets elliptic, subequal; racemes 6–15 cm long, flexuous; lower glume 2.5–3 mm long; perennial ..................................................... C. elata

Chloris barbata Sw., Fl. Ind. Occid. 1: 200. 1797. Andropogon barbatus L., Mant. Pl. 2: 302. 1771, “barbatum.” non L. 1759. Chloris inflata Link, Enum. Hort. Berol. Alt. 1: 105. 1821. Tufted annual, the culms 60–80 cm tall. Disturbed areas, 50–300 m; Delta Amacuro (Tucupita), Bolívar (El Dorado, Río Cuyuni, San Martín de Turumbán southwest of Tumeremo, 8 km southeast of Upata). Widespread elsewhere in Venezuela; southern U.S.A. (Florida), Mexico, Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina, Uruguay, Paleotropics. ◆Fig. 46. Chloris elata Desv., Mem. Soc. Agric. Angers 1: 177. 1831. —Pata gallina. Andropogon barbatus L., Syst. Nat. ed. 10, 1305. 1759, “barbatum.” —Chloris barbata (L.) Nash, Bull. Torrey Bot. Club 25: 443. 1898. Andropogon polydactylon L., Sp. Pl. ed. 2, 1483. 1763, nom. superfl. pro Andropogon fasciculatus L. —Chloris polydactyla (L.) Sw., Prodr. 26. 1788, nom. illeg. Chloris dandyana C.D. Adams, Phytologia 21: 408. 1971, nom. illeg. Tufted perennial 60–100 cm tall. Disturbed areas, 100–200 m; Bolívar (Mina de El Manganeso south of El Palmar). Widespread elsewhere in Venezuela; U.S.A. (Florida), Honduras, Guyana, Suriname, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina, Uruguay. Chloris radiata (L.) Sw., Prodr. 26. 1788. —Agrostis radiata L., Syst. Nat. ed. 10, 873. 1759. Plant tufted, stoloniferous, apparently short-lived, the culms 30–60 cm tall. Weedy areas, 50–100 m; Delta Amacuro (Tucupita), Bolívar (La Vergareña). Widespread elsewhere in Venezuela; U.S.A., southern Mexico, Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Chile, Paraguay, Pacific Ocean islands.

Fig. 46. Chloris barbata

Coelorachis 73

17. CHUSQUEA Kunth, J. Phys. Chim. Hist. Nat. Arts 95: 151. 1822. [Subfamily Bambusoideae, Tribe Bambuseae] by Emmet J. Judziewicz Small to medium-sized woody bamboos. Culms erect to arching or scandent, solid or occasionally with a small lumen; primary branch buds at midculm nodes usually developing into a single large primary branch, often with a cluster of numerous subsidiary branchlets below it, these each bearing complements of foliage leaves. Foliage leaves with inner and outer ligules present; oral setae absent; blades typically small, linear to elliptic. Inflorescence terminal, an open or contracted panicle, less commonly a raceme. Spikelets lanceolate to ovate, unawned or shortaristate, several-flowered; glumes 2, short; lowest 2 florets sterile, of lemmas only; uppermost floret bisexual, the lemma few- to many-veined; palea bicarinate but often with many veins; stamens 3; stigmas 2. Mexico, Central America, West Indies, Colombia, Guyana, Ecuador, Peru, Brazil (not in Amazonia), Bolivia, central Chile, Paraguay, Argentina; ca. 125 species, ca. 15 in Venezuela, 2 of these in the flora area. Key to the Species of Chusquea 1. 1.

Foliage leaf blades 2–10 cm × 3–10 mm ........................................ C. linearis Foliage leaf blades 17–25 cm × 25–35 mm ......................................... C. sp. A

Chusquea linearis N.E. Br., Trans. Linn. Soc. London, Bot. 6: 76. 1901. Erect or scandent shrub 0.5–2 m tall; branches at midculm nodes breaking through the sheaths; foliage leaf blades linear-lanceolate, cuneate at base, firm, striate; inflorescence (rarely seen) racemose, 4–7 cm long; spikelets 6.5–8 mm long. Streamsides, cloud forest edges, open rocks, Bonnetia forests of tepui summits, 1500–2800 m; Bolívar (Auyán-tepui, Kamarkawarai-tepui, southwest of Caraurín-tepui, Carrao-tepui, Cerro Guai-

quinima, Cerro Jaua, Kukenán-tepui, Macizo del Chimantá, Murisipán-tepui, Ptari-tepui, Roraima-tepui), Amazonas (Sierra Parima). Adjacent Guyana. ◆Fig. 47. Chusquea sp. A Shrub 2 m tall; branches at midculm nodes breaking through sheaths; foliage leaf blades lanceolate, rounded at base; inflorescence not seen. Forests at base of tepui bluff, 800–1300 m; Amazonas (eastern Cerro Huachamacari). Táchira.

18. COELORACHIS Brongn. in Duperrey, Voy. Monde (Phan.) 8: 64. 1829 [1831]. [Subfamily Panicoideae, Tribe Andropogoneae] by Emmet J. Judziewicz Cespitose perennials. Inflorescences terminal and axillary, numerous, each with a solitary, partially included, spathe-like, narrowly cylindrical raceme with densely imbricate spikelets; rachis internodes somewhat clavate, readily disarticulating along with the attached spikelets, prolonged basally into a short, peg-like, basal extension; spikelets paired, one sessile and the other pedicellate. Sessile spikelets ovate, sunken in the hollow formed by the rachis internode and spikelet pedicel; lower glume convex, coriaceous, erect and concealing the upper glume and florets, glabrous, the lower portion often pitted or rugose and obscurely veined, the upper portion with prominent wings and submarginal veins; upper glume slightly

74

P OACEAE

Fig. 47. Chusquea linearis

Coelorachis 75

Fig. 48. Coelorachis aurita

76

P OACEAE

shorter than the spikelet, membranous, strongly keeled, several-veined; florets distinctly shorter than the spikelet, hyaline; lower floret sterile, lacking a palea; upper floret bisexual, with a well-developed palea. Pedicel of pedicellate spikelet about as long as and similar to the rachis internode, often bearing a wing-like auricle on the margin opposite the sessile spikelet; spikelet sterile, staminate, or occasionally bisexual, similar to the sessile spikelet or rudimentary. Pantropics; ca. 20 species, 2 in Venezuela, 1 of these in the flora area. Coelorachis aurita (Steud.) A. Camus, Ann. Soc. Linn. Lyon sér. 2, 68: 197. 1921 [1922]. —Rottboellia aurita Steud., Syn. Pl. Glumac. 1: 361. 1855 [1854]. —Manisuris aurita (Steud.) Kuntze, Revis. Gen. Pl. 3: 356. 1898. Mnesithea aurita (Steud.) de Koning & Sosef, Blumea 31: 290. 1986. Coarse, cespitose perennial 1–2 m tall

from a knotty base; inflorescence 5–12 cm long; sessile spikelets 3.5–4.5 mm long. Savannas, 100–400 m; Bolívar (Altiplanicie de Nuria), Amazonas (base of Cerro Guanay, Puerto Ayacucho). Apure, Guárico; Mexico, Central America, Colombia, Trinidad-Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina. ◆Fig. 48.

19. COIX L., Sp. Pl. 972. 1753. [Subfamily Panicoideae, Tribe Andropogoneae] by Emmet J. Judziewicz Coarse, somewhat maize-like plants of indefinite duration. Inflorescences numerous, both axillary and terminal, long-pedunculate, compound, each composed of an ovate, bony bead enclosing a reduced pistillate branch, with a many-flowered staminate branch exserted from a terminal pore; staminate inflorescence deciduous from the summit of the bead, the bead at length deciduous from the summit of the peduncle. Pistillate branch consisting of a functional, sessile spikelet that nearly completely fills the bead, along with 2 pedicels, these usually bearing rudimentary spikelets; glumes fleshy, subequal, beaked, the apex slightly exserted from the terminal pore; florets hyaline, the lower sterile and lacking a palea, the upper pistillate and with a well-developed palea; stigmas 2, exserted. Staminate branch with spikelets in pairs or triads, 1 or 2 sessile and 1 pedicellate; rachis internodes slender, often disarticulating along with the persistent spikelets; spikelets membranous, greenish; lower glume flat to convex on back, usually winged above the middle, its margins inflexed over the edges of the keeled upper glume; florets shorter than the spikelet, both with well-developed lemmas and paleas. Southern Asia, now a pantropical weed; ca. 5 species, 1 cultivated and naturalized in the flora area. Coix lacryma-jobi L., Sp. Pl. 972. 1753. —Echepen yo’ (Yekwana). Maize-like plants, the culms 0.5–2 m × 1 cm; blades 20–40 × 2–4 cm; beads 9–12 × 6–9 mm, globose or ellipsoid, green when immature, becoming white, bluish, or black at full maturity, smooth and shiny or sometimes somewhat wrinkled. Disturbed places, near

sea level to 100 m; Delta Amacuro (Capure, Pedernales), Bolívar (Santa María de Erebato). Widespread elsewhere in Venezuela and throughout the tropics, native to southern Asia. ◆Fig. 49. Beads from Coix lacryma-jobi are strung and used as necklaces.

Coix 77

Fig. 49. Coix lacryma-jobi

78

P OACEAE

20. CORTADERIA Stapf, Gard. Chron. ser. 3, 22(570): 378. 1897. [Subfamily Arundinoideae, Tribe Arundineae] Moorea Lem., Ill. Hort. 2(Misc.): 15. 1855. by Gerrit Davidse Tall, dioecious perennials, forming large dense clumps. Basal leaves numerous; ligule a dense row of white hairs. Inflorescences large, plumy solitary panicles. Spikelets laterally compressed; florets 2–9; disarticulation above the glumes and near the base of each rachilla segment, the rachilla segment forming a callus below the detached floret; glumes subequal, 1-veined, nearly as long as the entire spikelet; lemma 3–7-veined, bearing numerous long, silky white hairs, either linear-lanceolate, tapering into an awn, or short and ovate, deeply bifid, the central awn arising between 2 awned lateral teeth; palea 1/3–1/2 as long as the lemma; callus bearded; pistillate flowers usually with sterile staminodes; staminate flowers with 3 large anthers and a rudimentary ovary. Costa Rica to Patagonia, New Zealand, New Guinea; 23 species, 7 in Venezuela, 1 of these in the flora area. Cortaderia roraimensis (N.E. Br.) Pilg., Notizbl. Bot. Gart. Berlin-Dahlem 6: 112. 1914. —Arundo roraimensis N.E. Br., Trans. Linn. Soc. London, Bot. 6: 74. 1901. Perennial tussocks 30–150 cm tall; leaves mostly basal, curling with age; panicle prominent. Widespread on the higher mountain summits and upper mountain slopes, as part of the herbaceous or open shrubby vegetation in rocky, humid areas, sometimes along streams, 1500–2800 m; Bolívar (Aprada-tepui, Auyán-tepui, Camarcaibarai-tepui, Macizo del Chimantá [Chimantá-tepui], Ilu-

tepui, Kukenán-tepui, Sierra de Maigualida, Ptari-tepui, Roraima-tepui, Tramen-tepui, Yuruaní-tepui), Amazonas (Cerro Marahuaka, Sierra de la Neblina). Guyana (Mount Roraima), Brazil (Amazonas). ◆Fig. 50. Cortaderia roraimensis is an outlying species endemic to the greater Guayana region and is a member of a genus otherwise confined to the Andean cordilleras and the central cordilleras of Costa Rica and Panama, although in temperate and subtropical southern South America some species descend to the lowlands.

21. CYNODON Rich. in Pers., Syn. Pl. 1: 85. 1805, nom. cons. [Subfamily Chloridoideae, Tribe Cynodonteae] Capriola Adans., Fam. Pl. 2: 31, 532. 1763, nom. rejic. by Emmet J. Judziewicz Perennials, the culms decumbent, creeping. Foliage often clustered near base of plant; ligules ciliate; blades linear, soft, short. Inflorescence terminal, consisting of 2–several slender, digitate or subdigitate racemes, each raceme bearing 2 rows of crowded, subsessile spikelets. Spikelets laterally compressed, 1-flowered, with a minute prolongation of the rachilla beyond the floret; glumes subequal, linear, subulate, 1-veined, shorter than the floret; floret with lemma broadly ovate, firmly membranous, 3-veined; palea narrow, shorter than the lemma; stamens 3; stigmas 2. Pantropics; 8 species, 1 in Venezuela. Cynodon dactylon (L.) Pers., Syn. Pl. 1: 85. 1805. —Panicum dactylon L., Sp. Pl. 58. 1753. —Capriola dactylon (L.) Kuntze, Revis. Gen. Pl. 2: 764. 1891.

Creeping perennial, forming large colonies by rhizomes and stolons. Inflorescence of 3–5 subequal racemes 2–6 cm long; spikelets 2–2.5 mm long. Weedy places, near sea level

Cortaderia 79

Fig. 50. Cortaderia roraimensis

80

P OACEAE

to 900 m; Delta Amacuro (Isla Mananito, Pedernales), Bolívar (Santa Elena de Uairén). Widespread elsewhere in Venezuela; worldwide in tropical and warm-temperate regions. ◆Fig. 51.

Fig. 51. Cynodon dactylon

22. DACTYLOCTENIUM Willd., Enum. Pl. 1029. 1809. [Subfamily Chloridoideae, Tribe Cynodonteae] by Emmet J. Judziewicz Cespitose to stoloniferous annuals or perennials. Leaves with ligules short, ciliate; blades linear. Inflorescence a terminal cluster of digitate, densely flowered racemes, the rachis triquetrous, bearing spikelets from 2 sides; rachis prolonged past the last spikelet as a naked bristle. Spikelets strongly laterally compressed, disarticulating above the lower glume, several-flowered, the uppermost florets rudimentary; glumes and lemmas ± similar in size and form, ovate, acuminate, 3veined, the keel prominent, scabrous, the lateral veins submarginal, inconspicuous; upper glume and often lowest lemma with a stout, short, flexuous awn; paleas shorter than lemmas; stamens 3; stigmas 2. Old World tropics and subtropics; 13 species, 1 of these a pantropical weed occurring in Venezuela.

Dichanthelium 81

Dactyloctenium aegyptium (L.) Willd., Enum. Pl. 1029. 1809. —Cynosurus aegyptius L., Sp. Pl. 72. 1753. Annual, culms creeping, the flowering culms 10–40 cm tall; inflorescence of 2–5 divergent, densely flowered racemes 1.5–5 cm long; spikelets 3–4 mm long, 3–5-flowered. Weedy places, savannas, thickets, 100–400 m; Bolívar (Altiplanicie de Nuria, 8 km southeast of Upata). Widespread elsewhere in Venezuela; worldwide in tropical areas. ◆Fig. 52.

Fig. 52. Dactyloctenium aegyptium

23. DICHANTHELIUM (Hitchc. and Chase) Gould, Brittonia 26: 59. 1974. —Panicum subg. Dichanthelium Hitchc. and Chase, Contr. U.S. Natl. Herb. 15: 142. 1910. [Subfamily Panicoideae, Tribe Paniceae] by Fernando O. Zuloaga Annual or perennial with or without a basal rosette of shorter, broader leaves than those of the culms. Ligules membranous-ciliate. Blades linear to ovate-lanceolate; foliar dimorphism present or absent. Inflorescences terminal and axillary, lax, open to occasionally contracted. Spikelets ellipsoid, cleistogamous spikelets present or absent. Lower glume 1–7-veined, upper glume and lower lemma subequal, (5–)7–15-veined; lower palea present, occasionally absent; lower flower staminate or absent. Upper anthecium indurate, smooth, with simple papillae all over its surface, shortly apiculate or crested. Canada, U.S.A., Mexico, Central America, most of South America to central Argentina and Chile; 58 species, 12 in Venezuela, 6 of these in the flora area. Dichanthelium is found from sea level to ca. 3000 m elevation. It is frequent in forest edges in wet habitats. Some species are found in open places on moist or dry sandy soils. Key to the Species of Dichanthelium 1. 1. 2(1). 2. 3(2).

Upper glume and lower lemma 10–14-veined .............................. D. davidsei Upper glume and lower lemma 7–9-veined .............................................. 2 Spikelets geminate, at least in part of the inflorescence; lower glume dimorphic; upper floret black; blades asymmetric basally ........... D. hebotes Spikelets never geminate; lower glume isomorphic; upper floret whitish to pale; blades symmetric basally ......................................................... 3 Lower palea conspicuous, as long at the lower lemma, 2 mm long; spikelets 3–3.2 mm long, ellipsoid; blades rounded at the base .... D. telmatum

82

3.

4(3). 4. 5(4).

5.

P OACEAE

Lower palea small, ca. 1/2 the length of the lower lemma, 0.7–1.8 mm long; spikelets 1.5–2.9 mm long (to 3.3 mm long in D. pycnoclados but blades cordate and clasping at the base) .......................................................... 4 Spikelets 2.2–3(–3.3) mm long, with a stipe between the lower and upper glume; panicle rachis glabrous ........................................... D. pycnoclados Spikelets 1.5–2.5 mm long, without a stipe between the lower and upper glume; panicle rachis hirsute ................................................................ 5 Spikelets short hispid, 1.5–1.9 mm long; inflorescences 2.5–9 cm long; lower palea 0.7–1 mm long; main axis of the inflorescence densely hirsute .................................................................................. D. sciurotoides Spikelets glabrous, 1.7–2.5 mm long; inflorescences 10–15 cm long; main axis of the inflorescence glabrous ................................ D. aequivaginatum

Dichanthelium aequivaginatum (Swallen) Zuloaga, Amer. J. Bot. 90: 816. 2003. —Panicum aequivaginatum Swallen, Contr. U.S. Natl. Herb. 29: 271. 1949. Panicum appressifolium Swallen, Mem. New York Bot. Gard. 9: 258. 1957. Cespitose to decumbent annual; blades linear-lanceolate, 3.5–12 × 0.5–1.3 cm; spikelets elliptic, 1.7–2.5 mm long, glabrous; upper glume and lower lemma 9-veined. Disturbed, open sandy areas in savannas, roadsides, 1100–1400 m; Bolívar (Gran Sabana, Uaipán-tepui), Amazonas (Río Yatúa, Sierra de la Neblina). Guyana, Suriname, French Guiana, northern Brazil. Dichanthelium aequivaginatum is related to D. sciurotoides and D. pycnoclados, with spikelets intermediate in size between the two species. Dichanthelium davidsei (Zuloaga & Morrone) Zuloaga, Amer. J. Bot. 90: 816. 2003. —Panicum davidsei Zuloaga & Morrone, Ann. Missouri Bot. Gard. 78: 158. 1991. Perennial, culms ca. 2.2 m long, scandent on vegetation; spikelets elliptic, 3–3.3 mm long, upper glume 11–14-veined, lower lemma 10–12-veined. Edges of forests, open places, 1200–1300 m; Bolívar (near Kavanayén, La Escalera). Brazil (Roraima). Dichanthelium hebotes (Trin.) Zuloaga, Amer. J. Bot. 90: 816. 2003. —Panicum hebotes Trin., Mém. Acad. Imp. Sci. St.Pétersbourg, Sér. 6, Sci. Math., Seconde Pt. Sci. Nat. 1: 301. 1834. Panicum mirandum Luces, J. Wash. Acad. Sci. 32: 163, fig. 8. 1942.

Scandent perennial, rooting and branching at the lower nodes; blades asymmetric basally; spikelets 1.5–1.9 mm long, usually geminate, lower glume dimorphic, 1/3–2/3 the length of the spikelet; upper floret apiculate, black. Forest margins, ca. 600 m; Bolívar (Altiplanicie de Nuria). Aragua, Sucre, Yaracuy; Ecuador, Brazil, Bolivia. Dichanthelium pycnoclados (Tutin) Davidse, Monogr. Syst. Bot. Missouri Bot. Gard. 45: 1258. 1993. —Panicum pycnoclados Tutin, J. Bot. 72: 340, fig. 10. 1934. Panicum albociliatum Swallen, Brittonia 3: 149. 1939. Panicum tiricaense Swallen, Mem. New York Bot. Gard. 9: 400. 1957. Panicum tiricaoides Swallen, Mem. New York Bot. Gard. 9: 401. 1957. Scandent perennial, creeping and rooting at lower nodes, 12–60 cm tall; blades lanceolate to ovate-lanceolate, cordate basally and stem-clasping, to 1.8 cm wide; spikelets elliptic, 2.2–3(–3.3) mm long, upper floret papillose. Forest margins, wet areas, (200–) 1000–2400 m; Bolívar (Gran Sabana and associated tepuis, Serranía de Imataca). Colombia, Guyana, Suriname, Ecuador, Peru, northern Brazil. ◆Fig. 53. Dichanthelium sciurotoides (Zuloaga & Morrone) Davidse, Novon 2: 104. 1992. —Panicum sciurotoides Zuloaga & Morrone, Novon 1: 1. 1991. Slender annual, decumbent and rooting at lower nodes; blades ovate-lanceolate, cordate basally and stem-clasping, 3–10 × 0.87–2 cm; spikelets elliptic, 1.5–1.9 mm long. Among

Dichanthelium 83

shrubs in humid places at forest edges, 800– 1400 m; Bolívar (Altiplanicie de Nuria, Gran Sabana, La Escalera). Falcón, Yaracuy; Panama, Guyana, Ecuador, Brazil, Bolivia. Dichanthelium telmatum (Swallen) Zuloaga, Amer. J. Bot. 90: 817. 2003. —Panicum telmatum Swallen, Phytologia 14: 81. 1966. Perennial, with culms branching and arching at the base; blades lanceolate, 3.5– 5.5 cm long; panicles short-exserted, 3–5 cm long; spikelets elliptic, 3–3.2 mm long, pilose, upper glume and lower lemma 7–9-veined. Burned areas of bogs, ca. 2100 m; Bolívar (Uei-tepui on Cerro El Sol). Endemic.

Fig. 53. Dichanthelium pycnoclados

84

P OACEAE

24. DIGITARIA Haller, Hist. Stirp. Helv. 2: 244. 1768, nom. cons. [Subfamily Panicoideae, Tribe Paniceae] Syntherisma Walter, Fl. Carol. 76. 1788. Trichachne Nees in Mart. et al., Fl. Bras. Enum. Pl. 2: 85. 1829. by Emmet J. Judziewicz Annuals or perennials. Leaves with membranous or uncommonly ciliate ligules; blades usually linear, soft and lax. Inflorescence a panicle of 1 to usually several or many slender, unilateral racemes, these digitate or disposed along a short axis; rachis triquetrous to flattened and winged, the spikelets borne on 2 of the 3 sides, all or in part pedicellate in pairs or triads, occasionally solitary, quadrate, or quinate, on pedicels of unequal lengths. Spikelets disarticulating from pedicel at maturity, lanceolate to elliptic, somewhat dorsally compressed, the upper glume facing away from the rachis, 2-flowered; lower glume absent or present as a veinless scale; upper glume 1/5 as long as to less commonly as long as spikelet, 3–5-veined; lower floret sterile, the lemma about as long as spikelet, 5–9-veined, flattened, the palea a minute rudiment, if present; upper floret slightly shorter than to as long as spikelet, the lemma lanceolate to narrowly ovate, soft, flexible, faintly 3-veined, glabrous, pale to nearly black, the margins thin and exposed, folded over the palea. Tropical and warm-temperate regions worldwide; ca. 175 species, ca. 15 in Venezuela, 9 of these in the flora area. Key to the Species of Digitaria 1. 1. 2(1). 2. 3(1). 3. 4(3).

4.

5(4). 5. 6(3). 6. 7(6). 7. 8(7).

Spikelets densely covered and nearly concealed by many silky hairs .... 2 Spikelets glabrous to pubescent but never concealed by silky hairs ....... 3 Spikelets 3.5–4.2 mm long ............................................................ D. insularis Spikelets 2.7–3.3 mm long ................................................................. D. tenuis Spikelets borne in pairs (binate), lanceolate, 2–3.6 mm long .................. 4 Spikelets borne in threes (ternate), at least in the central portion of each branch, elliptic to ovate, 1–2.2 mm long ............................................... 6 Rachis of racemes bearing scattered, elongate, papillae-based hairs; leaf blades finely pubescent throughout; axis of inflorescence 2–8 cm long .............................................................................................. D. horizontalis Rachis of racemes glabrous, not bearing elongate, papillae-based hairs; leaf blades glabrous to sparingly pubescent near base, uncommonly pubescent throughout; axis of inflorescence to 4 cm long .................... 5 Spikelets 2.5–3.5 mm long; lower glume minute, triangular ........ D. ciliaris Spikelets 2–2.4 mm long; lower glume absent .................................. D. nuda Spikelets 1.8–2.2 mm long; upper glume and lower lemma bearing golden brown, clavate hairs .............................................................. D. venezuelae Spikelets 1–1.6 mm long; upper glume and lower lemma glabrous or with whitish, nonclavate hairs ...................................................................... 7 Spikelets 1–1.3 mm long; racemes 4–15 ....................................... D. nervalis Spikelets 1.3–1.6 mm long; racemes 2(3) .................................................. 8 Spikelets glabrous; upper glume, lower lemma, and upper floret usually purple-tipped; upper floret slightly exceeding the lower lemma; plants extensively stoloniferous, usually with densely crowded, reflexed leaf blades ...................................................................................... D. fuscescens

Digitaria 85

8.

Spikelets minutely white-ciliate; upper glume, lower lemma, and upper floret not purple-tipped; upper floret shorter than to equaling the lower lemma; plants tufted to decumbent but never extensively stoloniferous with crowded, reflexed leaf blades .................. D. violascens

Digitaria ciliaris (Retz.) Koeler, Descr. Gram. 27. 1802. —Panicum ciliare Retz., Observ. Bot. 4: 16. 1791 [1786]. Digitaria sanguinalis auct. non (L.) Scop. 1771: sensu Luces, Gramíneas Distr. Fed. 94. 1963. Decumbent, creeping plant of indefinite duration, culms to 40 cm tall; racemes 2–8, each 4–10 cm long; lower lemma of pedicellate spikelet often fimbriate-margined. Weedy places, ca. 50 m; Delta Amacuro (Pedernales), Amazonas. Barinas, Distrito Federal, Falcón, Lara, Miranda, Portuguesa, Táchira, Zulia; widespread throughout tropical and warmtemperate America, Old World tropics. Digitaria fuscescens (J. Presl) Henrard, Meded. Rijks-Herb. 61: 8. 1930. —Paspalum fuscescens J. Presl in C. Presl, Reliq. Haenk. 1: 213. 1830. Plant extensively creeping, mat-forming, culms to 20 cm tall; spikelets 1.2–1.5 mm long, yellowish brown. Old fields, savannas, near sea level to 300 m; occasional in Delta Amacuro, Bolívar, and Amazonas. Occasional elsewhere in Venezuela and throughout tropical America and Africa, native of tropical Asia. ◆Fig. 54. Digitaria horizontalis Willd., Enum. Pl. 92. 1809. Sprawling, decumbent plant, culms 40– 100 cm tall; inflorescence 7–13 cm long, with 7–17 racemes; spikelets 2–2.7 mm long, strongly veined, the lower glume minute, triangular. Pastures and other weedy places, near sea level to 400 m; widespread in Delta Amacuro, Bolívar, and Amazonas. Widespread elsewhere in Venezuela; southeastern U.S.A., Mexico, Central America, West Indies, tropical and warm-temperate South America, western Africa, southern Asia. ◆Fig. 56. Digitaria insularis (L.) Mez ex Ekman, Ark. Bot. 11(4): 17. 1912. —Andropogon insularis L., Syst. Nat. ed. 10, 1304. 1759. —Trichachne insularis (L.) Nees in Mart. et al., Fl. Bras. Enum. Pl. 2: 86. 1829.

Acicarpa sacchariflora Raddi, Agrost. Bras. 31. 1823. —Trichachne sacchariflora (Raddi) Nees in Mart., Fl. Bras. Enum. Pl. 2: 87. 1829. —Digitaria sacchariflora (Raddi) Henrard, Blumea 1: 99. 1934. Coarse cespitose perennial 0.5–3 m tall; inflorescence with branches loosely ascending; spikelets lanceolate, concealed by hairs. Disturbed open areas, 100–800 m; Bolívar (Altiplanicie de Nuria, Cerro Bolívar, El Callao, Guairampai, Río Paramichi). Widespread elsewhere in Venezuela; southern U.S.A., Mexico, Central America, West Indies, tropical and warm-temperate South America. ◆Fig. 57. Digitaria nervalis Henrard, Monogr. Digitaria 490. 1950. Tufted plant, culms 20–40 cm tall; racemes 2–7 cm long on an axis to 3 cm long; spikelets elliptic, glabrous; upper floret brown, apiculate. Sandy pockets among rocks along Río Orinoco, ca. 100 m; Amazonas (Capuana, Puerto Ayacucho). Northeastern Brazil. Digitaria nuda Schumach., Beskr. Guin. Pl. 65. 1827. Decumbent, creeping plant, culms to 50 cm tall; leaf blades glabrous; inflorescence with racemes 2–5 on an axis to 1.5 cm long; spikelets with margins of lower lemma white-ciliate; lower lemma of pedicellate spikelet often fimbriate-margined. Uncommon weed, 100–1200 m; Bolívar (Gran Sabana), Amazonas (Santa Bárbara del Orinoco). Falcón; Mexico, Honduras, West Indies, Guyana, Suriname, French Guiana, Brazil, Bolivia, Paraguay, Argentina, tropical Africa and Asia. Digitaria tenuis (Nees) Henrard, Blumea 1: 99. 1934. —Trichachne tenuis Nees in Mart. et al., Fl. Bras. Enum. Pl. 2: 89. 1829. Cespitose perennial to 50 cm tall; inflorescence of 4–13 racemes, each 5–12 cm long; spikelets nearly concealed by white to tawny hairs. Savannas, ca. 300 m; Bolívar (Alti-

86

P OACEAE

planicie de Nuria). Nueva Esparta, Sucre; Guyana, northeastern Brazil. Digitaria venezuelae Henrard, Monogr. Digitaria 780. 1950. Digitaria glabriculmis Swallen, Fieldiana, Bot. 28: 18. 1951. Cespitose perennial ca. 50 cm tall; inflorescence of 2–5 racemes; spikelets plumply elliptic. Wooded streams in upland savannas, ca. 1000–1100 m; Bolívar (uncommon on Gran Sabana). Lara, Mérida; Suriname, French Guiana.

Fig. 54. Digitaria fuscescens

Digitaria violascens Link, Hort. Berol. 1: 229. 1827. —Ueni-yu. Cespitose perennial 30–70 cm tall, or some culms creeping; spikelets 1.2–1.6 mm long, the upper floret brownish black. Rare weed; Bolívar (Kavanayén). Widespread but uncommon in Venezuela; widespread in tropical and warm-temperate America, native of tropical Asia. ◆Fig. 55.

Fig. 55. Digitaria violascens

Digitaria 87

Fig. 56. Digitaria horizontalis

88

P OACEAE

Fig. 57. Digitaria insularis

Echinochloa 89

25. ECHINOCHLOA P. Beauv., Ess. Agrostogr. 53. 1812, nom. cons. [Subfamily Panicoideae, Tribe Paniceae] by Emmet J. Judziewicz Cespitose to rhizomatous annuals or perennials, some species aquatic or semiaquatic with aerenchymatous culms. Leaves with ligules absent or ciliate; blades linear to lanceolate. Inflorescence a panicle of racemes, the racemes varying from regularly flowered to irregular and bushy, the spikelets paired in 2 rows or irregularly fascicled. Spikelets elliptic to ovate, terete, falling entire, green or often blotched with purple, 2-flowered; glumes and lower lemma often appressedhispidulous along the veins; lower glume much shorter than the spikelet, ovate, acuminate to apiculate, several-veined; upper glume and especially the lower lemma acuminate to long-awned, as long as the spikelet, 5–7-veined, the upper glume rounded on the back, the lower lemma somewhat sulcate, with some of the lateral veins evident at the base and apex but obscure in the center; lower floret sterile or staminate, with a well-developed palea; upper floret elliptic to ovate, coriaceous, straw-colored, smooth to striate, shining, slightly apiculate, the margins of the lemma not strongly inrolled over the margins of the flat, slightly beaked palea. Temperate and tropical regions worldwide; 30–40 species, 3 in Venezuela, all in the flora area. Key to the Species of Echinochloa 1. 1. 2(1). 2.

Ligule present, ciliate, 2–4 mm long; spikelets 4–6 mm long (excluding awns), the lower floret staminate ....................................... E. polystachya Ligule absent; spikelets 2.3–3.3 mm long, the lower floret sterile .......... 2 Spikelets 2.3–2.6 mm long, awnless, in 4 regular rows in racemes 1.5– 3 cm long ....................................................................................... E. colona Spikelets 2.5–3.3 mm long, with awns 2–30 mm long, in irregular rows in racemes 3–7 cm long ........................................................... E. crus-pavonis

Echinochloa colona (L.) Link, Hort. Berol. 2: 209. 1833. —Panicum colonum L., Syst. Nat. ed. 10, 870. 1759. Tufted annual, culms decumbent and rooting at nodes; inflorescence 5–15 cm long, of 5–10 erect racemes. Stream banks, ditches, near sea level to 200 m; Delta Amacuro (common), Bolívar (El Dorado, La Vergareña, Upata), Amazonas (Puerto Ayacucho). Widespread elsewhere in Venezuela and the Neotropics. Echinochloa crus-pavonis (Kunth) Schult., Mantissa 2: 269. 1824. —Oplismenus crus-pavonis Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 108. 1815 [1816]. Decumbent, creeping and rooting plant, erect culms to 1 m tall, 1 cm diameter; leaf blades 10–30 × 0.5–2 cm. Stream sides, ditches, near sea level to 1000 m; Delta

Amacuro (common), Bolívar (Gran Sabana), Amazonas (Puerto Ayacucho). Widespread elsewhere in Venezuela and the Neotropics. ◆Fig. 59. Echinochloa polystachya (Kunth) Hitchc., Contr. U.S. Natl. Herb. 22: 135. 1920. —Oplismenus polystachyus Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 107. 1815 [1816]. —Paja pelua, Pasto chiguirera. Floating, extensively mat-forming aquatic, the culms spongy, rooting at nodes, erect portions 1–3 m tall, 1–2 cm diameter; leaf blades 35–75 × 1–2.5 cm. Rivers, ponds, near sea level to 50 m; Delta Amacuro (common), Amazonas (Isla Carestía in Río Orinoco). Anzoátegui, Falcón, Guárico, Portuguesa, Táchira, Zulia; widespread in the Neotropics. ◆Fig. 58.

90

P OACEAE

Fig. 58. Echinochloa polystachya

Echinochloa 91

Fig. 59. Echinochloa crus-pavonis

92

P OACEAE

26. ECHINOLAENA Desv., J. Bot. Agric. 1: 75. 1813. [Subfamily Panicoideae, Tribe Paniceae] by Emmet J. Judziewicz Annuals or perennials. Leaves with ligules ciliate; blades linear to more commonly lanceolate. Inflorescence of 1 (rarely several) deflexed, pectinate racemes, if solitary then subtended by a small bract; raceme(s) densely flowered, the spikelets in 2 rows on one side of the flattened rachis, the rachis terminating in a spikelet with a lower glume that simulates a sterile prolongation of the rachis. Spikelets lanceolate, awnless, 2-flowered; glumes rigid, often arching and inflexed, narrowly keeled, ribbed, attenuate, ± densely tuberculate-ciliate; lower glume 1/2 to as long as spikelet; upper glume about as long as to more commonly much shorter than the lower; lower floret staminate, membranous, the lemma about as long as the upper glume, the palea well developed; upper floret shorter than the lower floret, elliptic, acute, smooth, straw-colored, shining, the lemma with margins winged near the base, functioning as elaiosomes (oil-bearing bodies adapted for ant-dispersal). Central America, tropical South America, Madagascar; 7 species, 2 in Venezuela, both in the flora area. Key to the Species of Echinolaena 1. 1.

Leaf blades 20–40 × 4–6 mm; inflorescence 1.3–2.3 cm long ........ E. gracilis Leaf blades 50–100 × 5–15 mm; inflorescence (2–)2.5–5 cm long .....E. inflexa

Echinolaena gracilis Swallen, J. Wash. Acad. Sci. 23: 457. 1933. Decumbent, trailing, and rooting, the culms to 20 cm tall; raceme solitary, partially included in leaf sheath; spikelets 7–10 mm long. Savannas, 100–200 m; Bolívar (La Vergareña), Amazonas (Caño Guaviarito on Río Manapiare). Apure, Cojedes, Guárico; Central America, Colombia, Brazil, Bolivia. ◆Fig. 61. Echinolaena inflexa (Poir.) Chase, Proc. Biol. Soc. Wash. 24: 117. 1911. —Cenchrus inflexus Poir. in Lam., Encycl. 6:

50. 1804. —Saeta. Sprawling, decumbent, and rooting, the culms to 2 m long, sometimes scandent; raceme solitary, pedunculate, often deflexed; spikelets 8–15 mm long. Sandy savannas, lajas, wet open depressions, Mauritia palm swamps, lower slopes of tepuis on edges of forests, ant-dispersed and often growing on termite mounds, 100–1600 m; Bolívar (common on Gran Sabana), Amazonas (common in lowlands). Apure, Lara, Táchira; Colombia, Guyana, Suriname, French Guiana, Peru, Brazil (widespread), Bolivia. ◆Fig. 60.

27. ELEUSINE Gaertn., Fruct. Sem. Pl. 1: 7. 1788. [Subfamily Chloridoideae, Tribe Cynodonteae] by Emmet J. Judziewicz Cespitose to stoloniferous annuals or perennials. Leaves with linear blades. Inflorescence terminal, consisting of several to many digitate to more commonly subdigitate, densely flowered spikes; spikes flattened, unilateral, bearing 2 rows of spikelets on one side. Spikelets strongly laterally compressed, several-flowered, awnless, the uppermost florets reduced or rudimentary, disarticulation occurring above the glumes and between the florets; glumes shorter than the lowest floret, unequal, lanceolate, the lower 1-veined, the upper 3–5-veined; lemmas lanceolate, 3–5-veined; paleas shorter than lemmas; stamens 3; stigmas 2. Pantropics; 8 species, 1 in Venezuela.

Echinolaena 93

Fig. 60. Echinolaena inflexa

Fig. 61. Echinolaena gracilis

94

P OACEAE

Eleusine indica (L.) Gaertn., Fruct. Sem. Pl. 1: 8. 1788. —Cynosurus indicus L., Sp. Pl. 72. 1753. Annual, culms sprawling, creeping, erect portions 10–50 cm tall; leaves with sheaths strongly laterally compressed; inflorescence of 2–10 subdigitate racemes each 3–15 cm

long; spikelets 4–6 mm long, ovate, 4–7-flowered. Weedy in pastures, towns, and roadsides, 50–1100 m; widespread in Delta Amacuro, Bolívar, and Amazonas. Widespread elsewhere in Venezuela and worldwide in tropical and warm-temperate regions. ◆Fig. 62.

28. ELIONURUS Humb. & Bonpl. ex Willd., Sp. Pl. 4: 941. 1806. [Subfamily Panicoideae, Tribe Andropogoneae] by Emmet J. Judziewicz Cespitose perennials or rarely annuals. Inflorescences terminal and axillary, each consisting of an elongate peduncle bearing a solitary, erect, spicate raceme with a ± flexuous and strongly pilose rachis; rachis internodes disarticulating along with the sessile and pedicellate spikelets. Sessile spikelet with base prolonged into a prominent, oblique callus; lower glume firmly membranous, many-veined, strongly pilose all across the back or at least ciliate on the prominent marginal veins, the margins inflexed over the margins of the upper glume and florets, the apex bilobed, short-aristate, or entire; upper glume narrower than the lower glume, strongly keeled, 1–3-veined, usually pubescent on the back; florets hyaline, distinctly shorter than the spikelet, awnless, lacking a palea, the lower sterile, the upper bisexual. Pedicellate spikelet ± similar to sessile spikelet, but sterile or staminate, awnless to short-aristate. Tropical America, Africa, southwestern Asia, Australia; ca. 15 species, 1 in Venezuela. Elionurus muticus (Spreng.) Kuntze, Revis. Gen. Pl. 3(3): 350. 1898. —Lycurus muticus Spreng., Syst. Veg. 4(2): 32. 1827. Andropogon adustus Trin., Mém. Acad. Imp. Sci. St.-Pétersbourg, Sér. 6, Sci. Math. 2: 259. 1832. —Elionurus adustus (Trin.) Ekman, Ark. Bot. 13(10): 6. 1913, “Elyonurus.” Elionurus planifolius Renvoize, Kew Bull. 32: 669. 1978, “Elyonurus.”

Densely cespitose perennial 40–100 cm tall, from a hard, knotty base, the sheaths of the old leaves persistent, often burnt; inflorescence 3–7 cm long; subsessile spikelets 4– 5.5 mm long, ovate, ± densely pubescent. Savannas, often dominant, 100–1400 m; Bolívar (widespread), Amazonas (30 km north of Puerto Ayacucho, upper Río Ventuari). Widespread elsewhere in Venezuela; throughout tropical America and Africa. ◆Fig. 63.

29. ENTEROPOGON Nees in Lindl., Intr. Nat. Syst. Bot. ed. 2, 448. 1836. [Subfamily Chloridoideae, Tribe Cynodonteae] by Emmet J. Judziewicz Annuals or perennials. Leaves with ligules ciliate-membranous; blades linear. Inflorescence terminal, consisting of 1–several, subdigitate or approximate, often elongate, spreading or drooping racemes. Spikelets ascending, often somewhat crowded, 1- or 2-flowered, the lowest floret bisexual, the upper floret if present rudimentary, sterile; glumes subulate to lanceolate, subequal, 1-veined, divergent, acute to subaristate, the upper shorter than to about as long as the functional floret; functional floret noticeably dorsally compressed, 3-veined, minutely bifid, awned from between the teeth, disarticulating from the glumes; palea slightly shorter than

Eleusine 95

Fig. 62. Eleusine indica

96

P OACEAE

Fig. 63. Elionurus muticus

Fig. 64. Enteropogon mollis

Eragrostis 97

the lemma; rachilla internode ending in rudimentary floret(s) represented by awns; stamens 3; stigmas 2. Caryopsis dorsally compressed. Pantropics; 17 species, 1 in Venezuela. Enteropogon mollis (Nees) Clayton, Kew Bull. 37: 419. 1982. —Gymnopogon mollis Nees in Mart., Fl. Bras. Enum. Pl. 2: 427. 1829. —Chloris mollis (Nees) Swallen, N. Amer. Fl. 17(8): 596. 1939. Tufted perennial ca. 50 cm tall; inflorescence of 4–12 approximate, slender racemes 4–8 cm long; spikelets with functional floret 3–4.5 mm long, surmounted by an awn 3–6

mm long. Lowland forests, 100–200 m; Amazonas (between La Esmeralda and Cerro Duida). Aragua, Distrito Federal, Falcón, Lara, Zulia; Honduras, Nicaragua, Greater Antilles, Colombia, Ecuador, Peru, northeastern Brazil. ◆Fig. 64. The single flora area collection of Enteropogon mollis is overmature and is only tentatively identified as this species.

30. ERAGROSTIS Wolf, Gen. Pl. 23. 1776. [Subfamily Chloridoideae, Tribe Cynodonteae] Megastachya P. Beauv., Ess. Agrostogr. 74, 167. 1812. Neeragrostis Bush, Trans. Acad. Sci. St. Louis 13(7): 178. 1903. Erosion Lunell, Amer. Midl. Naturalist 4: 221. 1915. Diandrochloa De Winter, Bothalia 7(2): 387. 1960. Roshevitzia Tzvelev, Bot. Zurn. (Moscow & Leningrad) 53: 311. 1968. by Gerrit Davidse Annuals or perennials; cespitose, stoloniferous or rhizomatous. Ligule usually a line of hairs, rarely a membrane or ciliate membrane; blades linear. Inflorescence an open to spicate or glomerulate panicle. Spikelets laterally compressed, 2–manyflowered; disarticulation various, primarily (tropical America) from the base upward, the glumes falling first from the persistent rachilla followed by the lemma and caryopsis, the palea usually persistent, or the disarticulation from the tip downward, the rachilla breaking between the florets, the lemma, palea, and caryopsis falling together; florets reduced upward; glumes subequal, usually shorter than the lowest florets, usually lanceolate (as folded) or subulate, acute, usually 1veined, rarely 3-veined or the lower veinless, usually scaberulous or scabrous on the veins, never persistent; lemmas 3-veined, sharply or broadly keeled, herbaceous to membranous, usually glabrous or scaberulous, entire, obtuse or emarginate to acuminate, the lateral veins faint or conspicuous; palea keels ciliate, scabrous or glabrous; flowers usually bisexual, sometimes unisexual; stamens 2 or 3. Fruit a caryopsis, free from the lemma and palea; embryo usually ca. 1/2 as long as the caryopsis; hilum punctate. Cosmopolitan, especially common in the tropical and subtropical regions of the world; ca. 350 species, 27 in Venezuela, 14 of these in the flora area. Key to the Species of Eragrostis 1. 1. 2(1). 2.

Palea prominently papillose-ciliate on the keels; annuals ...................... 2 Palea glabrous to scabrous on the keels; annuals or perennials ............. 4 Anthers 2; panicle dense, spicate-cylindrical but interrupted below, the branches appressed ..................................................................... E. ciliaris Anthers 3; panicle open, oblong- to ovoid-cylindrical, the branches spreading ................................................................................................ 3

98

P OACEAE

3(2). 3. 4(1). 4. 5(4). 5. 6(5). 6. 7(6). 7. 8(7).

8.

9(7). 9.

10(9). 10. 11(10). 11. 12(11). 12. 13(11). 13.

Culms, leaves, and panicles not viscid; spikelets 0.9–1.4 mm wide; lemmas 0.8–1 mm long ................................................................... E. amabilis Culms, leaves, and panicles viscid; spikelets 1.5–2 mm wide; lemmas 1.1– 1.8 mm long .................................................................................. E. viscosa Ligule an eciliate membrane ......................................................... E. japonica Ligule a line of hairs .................................................................................. 5 Plants stoloniferous, the stolons rooting at the nodes, often forming a mat on recently exposed soil; longest leaf blades to 3.5 cm long .... E. hypnoides Plants cespitose, lacking stolons or rhizomes; longest leaf blades always > 6 cm ..................................................................................................... 6 Caryopsis grooved on the side opposite the embryo, apically truncate ................................................................................................. E. polytricha Caryopsis flat to obtuse on the side opposite the embryo, apically obtuse ................................................................................................................ 7 Lemmas acuminate to sharply acute, strongly keeled ............................. 8 Lemmas subacute to acute, indistinctly keeled to rounded on the back .... 9 Spikelets 1.1–1.8 mm wide; lemmas 1.7–2.4 mm long, the tips straight; paleas 4/5–5/6 as long as the lemmas, scabrous on the keels; caryopsis round in cross section, oblong-lanceolate in outline .............. E. acutiflora Spikelets 1.9–3.5 mm wide; lemmas 2.3–2.8 mm long, the tips divergent; palea 1/2–2/3 as long as the lemma, ciliolate on the keels; caryopsis oblong in cross section, oblong-ovate in outline .................... E. maypurensis Spikelets ovate to oblong-ovate, the lemmas widely divergent; caryopsis narrowly oblong in cross section, strongly flattened ............. E. unioloides Spikelets linear to oblong-lanceolate, the lemmas ascending; caryopsis round to elliptic in cross section, often wider than thick, but not strongly flattened ................................................................................ 10 Plants perennial; anthers 3 ........................................................ E. atrovirens Plants annual; anthers 2 or 3 .................................................................. 11 Anthers 2 .................................................................................................. 12 Anthers 3 .................................................................................................. 13 Spikelets 0.9–1.5 mm wide; pedicels appressed; plants 50–75 cm tall .................................................................................................. E. gangetica Spikelets 2–2.7 mm wide; pedicels divergent; plants 10–20 cm tall ................................................................................................ E. guianensis Spikelets 1.2–1.9 mm wide ......................................................... E. pectinacea Spikelets 0.8–1.1 mm wide ................................................................ E. pilosa

Eragrostis acutiflora (Kunth) Nees in Mart. et al., Fl. Bras. Enum. Pl. 2: 501. 1829. —Poa acutiflora Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 161. 1815 [1816]. —Megastachya acutiflora (Kunth) Roem. & Schult., Syst. Veg. 2: 588. 1817. Eragrostis acutiflora var. humilior J. Presl, Reliq. Haenk. 1: 277. 1830. Cespitose, short-lived perennial; anthers 2, 0.2–0.3 mm. Disturbed areas in savannas, high, open river banks, near sea level to 500

m; Delta Amacuro (Tucupita), Bolívar (widespread), Amazonas (Puerto Ayacucho). Apure, Guárico, Mérida, Monagas, Portuguesa, Táchira, Zulia; Mexico, Central America, Caribbean, Colombia, Guyana, Suriname, French Guiana, Ecuador, Brazil, Bolivia. ◆Fig. 69. Eragrostis amabilis (L.) Wight & Arn. ex Nees in Hook. & Arn., Bot. Beechey Voy. 251. 1838. —Poa amabilis L., Sp. Pl. 68. 1753.

Eragrostis 99

Poa tenella L., Sp. Pl. 69. 1753. —Eragrostis tenella (L.) P. Beauv. ex Roem. & Schult., Syst. Veg. 2: 576. 1817. Poa plumosa Retz., Observ. Bot. 4: 20. 1786. —Eragrostis amabilis var. plumosa (Retz.) E.G. Camus & A. Camus, Fl. Indo-Chine 7: 557. 1923. —Eragrostis plumosa (Retz.) Link, Hort. Berol. 1: 192. 1827. —Eragrostis tenella var. plumosa (Retz.) Stapf in Hook. f., Fl. Brit. India 7: 315. 1896. Eragrostis ciliaris var. patens Chapm. ex Beal, Grass. N. Amer. 2: 479. 1896. Glandular, but not viscid, annual weed. Roadsides, gardens, disturbed secondary areas around towns, 50–500 m; Bolívar (Ciudad Bolívar, Urimán), Amazonas (Puerto Ayacucho). Falcón, Mérida, Portuguesa, Táchira, Zulia; naturalized throughout the New World tropics and subtropics, native to the Old World tropics. Eragrostis atrovirens (Desf.) Trin. ex Steud., Nomencl. Bot. ed. 2, 1: 562. 1840. —Poa atrovirens Desf., Fl. Atlant. 1: 73, t. 14. 1798. Poa chariis Schult., Mantissa 2: 314. 1824. —Eragrostis chariis (Schult.) Hitchc., Lingnan Sci. J. 7: 193. 1929 [1931]. Glaucous perennial; anthers 3, 0.3–0.4 mm. Rocky river banks, 1100–1200 m; Bolívar (Gran Sabana, Río Arabopó). Naturalized in U.S.A., Mexico, Guatemala, Belize, Nicaragua, Caribbean, Colombia, Ecuador, Peru, Bolivia, native to Africa and Asia. Eragrostis ciliaris (L.) R. Br. in Tuckey, Narr. Exped. Zaire 478. 1818. —Poa ciliaris L., Syst. Nat. ed. 10, 2: 875. 1759. Pantropical weed native to the Old World tropics; 2 varieties, both in Venezuela, 1 of these in the flora area. Variety brachystachya Boiss., with a short, dense, compact panicle, is restricted to the arid coastal areas of northern Venezuela. E. ciliaris var. ciliaris Annual weed with an elongate, basally interrupted spicate panicle. Roadsides, disturbed secondary areas, near sea level to 200 m; common in Delta Amacuro, Bolívar, and Amazonas. Probably in all Venezuelan states; pantropical weed native to the Old World tropics.

Eragrostis gangetica (Roxb.) Steud., Syn. Pl. Glumac. 1: 266. 1855 [1854]. —Poa gangetica Roxb., Fl. Ind. ed. 1820, 1: 341. 1820. Poa elegantula Kunth, Révis. Gramin. 1: 114. 1829. —Eragrostis elegantula (Kunth) Nees ex Steud., Syn. Pl. Glumac. 1: 266. 1855 [1854]. Eragrostis stenoclada J. Presl, Reliq. Haenk. 1: 278. 1830. Eragrostis flamignii De Wild., Bull. Jard. Bot. État 6: 63, t. 1. 1919. Cespitose annual; anthers 2, 0.2–0.3 mm. Marsh margins, fields, roadsides, savannas, 50–300 m; Bolívar (Caicara, Cacurí, Hato la Vergareña). Naturalized in southern U.S.A., Belize, native to tropical Africa and Asia. Eragrostis gangetica is probably recently introduced and is uncommon. Eragrostis guianensis A. Hitchc., Contr. U.S. Natl. Herb. 22: 454. 1922. Delicate cespitose annual; anthers 2, 0.2– 0.3 mm. Open, sandy areas in savannas, 50– 300 m; Bolívar (Caicara, Urimán), Amazonas (Puerto Ayacucho). Anzoátegui, Monagas; Guyana, French Guiana, Brazil (Roraima). ◆Fig. 65. Eragrostis hypnoides (Lam.) Britton, Sterns & Poggenb., Prelim. Cat. 69. 1888. —Poa hypnoides Lam., Tabl. Encycl. 1: 185. 1791. —Megastachya hypnoides (Lam.) P. Beauv., Ess. Agrostogr. 74, 167, 175. 1812. —Neeragrostis hypnoides (Lam.) Bush, Trans. Acad. Sci. St. Louis 13: 180. 1903. —Erosion hypnoides (Lam.) Lunell, Amer. Midl. Naturalist 4: 221. 1915. Eragrostis reptans var. contracta Döll in Mart., Fl. Bras. 2(3): 148. 1878. Eragrostis reptans var. laxior Döll in Mart., Fl. Bras. 2(3): 148. 1878. Eragrostis reptans var. pygmaea Döll in Mart., Fl. Bras. 2(3): 149. 1878. Megastachya breviflora E. Fourn., Mexic. Pl. 2: 119. 1886. Megastachya corymbifera E. Fourn., Mexic. Pl. 2: 119. 1886. Megastachya gouinii E. Fourn., Mexic. Pl. 2: 119. 1886. Diminutive, creeping annual, often forming mats. River banks, sand bars, exposed lake margins, near sea level to 200 m; com-

100

P OACEAE

mon in Delta Amacuro, Bolívar, and Amazonas. Apure, Barinas, Monagas; Canada, U.S.A., Mexico, Central America, Caribbean, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina, Uruguay. Eragrostis hypnoides is a short-lived annual in which the spikelets become more numerous and more prominent than the leaves. In extreme cases plants appear to consist almost entirely of numerous-flowered spikelets. Eragrostis japonica (Thunb.) Trin., Mém. Acad. Imp. Sci. St. Pétersbourg, Sér. 6, Sci. Math. 1(4): 405. 1830. —Poa japonica Thunb., Fl. Jap. 51. 1784. —Eragrostis tenella var. japonica (Thunb.) Roem. & Schult., Syst. Veg. 2: 576. 1817. —Diandrochloa japonica (Thunb.) A.N. Henry, Bull. Bot. Surv. India 9: 290. 1968. —Roshevitzia japonica (Thunb.) Tzvelev, Novosti Sist, Vysš. Rast. 7: 50. 1970 [1971]. Eragrostis glomerata (Walter) L. Dewey, Contr. U.S. Natl. Herb. 2: 543. 1894. —Poa glomerata Walter, Fl. Carol. 1: 80. 1788. —Diandrochloa glomerata (Walter) Burkart, Bol. Soc. Argent. Bot. 12: 287. 1968. —Roshevitzia glomerata (Walter) Tzvelev, Novosti Sist, Vysš. Rast. 7: 50. 1970 [1971]. Eragrostis aturensis (Kunth) Trin. ex Steud., Nomencl. Bot. ed. 2, 1: 562. 1840. —Poa aturensis Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 161. 1815 [1816]. Eragrostis conferta (Elliott) Trin., Mém. Acad. Imp. Sci. St. Pétersbourg, Sér. 6, Sci. Math. 1(4): 409. 1830. —Poa conferta Elliott, Sketch Bot. S. Carolina 1: 158. 1816. Eragrostis hapalantha Trin., Mém. Acad. Imp. Sci. St. Pétersbourg, Sér. 6, Sci. Math. 1(4): 409. 1830. Eragrostis diplachnoides Steud., Syn. Pl. Glumac. 1: 268. 1855 [1854]. —Eragrostis interrupta var. diplachnoides (Steud.) Stapf in Hook. f., Fl. Brit. India 7: 316. 1896. —Diandrochloa diplachnoides (Steud.) A.N. Henry, Bull. Bot. Surv. India 9: 290. 1968. —Eragrostis namaquensis var. diplachnoides (Steud.) Clayton, Kew Bull. 25(2): 251. 1971.

Eragrostis interrupta var. laxiflora Döll in Mart., Fl. Bras. 2(3): 158. 1878. Eragrostis interrupta var. parviflora Döll in Mart., Fl. Bras. 2(3): 158. 1878. Eragrostis pallida Vasey, Contr. U.S. Natl. Herb. 1: 285. 1893. Annual weed to 1 m tall. River banks, sand bars, roadsides, weedy fields, near sea level to 200 m; common in Delta Amacuro, Bolívar, and Amazonas. Apure, Barinas, Guárico, Yaracuy; pantropical weed native to eastern Asia. ◆Fig. 68. Eragrostis japonica is the only Venezuelan species of Eragrostis with a membranous ligule. This character has been used to segregate the small cosmopolitan genus Diandrochloa from Eragrostis, which has mostly ciliate ligules. However, the existence of other Eragrostis species with ciliate membranous ligules diminishes the importance of the character. Eragrostis maypurensis (Kunth) Steud., Syn. Pl. Glumac. 1: 276. 1855 [1854]. —Poa racemosa Vahl, Eclog. Amer. 1: 7. 1796, non Thunb. —Poa maypurensis Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 161. 1815 [1816]. —Megastachya maypurensis (Kunth) Roem. & Schult., Syst. Veg. 2: 588. 1817. Eragrostis vahlii (Roem. & Schult.) Nees in Mart. et al., Fl. Bras. Enum. Pl. 2: 499. 1829. —Poa vahlii Roem. & Schult., Syst. Veg. 2: 563. 1817. Eragrostis amoena J. Presl, Reliq. Haenk. 1: 275. 1830. Eragrostis panamensis J. Presl, Reliq. Haenk. 1: 277. 1830. Poa meratiana Kunth, Révis. Gramin. 2: 539, t. 184. 1832. —Eragrostis maypurensis var. meratiana (Kunth) Pilg., Bot. Jahrb. Syst. 70: 348. 1939. —Eragrostis meratiana (Kunth) Steud., Syn. Pl. Glumac. 1: 276. 1855 [1854]. Eragrostis lindeniana Steud., Syn. Pl. Glumac. 1: 278. 1855 [1854]. Eragrostis acuminata Döll in Mart., Fl. Bras. 2(3): 153. 1878. Eragrostis vahlii var. polyantha Döll in Mart., Fl. Bras. 2(3): 155. 1878. Eragrostis maypurensis var. densiuscula Pilg., Bot. Jahrb. Syst. 70: 349. 1939. Common cespitose annual; anthers 2, 0.2– 0.4 mm. Disturbed areas in savannas, road-

Eragrostis 101

sides, 50–1100 m; common in Bolívar and Amazonas. Anzoátegui, Apure, Aragua, Barinas, Falcón, Guárico, Monagas, Portuguesa, Sucre, Yaracuy, Zulia; Mexico, U.S.A., Central America, Colombia, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia. Eragrostis pectinacea (Michx.) Nees, Fl. Afr. Austral. Ill. 406. 1841. —Poa pectinacea Michx., Fl. Bor.-Amer. 1: 69. 1803. Canada, U.S.A., Mexico, Guatemala, Honduras, El Salvador, Nicaragua, Costa Rica, Panama, Caribbean, Venezuela, Suriname, French Guiana, Ecuador, Peru, Brazil, Paraguay, Argentina, Uruguay; 2 varieties, both in Venezuela, 1 of these in the flora area. Variety miserrima (Fourn.) Reeder, distinguished by its divergent pedicels, has been reported from the Guianas but has not yet been found in the flora area, although it is to be expected there. E. pectinacea var. pectinacea Cespitose annual with the pedicels appressed to the panicle; anthers 3, 0.2–0.4 mm. Weedy open areas, 50–1400 m; Bolívar (Gran Sabana), Amazonas (Puerto Ayacucho). Aragua, Distrito Federal, Falcón, Lara, Portuguesa, Táchira; Canada, U.S.A., Mexico, Guatemala, Honduras, El Salvador, Nicaragua, Costa Rica, Panama, Caribbean, Suriname, French Guiana, Ecuador, Peru, Brazil, Paraguay, Argentina, Uruguay. Eragrostis pilosa (L.) P. Beauv., Ess. Agrostogr. 71, 162, 175. 1812. —Poa pilosa L., Sp. Pl. 68. 1753. Eragrostis filiformis Link, Hort. Berol. 1: 191. 1827. Poa linkii Kunth, Révis. Gramin. 1: 113. 1829. —Eragrostis linkii (Kunth) Steud., Syn. Pl. Glumac. 1: 273. 1855 [1854]. Cespitose annual; lower panicle branches whorled; anthers 3, 0.2–0.3 mm. Open, disturbed weedy areas, open river banks, 50– 300 m; Bolívar (Ciudad Bolívar, Río Cinaruco), Amazonas (vicinity of Puerto Ayacucho). Falcón, Guárico, Lara, Zulia; Canada, U.S.A., Mexico, Honduras, El Salvador, Nicaragua, Costa Rica, Caribbean, Guyana, French Guiana, Ecuador, Peru, Brazil, Bolivia, Chile, Paraguay, Argentina, Uruguay,

native to Eurasia, naturalized on all continents except Antarctica. Eragrostis polytricha Nees in Mart. et al., Fl. Bras. Enum. Pl. 2: 507. 1829. —Poa polytricha (Nees) Kunth, Enum. Pl. 1: 331. 1833. Eragrostis polytricha var. glabrior Döll in Mart., Fl. Bras. 2(3): 140. 1878. Eragrostis polytricha var. hirsutior Döll in Mart., Fl. Bras. 2(3): 140. 1878. Eragrostis trichocolea Hack. & Arechav., Anales Mus. Nac. Montevideo 1: 44. 1896. Eragrostis floridana Hitchc., Amer. J. Bot. 2: 308. 1915. —Eragrostis trichocolea var. floridana (Hitchc.) Witherspoon, Ann. Missouri Bot. Gard. 64: 328. 1977. Eragrostis purpusii Jedwabn., Bot. Arch. 5(3–4): 201. 1924. Eragrostis fragilis Swallen, Fieldiana, Bot. 28: 18. 1951. Cespitose perennial; inflorescence becoming a tumble weed; anthers 3, ca. 0.5 mm. Trachypogon-Curatella savannas, 300–1300 m; Bolívar (Altiplanicie de Nuria, Gran Sabana, vicinity of El Manteco, Upata). Anzoátegui, Distrito Federal, Falcón, Lara, Monagas; U.S.A., Mexico, Guatemala, Belize, Honduras, Nicaragua, Colombia, Guyana, Brazil, Bolivia, Chile, Paraguay, Argentina, Uruguay. Eragrostis unioloides (Retz.) Nees ex Steud., Syn. Pl. Glumac. 1: 264. 1855 [1854]. —Poa unioloides Retz., Observ. Bot. 5: 19. 1788. Cespitose annual; anthers 2 or 3, 0.2–0.3 mm. Disturbed roadsides and secondary vegetation, sandy river banks, 100–1100 m; becoming common in Bolívar. Naturalized in southern U.S.A., Mexico, Panama, the Caribbean, Colombia, Guyana, French Guiana, Ecuador, and tropical West Africa, native to the Asian tropics. ◆Fig. 66. Eragrostis viscosa (Retz.) Trin., Mém. Acad. Imp. Sci. St.-Pétersbourg, Sér. 6, Sci. Math. 1: 397. 1830. —Poa viscosa Retz., Observ. Bot. 4: 20. 1786. —Eragrostis tenella var. viscosa (Retz.) Stapf in Hook. f., Fl. Brit. India 7: 315. 1896. Poa viscosa var. pilosissima Hochst. ex A. Rich., Tent. Fl. Abyss. 2: 424. 1850. —Eragrostis viscosa var. pilosissima

102

P OACEAE

(Hochst. ex A. Rich.) Hochst., Flora 38: 329. 1855. Eragrostis warmingii Hack., Oesterr. Bot. 2. 52: 305. 1902. Glandular, viscid annual weed with soil particles and dehisced anthers usually attached to the foliage and panicle branches. Roadsides, disturbed open areas, 50–700 m; Bolívar (Canaima, Ciudad Piar, Hato La Vergeña, Maripa). Anzoátegui, Apure, Aragua, Falcón, Guárico, Lara, Nueva Esparta,

Fig. 65. Eragrostis guianensis

Sucre, Zulia; naturalized in Mexico, Guatemala, Honduras, El Salvador, Nicaragua, Costa Rica, Panama, and Ecuador, native to the Old World tropics. ◆Fig. 67. Eragrostis viscosa has variable inflorescence morphology. An extreme form, E. warmingii, previously identified from Bolívar, with dense inflorescences is not recognized because of the existence of intermediate inflorescence morphologies in numerous specimens.

Fig. 66. Eragrostis unioloides

Eragrostis 103

Fig. 67. Eragrostis viscosa

Fig. 68. Eragrostis japonica

Fig. 69. Eragrostis acutiflora

104

P OACEAE

31. ERIOCHLOA Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 94. 1815 [1816]. [Subfamily Panicoideae, Tribe Paniceae] by Emmet J. Judziewicz Annuals or perennials. Leaves with ligules short-membranous, long-ciliate; bracts linear. Inflorescence of 1–many racemes or loosely rebranching racemose branches; spikelets solitary or paired, borne on the lower side of the rachis, disarticulating below the glumes. Spikelets lanceolate to ovate, dorsally compressed, 2flowered; lower glume and the thickened internode between the glumes united to form a knob-like swelling at the base of the spikelet, the lower glume therefore absent or reduced to a minute cuff-like scale; upper glume as long as the spikelet, 5veined; lower floret slightly shorter than the spikelet, sterile, the lemma similar to the upper glume, the palea absent to well developed; upper floret distinctly shorter than the spikelet, typically elliptic, cartilaginous, straw-colored, glabrous, and granular-striate, the lemma mucronate or bearing an abrupt, delicate awn, its margins loosely covering the margins of the palea. Tropical and warm-temperate areas worldwide; ca. 30 species, 3 in Venezuela, 2 of these in the flora area. Key to the Species of Eriochloa 1. 1.

Inflorescence of 1–3 unbranched racemes; spikelets 3–3.5 mm long .................................................................................................. E. distachya Inflorescence of 8–20 branched racemes; spikelets 4–5 mm long ................................................................................................... E. punctata

Eriochloa distachya Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 95, t. 30. 1815 [1816]. Tufted annual 30–50 cm tall; leaf blades 2–5 mm wide; racemes 1.5–3 cm long, the rachis with spreading silky hairs. Savannas, 50–200 m; Bolívar (La Vergareña, Upata), Amazonas (Caño Guaviarito on Río Manapiare). Aragua, Carabobo, Distrito Federal, Guárico, Mérida, Miranda; Central America, Colombia, Ecuador, Peru, Brazil, Bolivia, Paraguay. ◆Fig. 70.

Eriochloa punctata (L.) Desv. ex Ham., Prodr. Pl. Ind. Occid. 5. 1825. —Milium punctatum L., Syst. Nat. ed. 10, 872. 1759. Cespitose perennial 50–150 cm tall; leaf blades 6–15 mm wide; racemes 2–4 cm long, the rachis sparingly appressed-pubescent. Steamsides, pastures, wet savannas, near sea level to 100 m; Delta Amacuro (mouth of Caño Guiniquina, Isla de Tortola), Amazonas (Isla Carestía in Río Orinoco). Aragua, Barinas, Distrito Federal, Guárico, Lara, Portuguesa, Yaracuy; southeastern U.S.A., Mexico, Central America, West Indies, tropical South America.

32. ERIOCHRYSIS P. Beauv., Ess. Agrostogr. 8. 1812. [Subfamily Panicoideae, Tribe Andropogoneae] by Emmet J. Judziewicz Cespitose perennials. Inflorescence a contracted, fuzzy, yellowish to brown panicle with numerous, short, erect branches; rachilla internodes short, slender, with flaring, cupulate apices, falling attached at base to both the sessile and pedicellate spikelets or often the pedicellate spikelet falling first. Spikelets awnless,

Eriochrysis 105

Fig. 70. Eriochloa distachya

Fig. 71. Eriochrysis aff. holcoides

106

P OACEAE

similar in aspect; glumes as long as the spikelet and concealing the florets, coriaceous, pale, the veins evident or not, bearing lines of spreading, tawny or dark brown cilia; lower glume elliptic, often 7-veined, the margins inflexed over the edges of the upper glume; upper glume lanceolate, keeled, 3-veined; florets somewhat shorter than spikelets, hyaline, with a lemma only, the palea absent; lower floret sterile, the upper floret bisexual in sessile spikelet and pistillate in pedicellate spikelet. Tropical America, Africa, India; 10 species, 2 in Venezuela, both in the flora area. Key to the Species of Eriochrysis 1. 1.

Inflorescence deep chestnut brown, 7–17 cm long; spikelets 1.5–3.2 mm long ....................................................................................... E. cayennensis Inflorescence brownish yellow, 6–9 cm long; spikelets ca. 4 mm long ............................................................................................ E. aff. holcoides

Eriochrysis cayennensis P. Beauv., Ess. Agrostogr. 8, pl. 4, fig. 11. 1812. —Cariyarena. Cespitose perennial 0.7–1.5 m tall, the nodes bearded; inflorescence very fuzzy, the spikelets subtended by dense beards of coppery hairs; lower glume of sessile spikelet veinless. Moist savannas, Mauritia palm swamps, pond margins, 100–1500 m; Bolívar (Gran Sabana, 5 km north of Ciudad Piar, 2– 5 km west of Amaruay-tepui), Amazonas (Puerto Ayacucho, Río Ocamo). Anzoátegui, Apure, Guárico, Portuguesa, Trujillo; Mexico, Central America, Greater Antilles, Colombia, Trinidad-Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Argentina, Uruguay.

Eriochrysis aff. holcoides (Nees) Kuhlm., Relat. Comiss. Linhas Telegr. Estratég. Mato Grosso Amazonas publ. 67, annexo 5, Bot. pt. 11: 89. 1922. —Anatherum holcoides Nees in Mart. et al., Fl. Bras. Enum. Pl. 2: 324. 1829. Cespitose perennial ca. 1 m tall; inflorescence not very fuzzy, the spikelets subtended by sparse beards of yellowish hairs; lower glume of sessile spikelet veinless. Mauritia palm swamps, wet savannas, 100–900 m; Bolívar (Odreman 13 km north of Santa Elena de Uairén), Amazonas (La Esmeralda, Río Parucito). Apure; eastern Colombia, Brazil, Bolivia, Paraguay, Argentina. ◆Fig. 71. Flora area material appears to be intermediate to Eriochrysis cayennensis; typical E. holcoides (as from Apure) has larger, yellow or green spikelets 5–7 mm long, with the lower glume strongly veined.

33. GOUINIA Benth. in Benth. & Hook. f., Gen. Pl. 3: 1178. 1883. [Subfamily Chloridoideae, Tribe Cynodonteae] by Emmet J. Judziewicz Perennials. Leaves with ligules membranous or uncommonly ciliate, the blades linear. Inflorescence a terminal panicle of slender racemes, these often elongate, bare at the base, and with a few secondary branches. Spikelets several-flowered, laterally compressed, disarticulating above the glumes and between the florets; glumes much shorter than the lowest floret, lanceolate, acute, keeled, the lower 1–5-veined, the upper 3–7-veined; florets with lemmas 3–5-veined, usually ciliate on the margins, the apex entire or bidentate, the midvein often prolonged into a long awn; palea shorter than lemma; uppermost floret reduced in size, but with a terminal awn; stamens 3, stigmas 2.

Gouinia 107

Mexico, Central America, Colombia, Venezuela, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina; 12 species, 1 in Venezuela. Gouinia latifolia (Griseb.) Vasey, Contr. U.S. Natl. Herb. 1: 365. 1895. —Tricuspis latifolia Griseb., Abh. Königl. Ges. Wiss. Göttingen 19: 259. 1874. Cespitose perennial 1–1.5 m from knotty base; leaf blades 15–22 × 1–1.5 cm; inflorescence ca. 50 cm long, ovate, with lax

branches to 30 cm long; spikelets 9–13 mm long, 3–5-flowered, the florets 5–6 mm long, ciliate-margined, tipped with awns 3–6 mm long. Savannas, ca. 100 m; Bolívar (21 km east of Río Caura). Guárico; Mexico, Central America, Colombia, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina. ◆Fig. 72.

Fig. 72. Gouinia latifolia

108

P OACEAE

34. GUADUA Kunth, J. Phys. Chim. Hist. Nat. Arts 95: 151. 1822. [Subfamily Bambusoideae, Tribe Bambuseae] by Emmet J. Judziewicz Erect to scandent woody bamboos of sympodial habit. Culms hollow or uncommonly solid, often with a distinctive band of short, white hairs above and below the nodal line; internodes usually sulcate above and below the insertion of a bud or branch complement; branch thorns present (except G. ciliata), especially near the base of the plant; culm sheaths triangular, the margins of the sheaths and blades contiguous or nearly so; branching pattern with buds typically solitary, with a single branch dominant. Foliage leaves with outer and inner ligules present; oral setae often conspicuous; blades linear to ovate, pseudopetiolate, typically not tessellate. Inflorescences various, either borne on separate, leafless culms, or lateral or terminal to leafy culms, indeterminate, developing by way of pseudospikelets. Pseudospikelets sessile or pedunculate, forming sparsely to densely crowded, often capitate aggregations, comprising one prophyll, several transitional glumes, several bisexual florets, and several progressively smaller and sterile apical florets, disarticulation occurring between the florets; florets lanceolate to ovate, the lemmas usually many-veined, the paleas shorter than the lemmas, 2-keeled, the keels with prominent wings; lodicules 3; stamens 6; stigmas 2 or 3, hispid to more commonly plumose. Mexico, Central America, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina, Uruguay (center of diversity Amazonian); 25–30 species, ca. 8 in Venezuela, 6 of these in the flora area. The genus needs a taxonomic revision, so the following treatment is very preliminary. Key to the Species of Guadua 1. 1. 2(1). 2. 3(2). 3. 4(1).

4.

5(4). 5.

Largest foliage leaf blades 6–17 mm wide ................................................ 2 Largest foliage leaf blades 20–65 mm wide .............................................. 4 Inflorescences borne on separate, leafless shoots; pseudospikelets 1–2 cm long ......................................................................................... G. venezuelae Inflorescences borne on leafy shoots; pseudospikelets 3–7(–11) cm long ................................................................................................................ 3 Erect plants 15–30 m tall; culms smooth ............................... G. angustifolia Arching plants ca. 3 m tall; culms minutely scabrous ............G. fascicularis Pseudospikelets (at least some of them) slightly bent downward; margins of lemmas smooth, eciliate; culms with only a tiny lumen, nearly solid ........................................................................................... G. aff. glomerata Pseudospikelets erect to divergent but never bent downward; margins of lemmas with abundant, brown, whisker-like cilia ca. 1 mm long; culms with lumen 1/3–2/3 their diameter .......................................................... 5 Plants scandent, thornless; culm leaves 9–17 cm long, the blades 1– 2.5 cm long .................................................................................... G. ciliata Plants erect, thorny; culm leaves 20–25 cm long, the blades ca. 5 cm long .................................................................................................... G. latifolia

Guadua 109

Fig. 73. Guadua ciliata

110

P OACEAE

Fig. 74. Guadua venezuelae

Gymnopogon 111

Guadua angustifolia Kunth, Syn. Pl. 1: 253. 1822. —Bambusa guadua Bonpl. in Humb. & Bonpl., Pl. Aequinoct. 1: 68, pl. 20. 1806. Giant cespitose bamboo; culms to 30 m × 10 cm, the lower branches to 2 m long; culm leaves ca. 60 × 25 cm, with irritating hispid hairs; foliage leaf blades 20–25 cm long; pseudospikelets 4–7 cm long (not seen in the flora area material). Forming groves along river banks, ca. 100 m; Amazonas (Río Mawarinuma). Mérida, Táchira; Mexico, Central America, Colombia, Trinidad-Tobago, Guyana, Ecuador, Brazil. Guadua ciliata Londoño & Davidse, Novon 1: 21. 1991. —Carrizo. Slender, climbing bamboo, the culms erect at first, later scandent and pendent, 4–10 m × 0.8–2 cm, without branch thorns; foliage leaf blades 12–27 × 2–4.5 cm, slightly subcordate at base or on pseudopetioles to 1 cm long; pseudospikelets 3–7(–11) cm long. High, infrequently flooded, forested river banks, ca. 100 m; Amazonas (Caño Caname, Caño Yagua, Río Casiquiare, Río Negro, Río Puruname). Brazil (Amazonas: Rio Negro basin). ◆Fig. 73. Guadua fascicularis Döll in Mart., Fl. Bras. 2(3): 186. 1880. Arching bamboo to 3 m tall; foliage leaf blades 8–22 cm long, linear-lanceolate, with cuneate bases; pseudospikelets reportedly 2.5–4.5 cm long, erect, the margins of the lemma eciliate. Forested river banks, 200– 700 m; Bolívar (Río Emékuni in Río Mereware basin, 5–10 km south of Salto Pará). Endemic.

Guadua aff. glomerata Munro, Trans. Linn. Soc. London 26: 79. 1868. —Carrizo. Slender, climbing bamboo; culms 2.5–4 m tall, yellow; culm leaves ca. 15 cm long, the blades 4 cm long; foliage leaf blades 10–17 × 2–3.5 cm, rounded above the pseudopetiole; pseudospikelets to 4.5 cm long. Gallery forests, near sea level to 100 m; Delta Amacuro (Caño Jota-Sabuca), Bolívar (Ciudad Bolívar). Apure; Colombia, Guyana, French Guiana, Peru, Brazil, Bolivia. Guadua latifolia (Bonpl.) Kunth, Syn. Pl. 1: 254. 1822. —Bambusa latifolia Bonpl. in Humb. & Bonpl., Pl. Aequinoct. 1: 73, pl. 21. 1806. Cespitose bamboo, mostly erect, sometimes becoming scandent later; culms to 10 m × 4 cm; foliage leaf blades 15–23 × 2–4 cm, the lower surface glabrous to puberulent; pseudospikelets 3.5–5(–8) cm long. Seasonally flooded, forested river banks and gravel bars, 50–200 m; Bolívar (Río Caroní near Caruachi), Amazonas (Río Baría, Río Casiquiare, Río Mawarinuma). Guyana, Suriname, French Guiana, Brazil. Guadua venezuelae Munro, Trans. Linn. Soc. London 26: 86. 1868. —Bambusa venezuelae (Munro) McClure, Smithsonian Contr. Bot. 9: 68. 1973. —Bambusillo. Thicket-forming bamboo to 3.5 m tall; culm leaves 12–18 × 2–3 cm, with blades 5–8 cm long; foliage leaf blades 10–20 cm × 6–17 mm. Savanna edges, gallery forests, near sea level to 1000 m; Delta Amacuro (Caño Macareo, Los Castillos), Bolívar (Cerro Zamuro, Gran Sabana). Anzoátegui, Apure, Sucre; Brazil (Amazonas). ◆Fig. 74.

35. GYMNOPOGON P. Beauv., Ess. Agrostogr. 41, 164. 1812. [Subfamily Chloridoideae, Tribe Cynodonteae] by Emmet J. Judziewicz Cespitose to rhizomatous annuals or perennials. Foliage often strongly distichous; ligules ciliate-membranous; blades linear to lanceolate, relatively short. Inflorescence terminal, consisting of several subdigitate or approximate, spreading to reflexed racemes. Spikelets ascending, often somewhat crowded, 1–several-flowered, disarticulating entire from short pedicels; glumes lanceolate, subequal, 1veined, longer than the florets; lowest (functional) floret with lemma lanceolate, 3veined, minutely bifid, short- to long-awned from between the teeth; palea slightly shorter than the lemma; rachilla internode ending in rudimentary floret(s) represented by awns; stamens 3; stigmas 2. Eastern U.S.A., Mexico, Central America, West Indies, tropical and warmtemperate South America; 13 species, 3 in Venezuela, all in the flora area.

112

P OACEAE

Key to the Species of Gymnopogon 1. 1. 2(1).

2.

Inflorescence 10–25 cm long, of 10–20 racemes borne singly on a long rachis .......................................................................................... G. spicatus Inflorescence 2.5–7 cm long, of 3–10 racemes borne subdigitately on a short rachis ............................................................................................ 2 Perennial 40–80 cm tall, the culms with 3–7 rather distant leaves with blades 2–4 cm long; spikelet with functional lemma with ciliate margins, the ultimate rachilla segment 1-awned ....................... G. fastigiatus Annual 15–40 cm tall, the culms with 9–15 crowded leaves with blades 1– 2.5 cm long; spikelet with functional lemma with glabrous margins, the ultimate rachilla segment 3-awned ..................................... G. foliosus

Gymnopogon fastigiatus Nees in Mart., Fl. Bras. Enum. Pl. 2: 430. 1829. Cespitose perennial; inflorescence 5–7 cm long. Savannas, 100–400 m; Bolívar (La Vergareña, Río Parguaza), Amazonas (Laguna Maguari, Río Ocamo, Río Parucito basin). Apure, Guárico, Miranda; Costa Rica, Panama, Colombia, Brazil, Bolivia. Gymnopogon foliosus (Willd.) Nees in Mart., Fl. Bras. Enum. Pl. 2: 426. 1829. —Chloris foliosa Willd., Sp. Pl. 4: 924. 1806. Densely tufted, fibrous-rooted annual; inflorescence 2.5–6 cm long. Dry, sandy or gravelly savannas, 50–1000 m; widespread and common in Bolívar and Amazonas. Widespread elsewhere in Venezuela; southern Mexico, Puerto Rico, Colombia, Guyana, Suriname, French Guiana, Peru, Brazil. ◆Fig. 75. Gymnopogon spicatus (Spreng.) Kuntze, Revis. Gen. Pl. 3(3): 354. 1891. —Polypogon spicatus Spreng., Syst. Veg. 1: 243. 1825 [1824]. Tufted perennial 30–50 cm tall, reportedly rhizomatous; leaves not crowded, the blades 4–6 cm long. Savannas, 50–200 m; Bolívar (Paso Caruachi, Tumeremo). Anzoátegui, Carabobo, Distrito Federal, Miranda, Monagas, Sucre, Yaracuy, Zulia; southern Mexico, Guatemala, Belize, Honduras, Colombia, Trinidad-Tobago, Guyana, Brazil, Bolivia, Paraguay, Argentina, Uruguay.

Fig. 75. Gymnopogon foliosus

Hackelochloa 113

36. GYNERIUM Willd. ex P. Beauv., Ess. Agrostogr. 138, 153, t. 24. 1812. [Subfamily Arundinoidae, Tribe Arundineae] by Gerrit Davidse Giant, rhizomatous, colonial perennial reeds; dioecious. Culms erect or arching, solid, woody, the lower parts covered with bladeless sheaths, the upper parts with a group of distichous leaves in the form of an umbrella. Leaves cauline, long and flat, clustered toward the top of culm; ligule a ciliate membrane. Panicles large, solitary, terminal; branches slender, pendent, the pistillate panicles plumose. Spikelets unisexual, dimorphic, laterally compressed, disarticulation above the glumes and between the florets. Pistillate spikelets plumose, usually 2-flowered, without rachilla extension; glumes not persistent, membranous, the lower 1-veined, the upper much longer than lemma, 3–5-veined, caudate, recurved; callus linear, glabrous or hairy; lemmas plumose, long caudate, entire; palea 1/4 as long as the lemma; lodicules 2, truncate; staminodia 2; styles 2; hilum shortly oblong. Staminate spikelet glabrous, with 2–4 florets; glumes unequal, shorter than the spikelets, lanceolate, membranous, 1-veined, the lower shorter than the upper; lemmas 1veined; paleas almost as long as the lemmas; stamens 2; rudimentary ovary sometimes present. Neotropics; 1 species. Gynerium sagittatum (Aubl.) P. Beauv., Ess. Agrostr. 138. 1812. —Saccharum sagittatum Aubl., Hist. Pl. Guiane 50. 1775. —Caña brava. Gynerium saccharoides Humb. in Humb. & Bonpl., Pl. Aequinoct. 2: 112, t. 115. 1813. Reed-like, strongly rhizomatous perennial 2–8 m tall. River banks, near sea level to 1000 m; Delta Amacuro (Misión San Francisco de Guayo), Bolívar (Gran Sabana, Río

Caura, vicinity of Santa María de Erebato), Amazonas (Río Casiquare, Río Guasacavi, Río Orinoco). Barinas, Falcón, Zulia; U.S.A., Mexico, Central America, Caribbean, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina. ◆Fig. 76. Kallio and Renvoize (Kew Bull. 49: 305– 320. 1994) have recognized three varieties of this widespread species, but only the typical variety has been reported from Venezuela.

37. HACKELOCHLOA Kuntze, Revis. Gen. Pl. 2: 776. 1891. [Subfamily Panicoideae, Tribe Andropogoneae] by Emmet J. Judziewicz Sprawling, decumbent annuals. Inflorescences numerous, produced from the middle and upper nodes of the culm; peduncles mostly included, each bearing a solitary, erect raceme; rachis internodes fused to the pedicels of the pedicellate spikelets, flattened, falling attached to the paired sessile and pedicellate spikelets, the base prolonged into an oil-bearing peg. Sessile spikelets globose; lower glume indurate, inflated, the surface impressed with numerous squarish pits in longitudinal rows, 13–15-veined, the veins evident adaxially but obscure abaxially, externally opening only by an elliptic pore on the adaxial side which fits tightly against the rachis internode and spikelet pedicel; upper glume slightly shorter than the lower, elliptic, 3-veined, convex; florets shorter than spikelet, hyaline, the lower sterile, the upper bisexual. Pedicellate spikelet usually slightly longer than the sessile, not globose, firmly membranous; lower glume broadly ovate, flat to slightly concave on back, many-veined, with inflexed margins that clasp the edges of the upper glume;

114

P OACEAE

Fig. 76. Gynerium sagittatum

Hackelochloa 115

upper glume hemispherical, strongly keeled, the keel slightly winged; florets present or absent, sterile or staminate. Southeast Asia, now a pantropical weed; 1 species. Hackelochloa granularis (L.) Kuntze, Revis. Gen. Pl. 2: 776. 1891. —Cenchrus granularis L., Mant. Pl. 2: 575. 1771. —Paja canilla teu teu. Annual, the culms sprawling and forming tangled mats; leaves with pustulate-based cilia; inflorescences 7–15 mm; sessile spikelets ca. 1 mm diameter, globose, pitted, whitish becoming black. Disturbed places, 50–100 m; Bolívar (22 km southwestern of Caicara, 25 km southwest of El Manteco), Amazonas (Río Manapiare). Widespread elsewhere in Venezuela; native to southeast Asia, now in tropical regions worldwide. ◆Fig. 77.

Fig. 77. Hackelochloa granularis

116

P OACEAE

Fig. 78. Heteropogon contortus

Homolepis 117

38. HETEROPOGON Pers., Syn. Pl. 2: 533. 1807. [Subfamily Panicoideae, Tribe Andropogoneae] by Emmet J. Judziewicz Annuals or perennials. Inflorescences numerous, produced from the upper nodes, forming a compound inflorescence; racemes solitary on each peduncle, usually with the base included within the subtending spathe-like sheath; racemes with lowermost spikelets and upper pedicellate spikelets alike, awnless, staminate, much larger than and partially twisted to conceal the awned, bisexual, sessile spikelets; rachis internodes falling attached to the spikelets, or the sessile and pedicellate spikelets with the pedicel falling first. Sessile spikelet linear to elliptic, the base prolonged into a pungent, oblique callus; glumes as long as spikelet, indurate, dark brown to black, the florets hyaline, nearly as long as the spikelet, the lower sterile, the upper bisexual, the upper lemma with a stout, geniculate, antrorsely plumose awn. Pedicellate spikelets staminate; lower glume lanceolate, broad, chlorophyllous, finely longitudinally striate, often slightly winged and twisted in the upper part; upper glume slightly shorter and enclosed by lower glume, 3-veined; florets similar to those of sessile spikelet but awnless. Tropical and subtropical regions worldwide; 6 species, 2 in Venezuela, 1 of these in the flora area. Heteropogon contortus (L.) P. Beauv. ex Roem. & Schult., Syst. Veg. 2: 836. 1817. —Andropogon contortus L., Sp. Pl. 1045. 1753. Cespitose perennial 50–75 cm tall; inflorescence 20–30 × 10 cm; racemes 3–6 cm

long; sessile spikelets 7–8 mm long, the callus pungent, with a beard of coppery hairs; awn stout, 6–10 cm long, reflexed. Savannas, 50–200 m; Bolívar (between Upata and Villa Lola). Aragua, Distrito Federal, Lara, Sucre; pantropics. ◆Fig. 78.

39. HOMOLEPIS Chase, Proc. Biol. Soc. Wash. 24: 146. 1911. [Subfamily Panicoideae, Tribe Paniceae] by Emmet J. Judziewicz Plants perennial or of indefinite duration, often stoloniferous or decumbent. Leaves with ligules membranous, often ciliate; blades linear. Inflorescence an open to contracted panicle. Spikelets broadly elliptic, obovate, or narrowly lanceolate, disarticulating entire, ± dorsally compressed, 2-flowered; glumes subequal, about as long as the spikelet, usually glabrous, 5–9-veined; lower floret sterile or staminate, the lemma glume-like, several-veined, often with long hairs in longitudinal bands, or viscid, the palea usually well developed; upper floret bisexual, usually distinctly shorter than the spikelet, elliptic, acute, firmly membranous to indurate, pale, glabrous and shining below, the lemma often with a beard of minute hairs at the apex, not strongly clasping the palea. Neotropics; 5 species, 3 in Venezuela, all in the flora area. Key to the Species of Homolepis 1. 1.

Spikelets 3–3.2 mm long, broadly elliptic to obovate, strongly viscid .................................................................................................. H. glutinosa Spikelets 5.2–8 mm long, lanceolate to elliptic, viscid or not .................. 2

118

2(1). 2.

P OACEAE

Spikelets 6.5–8 mm long, narrowly lanceolate, not viscid ......... H. aturensis Spikelets 3–6.4 mm long, narrowly to broadly elliptic, viscid ..... H. isocalycia

Homolepis aturensis (Kunth) Chase, Proc. Biol. Soc. Wash. 24: 146. 1911. —Panicum aturense Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 103, t. 33. 1815 [1816]. Straggling, stoloniferous perennial, the culms to 2 m long, 80 cm tall; leaf blades ca. 1 cm wide; panicle 6–10 cm long, ovoid, compact; spikelets green. Savannas, pastures, 100–200 m; Bolívar (widespread), Amazonas (Auyán-tepui). Mérida, Táchira, Trujillo, Zulia; Mexico, Central America, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 79. Homolepis glutinosa (Sw.) Zuloaga & Soderstr., Smithsonian Contr. Bot. 59: 19. 1985. —Panicum glutinosum Sw., Prodr. 24. 1788. —Uríyu. Cespitose perennial to 2 m tall, the culms decumbent and rooting; leaf blades 1.8–2.5 cm wide; panicle 15–30 cm long, ovoid, open;

spikelets dark. Forest edges, cultivated areas, 900–1400 m; Bolívar (Gran Sabana). Aragua, Distrito Federal, Falcón, Lara, Miranda, Monagas, Portuguesa, Táchira, Yaracuy; Mexico, Central America, Greater Antilles, Colombia, Guyana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina. ◆Fig. 80. Homolepis isocalycia (G. Mey.) Chase, Proc. Biol. Soc. Wash. 24: 147. 1911. —Panicum isocalycium G. Mey., Prim. Fl. Esseq. 59. 1818. Rhizomatous perennial 0.5–1.5 m tall, the culms decumbent and rooting; leaf blades 1.3–2 cm wide; panicle 8–12 cm long, ovoid. Savannas, river sides, forest edges, 100–1100 m; Bolívar (Gran Sabana, upper Río Paragua, Upata), Amazonas (Puerto Ayacucho). Táchira, Trujillo; Mexico, Panama, Colombia, Guyana, Suriname, French Guiana, northern Brazil. ◆Fig. 81.

40. HYMENACHNE P. Beauv., Ess. Agrostogr. 48, 165. 1812. [Subfamily Panicoideae, Tribe Paniceae] by Emmet J. Judziewicz Robust aquatic or semiaquatic perennials, the culms sprawling, creeping and rooting at the nodes, spongy, aerenchymatous. Leaves with ligules membranous; blades typically lanceolate, cordate at the base. Inflorescence a loosely to densely spicate panicle. Spikelets ± lanceolate, somewhat dorsally compressed, glabrous, 2flowered; lower glume short, 1–3-veined, an evident internode present between the glumes; upper glume as long as spikelet, 3–5-veined; lower floret sterile, the lemma as long as the spikelet, 3–5-veined, acute to short-awned, the palea absent; upper floret distinctly shorter than the spikelet, papery, pale, glabrous, the margins of the lemma not inrolled over the edges of the palea. Pantropics; 5 species, 2 in Venezuela, both in the flora area. Key to the Species of Hymenachne 1.

1.

Spikelets 4–5.3 mm long, the lower lemma attenuate to subaristate; inflorescence densely spicate, 0.7–1.2 cm wide at base, the lower branches strictly erect, 1.5–5(–8) cm long ...................... H. amplexicaulis Spikelets 2.5–3.3 mm long, the lower lemma acuminate; inflorescence loosely contracted, 3–6 cm wide at base, the lower branches divergent, 4–8 cm long ........................................................................... H. donacifolia

Homolepis 119

Fig. 79. Homolepis aturensis Fig. 80. Homolepis glutinosa

Fig. 81. Homolepis isocalycia

120

P OACEAE

Fig. 82. Hymenachne donacifolia

Hyparrhenia 121

Hymenachne amplexicaulis (Rudge) Nees in Mart., Fl. Bras. Enum. Pl. 2: 276. 1829. —Panicum amplexicaule Rudge, Pl. Guian. 21, pl. 27. 1805. Cespitose perennial to 2 m tall, 1 cm diameter; inflorescence 15–40 cm long. Stream banks, marshes, often in shallow water or on sand bars, near sea level to 500 m; Delta Amacuro (common), Bolívar (base of Auyántepui, Río Orinoco, Río Parguaza, Upata). Widespread in lowland Venezuela and the Neotropics.

Hymenachne donacifolia (Raddi) Chase, J. Wash. Acad. Sci. 13: 177. 1923. —Panicum donacifolium Raddi, Agrostogr. Bras. 44. 1823. Cespitose perennial to 2 m tall, 1 cm diameter; inflorescence 25–40 cm long. Stream banks, often in shallow water, near sea level to 300 m; Delta Amacuro (common), Bolívar (between Caicara and Ciudad Bolívar, Río Toro). Apure, Cojedes, Miranda; Honduras, Costa Rica, Panama, Cuba, Trinidad-Tobago, tropical South America. ◆Fig. 82.

41. HYPARRHENIA Andersson ex Fourn., Mex. Pl. 2: 51, 67. 1886. [Subfamily Panicoideae, Tribe Andropogoneae] by Emmet J. Judziewicz Coarse perennials. Inflorescence compound, of few to numerous pairs of racemes in spathe-like bracts from the upper and terminal nodes; peduncle included in spathe-like bract or exserted; racemes unequal to subequal, appressed to divergent, of few to many spikelet pairs and terminating in a triad of 1 sessile and 2 pedicellate spikelets; lowest pair of spikelets on a raceme awnless, sterile or staminate; rachis internodes thin, filiform, disarticulating along with the basally attached spikelets. Sessile spikelet with glumes equal, firmly membranous; lower glume flat to slightly sulcate on back, obscurely many-veined, the apex entire to bifid, often scabrid-ciliolate on margins near apex, the margins embracing those of the upper glume; upper glume 1–3-veined, rounded on back; florets somewhat shorter than the spikelet, hyaline; lower floret sterile; upper floret bisexual, the lemma bifid, awned from the sinus, the awn elongate, geniculate, kinked shortly after its emergence from the spikelet. Pedicel of pedicellate spikelet similar to rachis internode; spikelet sterile or staminate, the upper lemma awnless. Pantropics, mostly Africa; ca. 55 species, 3 species in Venezuela, 2 of these in the flora area. Key to the Species of Hyparrhenia 1.

1.

Racemes 1–1.3 cm long, the rachis internodes and pedicels pubescent with white or tawny cilia, the spikelets glabrous; sessile spikelet of terminal triad 5.5–6.5 mm long; spathe-like bracts densely pubescent .................................................................................................. H. bracteata Racemes 1.8–3.5 cm long, the rachis internodes, pedicels, and lower glumes of spikelets pubescent with golden brown cilia; sessile spikelet of terminal triad 3.7–4.5 mm long; spathe-like bracts glabrous or less commonly puberulent ...................................................................... H. rufa

Hyparrhenia bracteata (Willd.) Stapf in Prain, Fl. Trop. Afr. 9: 360. 1919. —Andropogon bracteatus Humb. & Bonpl. ex Willd., Sp. Pl. 4: 914. 1806. —Cymbopogon bracteatus (Willd.)

Hitchc., Contr. U.S. Natl. Herb. 17: 209. 1913. —Canaotera. Coarse cespitose perennial 1–2 m tall; inflorescence 15–30 × 5–8 cm; racemes paired, sharply reflexed away from each other. Sea-

122

P OACEAE

Fig. 83. Hyparrhenia bracteata

Ichnanthus 123

sonally flooded savannas, sometimes dominant, 900–1100 m; Bolívar (Gran Sabana). Widespread elsewhere in Venezuela; southern Mexico, Central America, Colombia, Guyana, Suriname, Brazil, Bolivia, Paraguay, tropical Africa. ◆Fig. 83. Hyparrhenia rufa (Nees) Stapf in Prain, Fl. Trop. Afr. 9: 304. 1919. —Trachypogon rufus Nees in Mart. et al., Fl. Bras. Enum.

Pl. 2: 345. 1829. —Paja del Brasil. Cespitose perennial 1–2.5 m tall; inflorescence ca. 30 cm long; racemes paired, only slightly divergent from each other. Cultivated for forage, widely naturalized in pastures, aggressively weedy if frequently burned, 50–300 m; Bolívar (El Callao), Amazonas (El Venado, Puerto Ayacucho, Samariapo). Widespread elsewhere in Venezuela; pantropics, native of Africa.

42. ICHNANTHUS P. Beauv., Ess. Agrostogr. 56. 1812. [Subfamily Panicoideae, Tribe Paniceae] by Emmet J. Judziewicz Annuals or perennials; small straggling herbs, broad-leaved forest understory grasses, or rambling bamboo-like climbers. Leaves with ligules membranous; blades often lanceolate to ovate, asymmetrical at the base. Inflorescences paniculate, terminal and sometimes axillary, rather open, the loosely paired spikelets deciduous from the summit of the pedicels, or the upper floret sometimes falling first. Spikelets elliptic, ovate, or lanceolate, acute to attenuate, laterally compressed to terete, 2-flowered; lower glume about 1/2 as long as to nearly as long as spikelet, several-veined, usually keeled, separated from the upper glume by a distinct internode; upper glume and lower floret subequal, about as long as the spikelet, (3–)5–7veined; lower floret sterile or commonly staminate, the palea well developed; upper floret somewhat shorter than spikelet, elliptic to lanceolate or ovate, indurate, pale, glabrous or uncommonly pubescent, often rotated 90° within the spikelet at maturity; basal margins of upper lemma with minute, paired scars, or with conspicuous, paired, lanceolate appendages, these adnate to basal margins of lemma but arising from the rachilla internode between the florets. Neotropics; 29 species, 13 in Venezuela, 11 of these in the flora area. Key to the Species of Ichnanthus 1. 1. 2(1). 2. 3(2). 3. 4(3). 4. 5(4).

5.

Spikelets 8–10 mm long; upper floret silky-hairy ..................... I. panicoides Spikelets 3–6.5 mm long; upper floret glabrous ....................................... 2 Leaf blades 15–75 cm long, linear to lanceolate ....................................... 3 Leaf blades < 15 cm long, lanceolate to elliptic ........................................ 6 Leaf blades linear, 40–75 × 0.5–1.8 cm ......................... I. ephemeroblepharis Leaf blades lanceolate, 15–38 × 1.5–5 cm ................................................. 4 Spikelets 5–5.7 mm long; upper floret with rachilla internode appendages 2–3 mm long ................................................................... I. lancifolius Spikelets 3–4.7 mm long; upper floret with rachilla internode appendages < 1 mm long ................................................................................... 5 Apex of leaf sheath bearing auricles 3–5 mm long; leaf blades (2–)2.5– 5 cm wide, the lower surface bluish whitened; lower glume 1/2–2/3 as long as spikelet; upper floret lacking rachilla internode appendages ................................................................................................ I. breviscrobs Apex of leaf sheath lacking auricles; leaf blades 1–2.8 cm wide, the lower

124

P OACEAE

surface green; lower glume 3/4 as long as to nearly as long as spikelet; upper floret with rachilla internode appendages 0.6–0.8 mm long .................................................................................................. I. calvescens 6(2). Upper floret with rachilla internode appendages 1.5–2.5 mm long .................................................................................................. I. nemoralis 6. Upper floret with rachilla internode appendages absent, represented by scars only ............................................................................................... 7 7(6). Inflorescence branches naked at base, bearing 2 or 3 pairs of spikelets near the middle, and terminating in a long-pedicellate spikelet; annual ..................................................................................... I. procurrens 7. Inflorescence branches bearing spikelets all along their lengths; annual or perennial ............................................................................................ 8 8(7). Inflorescence with both primary and secondary branches divaricately spreading .................................................................................. I. ruprechtii 8. Inflorescence not with both primary and secondary branches divaricately spreading ................................................................................................ 9 9(8). Spikelets ovate, plump, the upper glume and lower lemma acuminate to abruptly subaristate ............................................................... I. nemorosus 9. Spikelets lanceolate to ovate, the upper glume and lower lemma acute to subaristate (if subaristate, then the spikelets lanceolate) ................ 10 10(9). Inflorescence congested, many-flowered, the relatively stout peduncle included or exserted to 10 cm; spikelets broadly lanceolate, usually glabrous, never with spreading cilia; perennial ............................... I. pallens 10. Inflorescence diffuse, few-flowered, long-exserted on a slender peduncle 5–15 cm long; spikelets narrowly lanceolate, usually with spreading cilia, uncommonly glabrous; annual .............................................. I. tenuis Ichnanthus breviscrobs Döll in Mart., Fl. Bras. 2(2): 294. 1877. —Caicará, Ceduichu (Yekwana), Suduchu (Yekwana), Waná. Much-branched perennial, the culms to 5 m long, 1.3 cm diameter, weak, sprawling, scandent, or pendent; leaf blades 22– 33 cm long, tapering at both ends; inflorescence 17–40 cm long; spikelets 3.5–4.7 mm long. Forming dense thickets on forest edges or in gaps, often where burned, savannas, Mauritia palm swamps, cultivated areas, 100–1000 m; Delta Amacuro (Serranía de Imataca), Bolívar (Gran Sabana, mouth of Río Nichare, tributaries of Río Orinoco), Amazonas (Río Cunucunuma, Río Orinoco, Sierra Parima). Colombia, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia. ◆Fig. 84. The culms of Ichnanthus breviscrobs are used by Amerindians as flutes. Ichnanthus calvescens (Nees) Döll in Mart., Fl. Bras. 2(2): 285. 1877. —Pani-

cum calvescens Nees in Trin., Gram. Panic. 193. 1826. Ichnanthus chaseae Swallen, Contr. U.S. Natl. Herb. 29: 271. 1949. Ichnanthus acuminatus Swallen, Fieldiana, Bot. 28: 28. 1951. Ichnanthus hispidus Swallen, Phytologia 11: 78. 1964. Ichnanthus vestitus Swallen, Phytologia 11: 75. 1964. Loosely cespitose perennial, the culms to 2 m long, 7 mm diameter, somewhat scandent; leaves cauline or nearly all basal, nearly glabrous to densely pubescent, the blades 10–38 cm long; inflorescence 10–45 cm long, the branches whorled; spikelets 3–4 mm long. Open savannas, shrub islands, forest edges, stream banks, dry thickets, open slopes of tepuis, cultivated areas, 100–1700 m; Delta Amacuro (Serranía de Imataca), Bolívar (common), Amazonas (common). Widespread in Venezuelan Coastal Cordillera; Colombia, Trinidad-Tobago, Guyana, Suriname, Peru, Brazil. ◆Fig. 85.

Ichnanthus 125

Ichnanthus ephemeroblepharis G.A. Black & Fróes ex G.A. Black & Pires, Bol. Técn. Inst. Agron. N. 15: 5. 1948. Ichnanthus longifolius Swallen, Fieldiana, Bot. 28: 29. 1951. Ichnanthus serratus Swallen, Fieldiana, Bot. 28: 30. 1951. Ichnanthus angustus Swallen, Mem. New York Bot. Gard. 9: 266. 1957. Ichnanthus tectus Swallen, Mem. New York Bot. Gard. 9: 265. 1957. Ichnanthus neblinaensis Swallen, Phytologia 14: 84. 1966. Cespitose perennial to 1 m tall, or culms decumbent and to 5 m long; leaves basal; blades coarse, with white, cartilaginous, scabrous margins; inflorescence 10–30 cm long; spikelets 3–5 mm long, the upper floret with rachilla internode appendages 1–1.5 mm long. Open forests, lajas, stream banks in savannas, sandy savanna edges, 100–2100 m; Bolívar (Cerro Sarisariñama, Río Canaracuni), Amazonas (Cerro Aracamuni, Cerro Duida, Cerro Guanacoco, Cerro Marahuaka, Cerro Parú, Río Mawarinuma, Sierra de la Neblina). Adjacent Brazil (Amazonas). ◆Fig. 87. Ichnanthus lancifolius Mez, Repert. Spec. Nov. Regni Veg. 15: 126. 1918. Peru, northeastern Brazil, Bolivia. 2 varieties, 1 in Venezuela. I. lancifolius var. weberbaueri (Mez) Stieber, Syst. Bot. 7: 106. 1982. —Ichnanthus weberbaueri Mez, Repert. Spec. Nov. Regni Veg. 15: 127. 1918. Ichnanthus duidensis Swallen, Fieldiana, Bot. 28: 28. 1951. Cespitose, rhizomatous perennial to 1 m tall; leaf blades 23–35 × 2.5–10 cm, tapering at both ends; inflorescence 20–30 cm long, the branches ascending at an angle of 45°. Gaps in forests, 200–1400 m; Amazonas (Cerro Duida, Cerro Sipapo, Cerro Yutajé, Sierra de la Neblina, Sierra Parima). Peru, Bolivia. ◆Fig. 89. Ichnanthus nemoralis (Schrad. ex Schult.) Hitchc. & Chase, Contr. U.S. Natl. Herb. 18: 334. 1917. —Panicum nemorale Schrad. ex Schult., Mantissa 2: 255. 1824. —Pitillo. Loosely tufted perennial 0.5–1 m tall, the culms decumbent, branching and rooting at nodes; leaf blades 8–13 × 1.5–3.5 cm; inflo-

rescence 15–25 cm long, with few, stiff branches; spikelets 4.3–6.5 mm long. Forest shade, 100–1200 m; Delta Amacuro (near Los Castillos), Bolívar (common on Gran Sabana, Represa Guri, Río Toro, Río Uarama, Río Venamo, Serranía de Imataca, San Pedro de las Bocas near mouth of Río Paragua, Upata). Aragua, Carabobo, Falcón, Miranda, Monagas, Nueva Esparta, Sucre, Yaracuy; Belize, Honduras, Panama, Colombia, Trinidad-Tobago, Guyana, Suriname, French Guiana, Brazil. Ichnanthus nemorosus (Sw.) Döll in Mart., Fl. Bras. 2(2): 289. 1877. —Panicum nemorosum Sw., Prodr. 22. 1788. Ichnanthus nubilis Chase, J. Wash. Acad. Sci. 42: 124, fig. 3. 1952. Ichnanthus tenuifolius K.E. Rogers, Phytologia 24: 417. 1972. Sprawling, creeping perennial, erect portions of culms to 50 cm tall; leaves as in Ichnanthus pallens. Upland to highland forests, 500–1300 m; Amazonas (lower slopes of Sierra de la Neblina). Widespread in Venezuelan Coastal Cordillera; southern Mexico, Central America, West Indies, Colombia, Ecuador, Peru, Argentina. Ichnanthus pallens (Sw.) Munro ex Benth., Fl. Hongk. 414. 1861. —Panicum pallens Sw., Prodr. 23. 1788. Panicum axillare Nees in Mart., Fl. Bras. Enum. Pl. 2: 141. 1829. —Ichnanthus axillaris (Nees) Hitchc. & Chase, Contr. U.S. Natl. Herb. 18: 334. 1917. Sprawling, creeping, mat-forming perennial, culms to 1 m long and 0.7 m tall; leaf blades 2.5–12 × 1–3.4 cm, lanceolate to nearly circular; inflorescence 2–20 cm long, often ovoid and compact; spikelets 2.5–4.5 mm long, lanceolate to ovate, the upper glume and lower lemma acute to acuminate. Edges of and gaps in forests, stream banks, weedy places, near sea level to 1300 m; widespread in Delta Amacuro, Bolívar, and Amazonas. Widespread elsewhere in Venezuela; Mexico, Central America, West Indies, tropical and warm-temperate South America, Africa, Asia, Malesia, Australia. Ichnanthus panicoides P. Beauv., Ess. Agrostogr. 56, 57, pl. 12, fig. 1. 1812. Cespitose, usually unbranched perennial 0.5–1 m tall; leaf blades 10–25 × (3–)4–7 cm,

126

P OACEAE

Fig. 84. Ichnanthus breviscrobs

Ichnanthus 127

Fig. 85. Ichnanthus calvescens

Fig. 86. Ichnanthus panicoides

128

P OACEAE

Fig. 87. Ichnanthus ephemeroblepharis

Fig. 88. Ichnanthus tenuis

Ichnanthus 129

Fig. 89. Ichnanthus lancifolius var. weberbaueri

130

P OACEAE

Fig. 90. Ichnanthus procurrens

Imperata 131

elliptic to ovate-lanceolate, cross-veined; upper floret with rachilla internode appendages 4–5.5 mm long. Forest shade and in gaps, 100–1100 m; Delta Amacuro (Río Cuyubini), Bolívar (Altiplanicie de Nuria, Amaruaytepui, El Paují, Macizo del Chimantá [Toronó-tepui], Río Ichún, Río Paramichi), Amazonas (base of Cerro Duida, base of Cerro Huachamacari, Río Orinoco, Río Siapa). Colombia, Guyana, Suriname, French Guiana, Peru, western Brazil, Bolivia. ◆Fig. 86. Ichnanthus procurrens (Nees ex Trin.) Swallen, Phytologia 11: 149. 1964. —Panicum procurrens Nees ex Trin., Gram. Panic. 183. 1826. Sprawling, decumbent annual; leaf blades 4–7 × 1–1.5 cm; inflorescence 2.5–7 cm long, the few branches ascending at a 45° angle; spikelets with spreading cilia. Moist, rocky, savanna-covered slopes, 100–200 m; northwestern Bolívar. Guárico, Portuguesa; Brazil, Bolivia, Paraguay, Argentina. ◆Fig. 90. Ichnanthus ruprechtii Döll in Mart., Fl. Bras. 2(2): 293. 1877. Sprawling, creeping, mat-forming peren-

nial. Forest edges, ca. 200 m; Bolívar (east of Urimán). Sucre; Guyana, Peru, Brazil, Bolivia, Paraguay. Ichnanthus tenuis (J. Presl) Hitchc. & Chase, Contr. U.S. Natl. Herb. 18: 334. 1917. —Oplismenus tenuis J. Presl in C. Presl, Reliq. Haenk. 1: 319. 1830. —Jiba, Soowo. Ichnanthus venezuelanus Mez, Repert. Spec. Nov. Regni Veg. 15: 132. 1918. Ichnanthus tamayonis Chase, J. Wash. Acad. Sci. 42: 122, fig. 2. 1952. Ichnanthus sabulosus K.E. Rogers, Phytologia 26: 61. 1973. Ichnanthus sucrensis K.E. Rogers, Phytologia 26: 63. 1973. Weak, sprawling annual, the erect portions of culms to 30 cm tall; leaf blades (1.5–) 3–10 × 0.5–2 cm, lanceolate to ovate; inflorescences 2.5–10 cm long; spikelets 3.5–5.3 mm long. Mauritia palm swamps, forest edges, often along streams near waterfalls, in sinkholes, 100–1300 m; Bolívar (widespread), Amazonas (widespread). Venezuelan Coastal Cordillera; southern Mexico, Central America, Dominican Republic, tropical and warmtemperate South America. ◆Fig. 88.

43. IMPERATA Cirillo, Pl. Rar. Neapol. 2: 26. 1792. [Subfamily Panicoideae, Tribe Andropogoneae] by Emmet J. Judziewicz Coarse, stoloniferous perennials. Inflorescence a cylindrical to narrowly pyramidal panicle of erect or ascending racemes, usually densely white-silky; racemes bearing pairs of spikelets on unequal pedicels, the pedicels filiform, flaring and cupulate at apex, the spikelets falling individually from the persistent rachis. Spikelets all alike, narrowly lanceolate (slightly gaping at maturity), delicately membranous, the truncate callus bearing a long beard of silky hairs much longer than the spikelet; glumes as long as spikelet, subequal, 1–5-veined, silky-hairy on the back, the lower glume often distinctly shorter than the upper; florets hyaline, much shorter than the spikelet, glabrous; lower floret sterile, lacking a palea; upper floret bisexual, the lemma sometimes absent, the palea usually well developed, broad; stamen often solitary. Tropical and subtropical regions worldwide; 9 species, 2 in Venezuela, both in the flora area.

132

P OACEAE

Fig. 91. Imperata brasiliensis

Isachne 133

Key to the Species of Imperata 1. 1.

Inflorescence 7–16 cm long, the lower branches 2–3 cm long, erect; foliage predominantly basal .............................................................. I. brasiliensis Inflorescence ca. 55 cm long, the lower branches 6–12 cm long, slightly divergent; foliage predominantly cauline ................................ I. contracta

Imperata brasiliensis Trin., Mém. Acad. Imp. Sci. St.-Pétersbourg, Sér. 6, Sci. Math., Seconde Pt. Sci. Nat. 2: 331. 1832. Cespitose, stoloniferous perennial 0.5–1 m tall; spikelets 3.3–4.5 mm long. Savannas, sometimes dominant, 100–1400 m; Bolívar (Río Caroni, Río Marirupá, Uaipán-tepui), Amazonas (San Pedro del Orinoco, Sierra Parima). Widespread elsewhere in Venezuela; U.S.A. (Florida), Mexico, Central America, Cuba, Colombia, Guyana, Suriname, French Guiana, Brazil, Bolivia, Paraguay, Argentina, Uruguay. ◆Fig. 91.

Imperata contracta (Kunth) Hitchc., Annual Rep. Missouri Bot. Gard. 1893: 146. 1893. —Saccharum contractum Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 182. 1815 [1816]. Imperata flexuosa Swallen, Phytologia 14: 87. 1966. Stoloniferous perennial 0.5–1.2 m tall; spikelets 3–4 mm long. Savannas, ca. 900 m; Bolívar (Oparuma in Gran Sabana). Anzoátegui, Apure, Barinas, Cojedes, Distrito Federal, Falcón, Guárico, Lara, Mérida, Monagas, Táchira; Mexico, Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia.

44. ISACHNE R. Br., Prodr. 196. 1810. [Subfamily Panicoideae, Tribe Isachneae] by Emmet J. Judziewicz Plants annual or perennial. Leaves with ligules ciliate; blades linear to lanceolate. Inflorescence usually an open panicle. Spikelets falling entire or the glumes persistent, the florets falling as a unit, biconvex, 2-flowered; lower glume nearly as long as spikelet, few- to many-veined; internode between glumes evident; upper glume about as long as spikelet, many-veined; lower floret staminate or bisexual, membranous or papery to indurate, distinctly shorter than to as long as spikelet, the palea well developed; rachilla internode evident between florets; upper floret pistillate or bisexual, about as long as spikelet, plano-convex, indurate, pale. Pantropics; ca. 100 species, 2 in Venezuela, both in the flora area. Key to the Species of Isachne 1. 1.

Spikelets with upper floret pubescent; leaf blades 2–3.5 cm long, lanceolate-ovate ........................................................................ I. polygonoides Spikelets with upper floret glabrous; leaf blades 6–12 cm long, linear-lanceolate ............................................................................................. I. rigens

Isachne polygonoides (Lam.) Döll in Mart., Fl. Bras. 2(2): 273. 1877. —Panicum polygonoides Lam., Encycl. 4: 742. 1798. Sprawling, decumbent, clone-forming plant, the erect culms to 30 cm tall; inflorescence 3–9 cm long, compact, the branches

with minute yellow bands; spikelets 1.3–1.8 mm long. Swamps, marshes, stream sides, ponds, seasonally flooded savannas, 100–500 m; Bolívar (Altiplanicie de Nuria, southeastern base of Auyán-tepui, south of Caicara, Caño Azul, 10 km south of Ciudad Bolívar, La Vergareña), Amazonas (Puerto Ayacucho).

134

P OACEAE

Fig. 92. Isachne rigens

Fig. 93. Isachne polygonoides

Ischaemum 135

Anzoátegui, Apure, Guárico, Monagas, Portuguesa, Táchira; Neotropics. ◆Fig. 93. Isachne rigens (Sw.) Trin., Gram. Panic. 252. 1826. —Panicum rigens Sw., Prodr. 23. 1788. —Uiri-yu-yek. Isachne ligulata Swallen, Caldasia 2: 305. 1943. Isachne arundinacea auct. non (Sw.) Griseb. 1864: sensu Luces, Gramíneas Distr. Fed. 171. 1963.

Perennial to 1.5 m tall, the culms weak and semi-scandent; inflorescence 6–13 cm long, open; spikelets 1.6–2.3 mm long. Upper slopes and summits of tepuis, often on bare rock, 1400–2500 m; Bolívar (Auyán-tepui, Cerro Venamo, Macizo del Chimantá, Roraima-tepui, Sororopán-tepui), Amazonas (Sierra Tapirapecó). Anzoátegui, Distrito Federal, Lara, Mérida, Monagas, Sucre, Táchira, Trujillo; Jamaica, Lesser Antilles, Colombia, Guyana, Ecuador, Peru. ◆Fig. 92.

45. ISCHAEMUM L., Sp. Pl. 1049. 1753. [Subfamily Panicoideae, Tribe Andropogoneae] Meoschium P. Beauv., Ess. Agrostogr. 111, 167. 1812. Ischaemopogon Griseb., Fl. Brit. W. I. 560. 1864. by Gerrit Davidse Annuals or perennials, cespitose. Culms hollow or solid. Sheaths keeled; ligule a membrane; leaf blades linear to linear-lanceolate, flat. Inflorescence 2–many, digitate or racemose racemes; racemes articulated, the spikelets paired, the 2 spikelets of a pair and an internode segment of the raceme falling as 1 unit or the pedicellate spikelets falling separately. Spikelets dorsally compressed, unequally pedicellate or 1 sessile, equal, with 2 florets. Sessile spikelets bisexual; callus obtuse, inserted in the concave apex of the raceme internode segment; glumes longer than the florets; lower glume rigid in the lower part, herbaceous in the upper part, flat; upper glume slightly larger than the lower glume, membranous, keeled, 5–9-veined; lower floret staminate; upper floret pistillate or bisexual, the upper lemma hyaline, 2-lobed to the middle, awned from between the lobes, the awn geniculate or twisted; upper palea hyaline, nearly as long as the upper lemma; lodicules 2; stamens 3; styles 2; hilum punctate. Pedicellate staminate spikelets, similar to the sessile spikelets, but frequently somewhat smaller; pedicels free; both florets staminate. Tropics, mostly in the Old World; ca. 65 species, 5 in Venezuela, 4 of these in the flora area. Key to the Species of Ischaemum 1. 1. 2(1). 2. 3(2).

3.

Lower glume of sessile spikelets deeply transversely rugose ...... I. rugosum Lower glume of sessile spikelets smooth, not rugose ............................... 2 Leaf blades 4–7 mm wide; racemes 2 ........................................... I. arenosum Leaf blades 8–20 mm wide; racemes 2–25 ................................................ 3 Racemes 2–5(–9); subsessile spikelets 4.5–6.2 mm long; rachis internodes 0.5–0.9 mm wide; callus hairs usually shorter than the basal 1/4 of the adjoining subsessile spikelet, rarely reaching the middle of the subsessile spikelets .................................................................. I. guianense Racemes 10–25; subsessile spikelets 3.4–3.6 mm long; rachis internodes 0.2–0.3 mm wide; callus hairs always longer than the basal 1/4 of the adjoining subsessile spikelet, usually reaching the middle of the subsessile spikelet and the longer hairs reaching the apex ........ I. latifolium

136

P OACEAE

Fig. 94. Ischaemum guianense

Fig. 95. Ischaemum arenosum

Lasiacis 137

Ischaemum arenosum Sohns, Mem. New York Bot. Gard. 9(3): 404, fig. 75. 1957. Sprawling perennial. In moist sand along rivers, 400–500 m; Bolívar (Macizo del Chimantá). Endemic. ◆Fig. 95. Ischaemum arenosum is only known with certainty from the type collection. It was described as having solitary racemes, but this was a mistake as the holotype (US) and an isotype (VEN) clearly have two racemes. As in other species with paired racemes, they may be held close together until anthesis. Ischaemum arenosum is similar to I. timorense Kunth, an Asian species introduced as a forage plant into the American tropics but not yet known from the flora area. Ischaemum arenosum differs in its long-pilose glumes, and more robust and tougher culms. Ischaemum guianense Kunth ex Hack. in A. DC., Monogr. Phan. 6: 235. 1889. Ischaemum guianense var. schomburgkii Hack. in A. DC., Monogr. Phan. 6: 235. 1889. Perennial to 170 cm tall. Savannas, river banks, granite outcrops, 50–1900 m; common in Bolívar and Amazonas. Anzoátegui; Guyana, Suriname, French Guiana, Brazil (Amapá). ◆Fig. 94. Ischaemum guianense is a very variable species that needs additional study. As here interpreted, it includes decumbent plants with soft spreading foliage from waterfall basins to tall erect plants with stiff erect foliage from savannas and river banks. Spikelets are usually scabrous, at least in the upper half, but other plants may have glumes that are densely long-pilose. It may be that populations stretching from Cojedes, Barinas, Trujillo, and Mérida to Norte de Santander in Colombia are another variant of this species. They are remarkably pilose on the glumes and rachis segments, but spikelet size, raceme number, and the morphology of

the pedicels and rachis segments are similar to those of Ischaemum guianense. Ischaemum latifolium (Spreng.) Kunth, Révis. Gramin. 168. 1829. —Andropogon latifolius Spreng., Syst. Veg. 1: 286. 1825. —Ischaemopogon latifolius (Spreng.) Griseb., Fl. Brit. W. I. 560. 1864. Ischaemum latifolium var. minus E. Fourn., Mexic. Pl. 2: 55. 1886. Ischaemum latifolium subsp. hirtivaginum Hitchc., Contr. U.S. Natl. Herb. 24: 506. 1927. Perennial to 150 cm tall. Rocky outcrops, 100–1500 m; rare in Amazonas (Río Siapa, Sierra Tapirapecó). Anzoátegui, Barinas, Cojedes, Lara, Mérida, Táchira, Trujillo, Yaracuy; Mexico, Central America, Caribbean, Colombia, Ecuador, Peru, Brazil. Ischaemum rugosum Salisb., Icon. Stirp. Rar. 1, t. 1. 1791. —Meoschium rugosum (Salisb.) Nees, Gramineae 68. 1841. Ischaemum segetum Trin., Mém. Acad. Imp. Sci. St. Pétersbourg, Sér. 6, Sci. Math. 2(3): 294. 1832. —Andropogon segetum (Trin.) Steud., Syn. Pl. Glumac. 1: 376. 1855 [1854]. —Ischaemum rugosum var. segetum (Trin.) Hack. in A. DC. & C. DC., Monogr. Phan. 6(1): 208. 1889. Weedy annual 25–70 cm tall. Secondary, weedy vegetation, especially in open, seasonally wet sites, 100–200 m; Amazonas (vicinity of Puerto Ayacucho). Barinas, Carabobo, Guárico, Portuguesa; introduced from Mexico, Nicaragua, Costa Rica, Panama, Caribbean, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, native to tropical Asia, now a pantropical weed. Ischaemum rugosum is apparently a recently (1982) introduced weed in the flora area and can be expected to spread into all lowland areas that are regularly disturbed by human activities.

46. LASIACIS (Griseb.) Hitchc., Contr. U.S. Natl. Herb. 15: 16. 1910. —Panicum sect. Lasiacis Griseb., Fl. Brit. W. I. 551. 1864. —Carricillo, Taquari. [Subfamily Panicoideae, Tribe Paniceae] by Gerrit Davidse Plants perennial, rarely annual in L. procerrima; culms freely branched, cespitose, highly lignified and erect, arching or climbing, or procumbent and nearly

138

P OACEAE

herbaceous, creeping, and rooting at the nodes; internodes solid or hollow; culm pulvini well developed, sheaths rounded, the margins free and overlapping; collar sometimes somewhat enlarged to form a small pseudopetiole; blades linear to ovate, slightly to prominently asymmetrical at the base; inflorescence an open or contracted panicle; spikelets subglobose to globose, obovate or elliptic, placed obliquely on the pedicel; disarticulation below the glumes; glumes and sterile lemmas broad, abruptly apiculate, membranous, shiny black at maturity, woolly at the apex; first glume 1/3–2/3 as long as the spikelet, 5–13-veined, saccate at the base, the margins overlapping; second glume and sterile lemma subequal, about as long as the fertile floret, 7–15-veined, the sterile lemma enclosing a palea 1/4 as long as to equal to the length of the fertile floret; sterile floret with or without a staminate flower; a second sterile lemma present in L. anomala; cells of the inner epidermis of the glumes and sterile lemmas filled with oil globules at maturity; fertile lemma indurate, obtuse, the margins inrolled enclosing the edges of the indurate palea, usually dark brown at maturity, broadly elliptic to obovate; palea gibbous above, concave below, its margins lobed, overlapping or meeting on the dorsal side of the caryopsis; both fertile lemma and palea with woolly pubescence in slight excavations at their apices; rachilla sometimes prolonged beyond the base of the fertile floret; stamens 3; styles 2, the bases separate; lodicules 2, fleshy, truncate, vasculated, 1 limb within the fertile palea, the other situated between the fertile lemma and palea; caryopsis planoconvex, ovate, obovate, or nearly orbicular, the apex rounded, sometimes broadly grooved on the hilum side; hilum oblong or nearly round; embryo approximately 1/2 the length of the caryopsis. U.S.A. (southern Florida), Mexico, Central America, the Caribbean, South America south to Argentina; 16 species, 12 in Venezuela, 8 of these in the flora area. Key to the Species of Lasiacis 1. 1. 2(1).

2.

3(2). 3. 4(3). 4. 5(3). 5. 6(5). 6.

Sterile florets 2 .............................................................................. L. anomala Sterile floret 1 ............................................................................................ 2 Leaf blades cordate at the base, 25–31 cm long, 35–43 mm wide; inflorescences 40–100 cm long; base of plants with prominent prop roots ................................................................................................ L. procerrima Leaf blades rounded to cuneate at the base, 4–19(–23) cm long, 3–45(–56) mm wide; inflorescences 3–35 cm long; base of plants without prominent prop roots ....................................................................................... 3 Ligules 1.6–7 mm long ............................................................................... 4 Ligules 0.3–1.5(–2.6) mm long .................................................................. 5 Inflorescences 1–9 cm long; ligules (3.5–)4–6(–7) mm long .......... L. scabrior Inflorescences 2–17(–21) cm long; ligules (1.6–)2–3(–3.7) mm long .......... ..................................................................................................... L. ligulata Leaf blades with obvious pubescence on at least one of the leaf surfaces ................................................................................................................ 6 Leaf blades glabrous on both surfaces, sometimes with a few, small, scattered hairs at the base of the upper and/or lower surfaces ................. 7 Spikelets (3.6–)4–5 mm long; panicles rather open with few spikelets, the pedicels widely spreading ............................................................... L. nigra Spikelets (3–)3.4–4.1(–4.3) mm long; panicles much denser, the pedicels not as widely spreading ......................................................... L. sorghoidea

Lasiacis 139

7(5). 7.

Main inflorescence branches sparsely branched, bearing few spikelets (3.6–)4–5(–5.3) mm long .............................................................. L. sloanei Main inflorescence branches highly branched, bearing numerous spikelets (2.6–)2.8–4.1 mm long ...................................................... L. ruscifolia

Lasiacis anomala Hitchc., J. Wash. Acad. Sci. 9: 37. 1919. Bamboo-like perennial 0.75–5 m tall. Margins of gallery forests, brush of secondary vegetation, tree islands in savannas, near sea level to 800 m; widespread in Delta Amacuro, Bolívar, and Amazonas. Anzoátegui, Apure, Aragua, Carabobo, Falcón, Guárico, Lara, Miranda, Monagas, Nueva Esparta, Portuguesa, Sucre, Yaracuy, Zulia; Caribbean, Colombia, Guyana, French Guiana, Suriname, Brazil, Bolivia. ◆Fig. 97. Lasiacis ligulata Hitchc. & Chase, Contr. U.S. Natl. Herb. 18: 337. 1917. Bamboo-like perennial 0.75–5 m tall. Forest margins, light gaps in forests, secondary vegetation, near sea level to 1700 m; Delta Amacuro (Antonio Diaz), Bolívar (Gran Sabana), Amazonas (widespread). Anzoátegui, Miranda, Nueva Esparta Sucre; Caribbean, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 96. Lasiacis nigra Davidse, Phytologia 29: 152. 1974. Bamboo-like perennial 1–8 m tall; ligule prominent. Forest margins, light gaps in forests, secondary vegetation, 900–1300 m; Bolívar (Gran Sabana). Aragua, Distrito Federal, Falcón, Mérida, Miranda, Sucre, Táchira, Trujillo, Yaracuy, Zulia; Mexico, Central America, Colombia, Ecuador, Peru. Lasiacis procerrima (Hack.) Hitchc., Proc. Biol. Soc. Wash. 24: 145. 1911. —Panicum procerrimum Hack, Oesterr. Bot. Z. 51: 431. 1901. Annual to 3 m tall; panicles large, widely spreading; prop roots conspicuous. Forest margins, roadsides, secondary vegetation, 50– 1400 m; Bolívar (widespread), Amazonas (Atabapo, headwaters of Río Orinoco). Anzoátegui, Aragua, Barinas, Carabobo, Cojedes, Distrito Federal, Falcón, Guárico, Lara, Mérida, Miranda, Portuguesa, Táchira, Trujillo, Yaracuy, Zulia; Mexico, Central America, Colombia, Guyana, Ecuador, Peru, Brazil. ◆Fig. 98.

Lasiacis ruscifolia (Kunth) Hitchc., Proc. Biol. Soc. Wash. 24: 145. 1911. —Panicum ruscifolium Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 101. 1815 [1816]. Lasiacis compacta (Sw. ex Wikstr.) Hitchc., Bot. Gaz. 51(4): 301. 1911. —Panicum compactum Sw. ex Wikstr. in Sw., Adnot. Bot. 14. 1829. Panicum liebmannianum E. Fourn., Mexic. Pl. 2: 33. 1886. —Lasiacis liebmanniana (E. Fourn.) Hitchc., Proc. Biol. Soc. Wash. 24: 145. 1911. Lasiacis glabra Swallen, Ceiba 4(5): 287. 1955. Bamboo-like perennial, culms 1–8 m tall. Forest margins, roadsides, secondary vegetation, 100–200 m; Amazonas (Nericagua). Anzoátegui, Aragua, Carabobo, Cojedes, Falcón, Guárico, Lara, Miranda, Yaracuy, Zulia; U.S.A., Mexico, Belize, Honduras, El Salvador, Nicaragua, Costa Rica, Panama, Caribbean, Colombia, Guyana, Ecuador. Lasiacis scabrior Hitchc., Proc. Biol. Soc. Wash. 40: 85. 1927. Bamboo-like perennial, culms 1–8 m tall; ligule prominent. Forest margins, roadsides, secondary vegetation, 50–400 m; Amazonas (Atabapo, upper Río Orinoco). Mexico, Central America, Colombia, Ecuador, Peru. ◆Fig. 100. Lasiacis sloanei (Griseb.) Hitchc., Bot. Gaz. 51: 302. 1911. —Panicum sloanei Griseb., Fl. Brit. W. I. 551. 1864. Panicum latifolium Ham., Prodr. Pl. Ind. Occid. 10. 1825. Bamboo-like perennial, culms 1–6 m tall. Forest margins, roadsides, secondary vegetation, 50–900 m; Bolívar (Altiplanicie de Nuria, Pueblo Viejo). Aragua, Distrito Federal, Lara, Mérida; Mexico, Guatemala, Belize, Honduras, Nicaragua, Costa Rica, Caribbean, Colombia, Ecuador. ◆Fig. 99. Lasiacis sorghoidea (Desv. ex Ham.) Hitchc. & Chase, Contr. U.S. Natl. Herb. 18: 338. 1917. —Panicum sorghoideum

140

P OACEAE

Fig. 96. Lasiacis ligulata

Fig. 97. Lasiacis anomala

Lasiacis 141

Fig. 98. Lasiacis procerrima

142

P OACEAE

Fig. 99. Lasiacis sloanei

Fig. 100. Lasiacis scabrior

Leersia 143

Desv. ex Ham., Prodr. Pl. Ind. Occid. 10. 1825. —Panicum lanatum var. sorghoideum (Desv. ex Ham.) Griseb., Fl. Brit. W. I. 551. 1864. Panicum lanatum Sw., Prodr. 24. 1788, non Rottb. 1776. Panicum glutinosum Lam., Tabl. Encycl. 1: 74, t. 43, fig. 3. 1791, non Sw. 1788. —Panicum agglutinans Kunth, Enum. Pl. 1: 120. 1833. Panicum divaricatum var. latifolium Schltdl. & Cham., Linnaea 6: 33. 1831. Panicum divaricatum var. lanatum Schltdl. & Cham., Linnaea 6(1): 33. 1831. Panicum praegnans Steud., Pl. Glumac. 1: 74. 1854. Panicum guaraniticum Speg., Anales Soc. Ci. Argent. 16: 107. 1883. —Lasiacis guaraniticum (Speg.) Parodi, Notas Mus. La Plata, Bot. 8: 95. 1943. Panicum swartzianum Hitchc., Contr.

U.S. Natl. Herb. 12: 140. 1908. —Lasiacis swartziana (Hitchc.) Hitchc., Bot. Gaz. (Crawfordsville) 51: 302. 1911. Lasiacis patentiflora Hitchc. & Chase, Contr. U.S. Natl. Herb. 18: 338. 1917. —Lasiacis sorghoidea var. patentiflora (Hitchc. & Chase) Davidse, Ann. Missouri Bot. Gard. 64: 375. 1977 [1978]. Lasiacis acuminata Swallen, Mem. New York Bot. Gard. 9: 267. 1957. Bamboo-like perennial, culms 1–10 m tall. Forest margins, roadsides, secondary vegetation, 50–900 m; common in Bolívar and Amazonas. Anzoátegui, Apure, Aragua, Barinas, Carabobo, Distrito Federal, Falcón, Guárico, Lara, Mérida, Miranda, Monagas, Portuguesa, Sucre, Táchira, Trujillo, Yaracuy, Zulia; Mexico, Central America, Caribbean, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina.

47. LEERSIA Sw., Prodr. 21. 1788, nom. cons. [Subfamily Bambusoideae, Tribe Oryzeae] by Emmet J. Judziewicz Cespitose or rhizomatous perennials, often aquatic. Leaves with ligules membranous; blades linear. Inflorescences terminal panicles, the rice-like spikelets often overlapping on the branches. Spikelets elliptic to ovate, usually strongly laterally flattened, asymmetrically apiculate, bisexual, 1-flowered, falling entire; glumes reduced to a minute ridge or cupule at the summit of the pedicel; lemma as long as spikelet, often gibbous, 5-veined, the veins often pectinate-scabrid, the margins concealing most of palea; palea keeled, 3-veined; stamens 1–6; stigmas 2. Temperate to tropical areas worldwide; 18 species, 2 in Venezuela, both in the flora area. Key to the Species of Leersia 1. 1.

Spikelets 3–3.8 mm long, the lemma with veins pectinate-scabrid .................................................................................................. L. hexandra Spikelets 2–2.5 mm long, the lemma with veins smooth ............. L. ligularis

Leersia hexandra Sw., Prodr. 21. 1788. Rhizomatous perennial 0.3–0.6 cm tall; ligules 2–4 mm long; leaf blades 10–25 × 0.3– 1 cm. Swamps, marshes, stream banks, near sea level to 300 m; common in Delta Amacuro and northeastern Bolívar. Anzoátegui, Apure, Barinas, Cojedes, Distrito Federal, Guárico, Miranda, Portuguesa, Zulia; widespread worldwide in tropical and warm-temperate regions, often weedy in rice fields.

Leersia ligularis Trin., Mém. Acad. Imp. Sci. St.-Pétersbourg, Sér. 6, Sci. Math., Seconde Pt. Sci. Nat. 5: 168. 1840. Oryza monandra var. grandiflora Döll in Mart., Fl. Bras. 2(2): 9. 1871. —Leersia grandiflora (Döll) Prodoehl, Bot. Arch. 1: 219. 1922. —Leersia ligularis var. grandiflora (Döll) Pyrah, Iowa State J. Sci. 44: 236. 1969. Leersia distichophylla Balansa & Poitr.,

144

P OACEAE

Bull. Soc. Hist. Nat. Toulouse 12: 221, t. 1, fig. 2. 1878. Leersia ligularis var. glabriflora Pyrah, Iowa State J. Sci. 44: 238. 1969. Cespitose perennial 1–2 m tall; ligules 5– 10 mm long; leaf blades 25–40 × 1–1.5 cm. Upland forest gaps and edges, 100–500 m;

Bolívar (Altiplanicie de Nuria, Cerro Picacho). Distrito Federal, Falcón, Lara, Miranda; southern Mexico, Guatemala, Belize, Honduras, El Salvador, Costa Rica, Panama, Lesser Antilles, Colombia, Ecuador, Brazil, Paraguay, Argentina, Uruguay. ◆Fig. 101.

Fig. 101. Leersia ligularis

Leptochloa 145

48. LEPTOCHLOA P. Beauv., Ess. Agrostogr. 71. 1812. [Subfamily Chloridoideae, Tribe Cynodonteae] by Emmet J. Judziewicz Cespitose annuals or perennials. Leaves with ligules ciliate-membranous; blades linear. Inflorescence a terminal panicle of slender racemes; racemes triquetrous, with spikelets alternating on one side, loosely overlapping or slightly distant from each other. Spikelets several-flowered, laterally compressed, disarticulating above the glumes and also between the florets; glumes much shorter than to as long as spikelet, narrow, keeled, 1-veined; lemmas 3-veined, the lateral veins submarginal, the midvein awned or not; palea shorter than lemma; upper florets often reduced in size, the uppermost rudimentary. Worldwide in tropical and warm-temperate regions; ca. 40 species, 5 in Venezuela, 3 of these in the flora area. Key to the Species of Leptochloa 1.

1.

2(1).

2.

Spikelets 2- or 3-flowered, the florets ca. 1 mm long, elliptic; glumes narrowly lanceolate, subequal, widely divergent, nearly as long as the spikelet; leaf blades 3–5 mm wide ........................................... L. filiformis Spikelets (3)4–7-flowered, the florets 2–3 mm long, lanceolate; glumes lanceolate, unequal, not widely divergent, at most 2/3 as long as spikelet; leaf blades 5–15 mm wide ............................................................... 2 Ligules 1.5–2.5 mm long, the membranous part hyaline; glumes 1/4–2/5 as long as the spikelet, leaf blades 5–10 mm wide, scabrous; lemmas awnless ................................................................................................. L. scabra Ligules 0.4–0.8 mm long, the membranous part indurate; glumes 1/2–2/3 as long as the spikelet; leaf blades 8–15 mm wide, smooth; lemmas awned or awnless ......................................................................... L. virgata

Leptochloa filiformis (Lam.) P. Beauv., Ess. Agrostogr. 71, 163, 166. 1812. —Festuca filiformis Lam., Tabl. Encycl. 1: 191. 1791. Annual with weak culms ca. 40 cm tall. Weedy places, ca. 100 m; Bolívar (San Martín de Turumbán southwest of Tumeremo). Widespread elsewhere in Venezuela; eastern U.S.A., Mexico, Central America, West Indies, tropical and warm-temperate South America. Leptochloa scabra Nees in Mart., Fl. Bras. Enum. Pl. 2: 270. 1829. Tufted annual 40–80 cm tall. Disturbed areas, 50–100 m; Delta Amacuro (Güiniquina, Tucupita), Amazonas (Capuana on Río Orinoco, mouth of Río Parhueña). Widespread elsewhere in Venezuela and throughout tropical and warm-temperate America. ◆Fig. 102.

Leptochloa virgata (L.) P. Beauv., Ess. Agrostogr. 71, 166, pl. 5, fig. 1. 1812. —Cynosurus virgatus L., Syst. Nat. ed. 10, 876. 1759. Cynosurus domingensis Jacq., Misc. Austriac. 2: 363. 1781. —Leptochloa domingensis (Jacq.) Trin., Fund. Agrost. 133. 1820. Cespitose perennial 50–100 cm tall; spikelets with lemmas awnless or with awns to 2.5 mm long. Savannas, disturbed areas, 100– 1200 m; Delta Amacuro (Santa Catalina), Bolívar (Isla de Santa Bárbara, San Martín de Turumbán southwest of Tumeremo, Santa Elena de Uairén), Amazonas (headwaters Río Orinoco). Widespread elsewhere in Venezuela and throughout tropical and warmtemperate America. ◆Fig. 103.

146

P OACEAE

Fig. 102. Leptochloa scabra

Fig. 103. Leptochloa virgata

Lithachne 147

49. LEPTOCORYPHIUM Nees in Mart. et al., Fl. Bras. Enum. Pl. 2: 83. 1829. [Subfamily Panicoideae, Tribe Paniceae] Anthaenantia P. Beauv., Ess. Agrostogr. 48, t. 10. 1812. by Emmet J. Judziewicz Cespitose, rarely stoloniferous perennials. Leaves with ligules membranousciliate; blades linear. Inflorescence an open, narrowly pyramidal panicle, the spikelet pedicels elongate. Spikelets lanceolate, dorsally compressed, falling entire, 2flowered; lower glume absent or represented by a cupule; upper glume and lower lemma equal, as long as the spikelet, strongly 5–7-veined, the veins bearing elongate silky hairs; lower floret sterile, the palea absent; upper floret lanceolate, papery, dark brown, 3-veined, glabrous, the margins of the lemma not inrolled over the edges of the palea. Neotropics; 2 species, 1 in Venezuela. Leptocoryphium lanatum (Kunth) Nees in Mart., Fl. Bras. Enum. Pl. 2: 84. 1829. —Paspalum lanatum Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 94, t. 29. 1815 [1816]. —Anthaenantia lanata (Kunth) Benth., J. Linn. Soc. Bot. 19: 39. 1881. Densely cespitose perennial to 80 cm tall, the culms swollen at base; leaves coarse, the

sheaths persistent, shredding, the blades usually involute; inflorescence 9–15 cm long; spikelets 4–5 mm long, with abundant spreading white or tawny hairs. Common and often dominant in dry savannas, 100– 500(–1100) m; Bolívar (widespread), Amazonas (widespread). Throughout the rest of Venezuela and the Neotropics. ◆Fig. 104.

50. LITHACHNE P. Beauv., Ess. Agrostogr. 135, 166, 168. 1812. [Subfamily Bambusoideae, Tribe Olyreae] by Emmet J. Judziewicz Monoecious, cespitose perennials. Leaves exhibiting sleep movements, drooping downwards at night; ligules membranous; pseudopetioles short; blades lanceolate to ovate, asymmetrically truncate at base. Inflorescences racemiform, lateral and terminal, only partially exserted from the sheaths; lateral inflorescences bearing 1–several pistillate spikelets at the apex, accompanied by several subterminal staminate spikelets; terminal inflorescence staminate only. Spikelets 1-flowered, the pistillate spikelets much larger than the staminates. Pistillate spikelet pedicels clavate; glumes persistent or not, subequal, ovate, caudate, many-veined; floret shorter than glumes, obpyramidal, indurate, laterally compressed, hooded, white becoming marbled with dark spots at maturity, borne on a peg-like persistent rachilla internode, the margins of the lemma strongly covering and nearly concealing the slightly protruding palea; stigmas 2, plumose. Staminate spikelets on filiform pedicels, caducous, hyaline, lacking glumes; glumes absent; floret linear, translucent, the lemma 3-veined; stamens 3. Mexico, Central America, West Indies, Colombia, Venezuela, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina (absent from Amazonian area); 4 species, 1 in Venezuela. Lithachne pauciflora (Sw.) P. Beauv., Ess. Agrost. 135, 168. 1812. —Olyra pauciflora Sw., Prodr. 21. 1788. Cespitose perennial 25–60 cm; leaf blades ca. 7 × 2.5 cm; pistillate florets 4–5 mm long.

Forest understory, ca. 500 m; Bolívar (Río Venamo between Isla Anacoco and Puesto Venamo). Falcón, Lara, Zulia; Mexico, Central America, West Indies, tropical South America (except Suriname and Bolivia). ◆Fig. 105.

148

P OACEAE

Fig. 104. Leptocoryphium lanatum

Fig. 105. Lithachne pauciflora

Luziola 149

51. LUZIOLA Juss., Gen. Pl. 33. 1789. [Subfamily Bambusoideae, Tribe Oryzeae] by Emmet J. Judziewicz Monoecious aquatic perennials. Spikelets unisexual, 1-flowered, usually borne in separate panicles, the glumes absent or represented by a minute cupule, the spikelets usually falling entire from the cupule. Pistillate panicles open to umbelliform, axillary, often several from each node; spikelets linear to elliptic when young, ovate when mature, the lemma and palea equal, similar, several- to manyveined; stigmas 2, white, plumose. Staminate panicles solitary, terminal, open to racemose; spikelets narrow, hyaline to membranous, the lemma and palea equal, several-veined; stamens 6(–18), the anthers often showy, yellow to reddish. Caryopsis ± globose, persistent within the floret or sometimes falling before the spikelet, beaked, longitudinally striate. Southeastern U.S.A., Mexico, Central America, West Indies, tropical and warm-temperate South America; 11–15 species, 3 in Venezuela, all in the flora area. Key to the Species of Luziola 1. 1. 2(1). 2.

Pistillate spikelets 4–5 mm long; pistillate inflorescence umbelliform, exserted from a 6–12 mm wide leaf sheath .......................... L. subintegra Pistillate spikelets 1–3.7 mm long; pistillate inflorescence paniculate, exserted from a 2–4 mm wide leaf sheath ................................................ 2 Pistillate spikelets 3–3.7 mm long in a panicle 3–5 cm long ..... L. bahiensis Pistillate spikelets 1–1.5 mm long in a panicle 5–10 cm long ....L. doelliana

Luziola bahiensis (Steud.) Hitchc., Contr. U.S. Natl. Herb. 12: 234. 1909. —Caryochloa bahiensis Steud., Syn. Pl. Glumac. 1: 5. 1855 [1853]. Perennial 30–50 cm tall; leaves erect; pistillate panicles often contracted, the spikelets strongly 9–13-veined. Stream banks, 400–1200 m; Bolívar (Altiplanicie de Nuria, Gran Sabana, Hato Divina Pastora). Apure, Guárico, Táchira; southeastern U.S.A., Mexico, Central America, Cuba, Dominican Republic, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina. Luziola doelliana Prodoehl, Bot. Arch. 1: 240. 1922. Luziola pittieri Luces, J. Wash. Acad. Sci. 32: 159, fig. 3. 1942. Perennial 25–60 cm tall; leaves spreading; pistillate panicles open, the spikelets weakly 11–13-veined. Seasonally flooded savannas, ponds, near sea level to 200 m; Bolívar (east of Caicara, Moitaco, Oríllas del Orinoco near

Cabruta). Apure, Barinas, Guárico, Portuguesa; northeastern Brazil. ◆Fig. 106. If interpreted in a very broad sense, the older name Luziola brasiliana Moric. may apply to this taxon. Luziola subintegra Swallen, Ann. Missouri Bot. Gard. 30: 165. 1943. Luziola spruceana auct. non Benth. ex Döll 1871: sensu Hitchc., Contr. U.S. Natl. Herb. 22: 463. 1922; Amshoff & Henrard in Pulle, Fl. Suriname 1(1): 310. 1943. Floating, mat-forming perennial; pistillate inflorescences 4–6 cm, the branches becoming strongly bent downward, the spikelets strongly 5–7-veined. Streams, ponds, near sea level to 100 m; Delta Amacuro (east of Isla de Guaro), Amazonas (Ocamo). Apure, Falcón, Guárico, Portuguesa; Mexico, Central America, Greater Antilles, Colombia, Trinidad-Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina. ◆Fig. 107.

150

P OACEAE

Fig. 106. Luziola doelliana

Fig. 107. Luziola subintegra

Melinis 151

52. MELINIS P. Beauv., Ess. Agrostogr. 54. 1812. [Subfamily Panicoideae, Tribe Paniceae] by Emmet J. Judziewicz Plants sprawling, decumbent. Leaves pubescent, viscid, aromatic, the ligules ciliate; blades linear. Inflorescence a bushy panicle. Spikelets oblong, laterally compressed, purplish, falling entire, 2-flowered; lower glume a minute, veinless scale; upper glume and lower lemma as long as spikelet, membranous, emarginate, strongly veined; upper glume oblong-elliptic, awned from the back or not; lower floret sterile, the lemma lanceolate to elliptic, 3–5-veined, with a delicate awn produced from the back just below the emarginate apex, the palea absent; upper floret shorter than spikelet, lanceolate, hyaline. Worldwide in tropical regions, natives of tropical and South Africa; 11 species, 1 naturalized worldwide in tropical regions. Melinis minutiflora P. Beauv., Ess. Agrostogr. 54, pl. 11, fig. 4. 1812. —Capim melas, Paja gordura. Sprawling, decumbent perennial to 70 cm tall; foliage velvety, aromatic (smelling of linseed oil); inflorescence 15–20 cm long, green becoming purple; spikelets 1.5–2 mm long, the lower lemma with an awn 5–10 mm long. Cultivated for forage and frequently naturalized, 100–1300 m; Bolívar (Gran Sabana), Amazonas (Río Casiquiare, Río Cunucunuma, Río Orinoco). Native of tropical Africa, naturalized throughout tropical regions. ◆Fig. 108.

Fig. 108. Melinis minutiflora

152

P OACEAE

53. MEROSTACHYS Spreng., Syst. Veg. 1: 132. 1825 [1824]. [Subfamily Bambusoideae, Tribe Bambuseae] by Emmet J. Judziewicz Cespitose woody bamboos. Rhizomes determinate. Culms unarmed, hollow or rarely pithy, erect at base, becoming scandent or pendent in the upper portions. Culm leaves with blades usually narrow and bent downward. Primary buds solitary at the midculm nodes, when developing, soon producing numerous small subsidiary branches from a truncate to shallowly fan-shaped meristem, with no supranodal ridge evident. Foliage leaves in complements; inner ligules, outer ligules, and oral setae all present; blades deciduous, usually broad. Inflorescences racemose, determinate, terminating leafy branches, often densely secund, the sessile or short-pedicellate spikelets borne in 1 or 2 rows from the lower side of the rachis. Spikelets with glumes absent or rudimentary; sterile florets 2, glume-like, unequal, much shorter than the spikelet; functional florets 1(2), the lemmas many-veined, the paleas strongly 2-keeled, sulcate, the keels clasping the rachilla internode; stamens 3, stigmas 2; rachilla prolonged beyond functional floret(s) to form a minute bristle, or sometimes terminating in a rudimentary spikelet. Central America, Colombia, Venezuela, Guyana, Peru, Brazil, Bolivia, Paraguay, Argentina; ca. 45 poorly known species, 3 or 4 in Venezuela, 2 of these in the flora area. Key to the Species of Merostachys 1. 1.

Foliage leaf blades 5–6 × 1–1.5 cm; rachis densely silky-hairy ................. .......................................................................................... M. maguireorum Foliage leaf blades 10–21 × 2–4.5 cm; rachis sparsely short-hairy ........... .................................................................................................... M. retrorsa

Merostachys maguireorum McClure, Mem. New York Bot. Gard. 10(5): 5. 1964. Cespitose bamboo with arching, glabrous culms 12–15 m × 1.5–3 cm, the lowest internode 1–1.5 m long; inflorescence 5–7 cm long, of 8–13 secund, divergent spikelets ca. 16 × 3 mm. Cloud forests, 1500–1800 m; Bolívar (Cerro Marahuaka). Endemic. ◆Fig. 109. The protologue gives the type locality as the upper Río Cunucunuma. However, the locality on the type specimen gives Cerro Marahuaka and we are following the latter.

Merostachys retrorsa McClure, Mem. New York Bot. Gard. 10(5): 6. 1964. Erect to sprawling bamboo with culms to 4 m long; inflorescence 6–10 cm long, of 10–15 secund, bent downward spikelets ca. 16 × 2.5 mm. Forested river sides in lowland and montane forests, 50–1000 m; Bolívar (mouth of Río Nichare, Río Uonán, Río Tirica), Amazonas (saddle between Cerro Duida and Cerro Marahuaka). Southwestern Guyana. ◆Fig. 110.

54. MESOSETUM Steud., Syn. Pl. Glumac. 1: 118. [1854]. [Subfamily Panicoideae, Tribe Paniceae] by Emmet J. Judziewicz Cespitose or stoloniferous perennials or annuals, the foliage mostly basal. Leaves with ligules membranous-ciliate; blades linear. Inflorescence terminal, a solitary spike or spike-like raceme; rachis triquetrous, bearing crowded, appressed, sessile to short-pedicellate spikelets in pairs on 2 sides. Spikelets falling entire, usu-

Merostachys 153

Fig. 109. Merostachys maguireorum

154

P OACEAE

Fig. 110. Merostachys retrorsa

Mesosetum 155

ally somewhat diamond- or fan-shaped, laterally compressed, glabrous or often pubescent in complex patterns, 2-flowered; lower glume distinctly shorter than to nearly as long as spikelet, facing the rachis, flat to turgid, keeled or not, 1–3-veined; upper glume somewhat shorter than to about as long as spikelet, 3–5-veined; lower floret usually sterile, about as long as spikelet, the lemma 3–5-veined, the palea usually well developed; upper floret distinctly shorter than spikelet, lanceolate, acuminate, slightly beaked, coriaceous, pale, glabrous, the lemma only loosely enfolding the palea. Southern Mexico, Central America, Cuba, Jamaica, Colombia, Venezuela, Trinidad-Tobago, Guyana, Suriname, French Guiana, Brazil, Bolivia, Paraguay, northeastern Argentina; 25 species, 7 in Venezuela, 6 of these in the flora area. Key to the Species of Mesosetum 1. 1. 2(1). 2. 3(1). 3. 4(3). 4. 5(4). 5.

Spikelets pubescent with tawny or brown hairs ...................................... 2 Spikelets pubescent with clear or white hairs.......................................... 3 Apex of lower glume erose, irregular, or bidentate; rare ........ M. cayennense Apex of lower glume obtuse to acute; common ................... M. rottboellioides Leaf blades 15–35 cm long, involute (rarely flat); very densely cespitose perennial ................................................................................. M. filifolium Leaf blades < 15 cm long, involute or flat; annuals or tufted perennials, often stoloniferous ................................................................................. 4 Lower glume mucronate or aristate with an awn to 3 mm long; rachis of racemes 0.8–1.8 mm wide ......................................................... M. chaseae Lower glume obtuse to acute; rachis of racemes 0.2–0.5 mm wide ......... 5 Spikelets 2.5–3.3 mm long, in 1 row on the rachis; annual ....................... ....................................................................................... M. chlorostachyum Spikelets 3.5–4 mm long, in 2 rows on the rachis; perennial .... M. loliiforme

Mesosetum cayennense Steud., Syn. Pl. Glumac. 1: 118. 1855 [1854]. Cespitose perennial ca. 50 cm tall; leaf blades 5–8 cm × 2–4 mm; inflorescence 3–6 cm long, the rachis 0.3–0.4 mm wide; spikelets 3.2–3.6 mm long, the bracts with tawny or brown hairs in tufts. Savannas, 100–500 m; Bolívar (La Vergareña, middle Río Caura). Colombia, Guyana, Suriname, French Guiana, Brazil, Bolivia. ◆Fig. 113. Mesosetum chaseae Luces, J. Wash. Acad. Sci. 32: 160. 1942. —Wana’ ipun. Mesosetum cardonum Luces, Bol. Soc. Venez. Ci. Nat. 15: 19, fig. 12. 1953. Tufted perennial 25–65 cm tall, developing elongate rhizomes and stolons to 1 m long; leaf blades 5–13 cm × 2–5 mm; inflorescence 5–8 cm long; spikelets (3.5–)4.5–5.8 mm long. Savannas, sometimes on granite, 100–1100 m; Bolívar (near Caicara, Canaima, Kurún-tepui near Cerro Venado, Río Am-

buituir south of Auyán-tepui, Río Maniapure, Río Paragua, Río Topopo, Uaipán-tepui). Anzoátegui, Guárico; Guatemala, Honduras, Trinidad-Tobago, Guyana, Brazil, Paraguay. ◆Fig. 111. Mesosetum chlorostachyum (Döll) Chase, Proc. Biol. Soc. Wash. 24: 122. 1911. —Panicum chlorostachyum Döll in Mart., Fl. Bras. 2(2): 175, pl. 28a. 1877. Slender, tufted annual 25–40 cm tall; leaves all basal, the blades 7–12 cm × 2–4 mm; inflorescence 4–8 cm long. Savannas, ca. 100 m; Amazonas (Caño Caname). Brazil (Amazonas, Mato Grosso, Pará). ◆Fig. 114. Mesosetum filifolium Hubb., Proc. Amer. Acad. Arts 49: 493. 1913. Coarse, densely cespitose perennial 40–60 cm tall; leaf blades involute, curving, 1 mm wide, or rarely flat and to 3 mm wide; inflorescence 4–9 cm long; spikelets 3.3–5(–6) mm

156

P OACEAE

long. White-sand savannas, 100–200 m; Bolívar (Raudal Maturín on Río Asa), Amazonas (Caño Camani, Caño Cotúa near base of Cerro Yapacana, Caño Yagua, Río Ventuari). Apure; Belize, Nicaragua. ◆Fig. 112. Mesosetum loliiforme (Hochst. ex Steud.) Chase, Bot. Gaz. (Crawfordsville) 51: 302. 1911. —Panicum loliiforme Hochst. ex Steud., Syn. Pl. Glumac. 1: 56. 1855 [1853]. Tufted perennial 40–60 cm tall, the culms often somewhat decumbent; leaves mostly basal, the blades 4–7 cm × 2–5 mm, flat or less commonly involute; inflorescence 4–7 cm long. Savannas, 100–300 m; Bolívar (near El Manteco), Amazonas (30 km north of Puerto Ayacucho). Barinas; Cuba, Colombia, Trinidad-Tobago, Guyana, Suriname, French Guiana, Brazil, Bolivia. Mesosetum rottboellioides (Kunth) Hitchc., Contr. U.S. Natl. Herb. 12: 211. 1909. —Panicum rottboellioides Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 96, t. 32. 1815 [1816]. Cespitose perennial 50–75 cm tall; leaf blades 4–20 cm × 1–7 mm, flat; inflorescence 6–22 cm long, the rachis 0.2–0.3 mm wide; spikelets 3.2–5 mm long, the bracts with tawny or brown hairs in tufts. Savannas, lajas, 50–500(–1300) m; Bolívar (Gran Sabana, Río Orinoco basin), Amazonas (throughout). Apure; eastern Colombia, Guyana, Suriname, French Guiana, western Brazil, Bolivia. ◆Fig. 115.

Fig. 111. Mesosetum chaseae

Mesosetum 157

Fig. 112. Mesosetum filifolium

Fig. 113. Mesosetum cayennense

158

P OACEAE

Fig. 114. Mesosetum chlorostachyum

Fig. 115. Mesosetum rottboellioides

Myriocladus 159

55. MYRIOCLADUS Swallen, Fieldiana, Bot. 28: 34. 1951. [Subfamily Bambusoideae, Tribe Bambuseae] by Emmet J. Judziewicz Woody bamboos from short, determinate rhizomes. Culms erect, hollow to solid; lowest internode usually very long, the succeeding nodes placed very close together and bearing a leaf complement; branching often occurring in upper part of plant at or near the leaf complement, the branches appressed, borne singly at the nodes but often appearing clustered because of the very short internodes. Culm and foliage leaves not differentiated; sheaths persistent, imbricate, greatly thickened at the rounded to keeled summit; inner ligule short, membranous; outer ligule short, rim-like; oral setae often present; pseudopetiole present in most species, short; blades small or more commonly long and broad, coriaceous, deciduous. Inflorescence a raceme or more commonly an ovoid to pyramidal terminal panicle. Spikelets with glumes 2, awned or awnless; lowest floret sterile, lacking a palea and flower; fertile florets 1–several(–10); uppermost floret rudimentary; lodicules 3; stamens 3; stigmas 2; style 1. Tepuis of southern Venezuela and adjacent Brazil; 13 species, all in the flora area. Myriocladus may be most closely related to Aulonemia, even though the type species M. virgatus bears a striking vegetative resemblance to Neurolepis. The key does not account for the dwarf-flowering forms that may occur in populations of normal-sized individuals; these have been observed in M. distantiflorus, M. paludicolus, and M. virgatus. Key to the Species of Myriocladus 1. 1. 2(1). 2. 3(1). 3. 4(3). 4. 5(4). 5. 6(5). 6.

Leaf blades 0.7–3.5 mm wide; inflorescence 2–3.5 cm long, racemose; internodes of approximately equal lengths, the lowest not elongated ... 2 Leaf blades 6–150 mm wide; inflorescence 4–110 cm long, paniculate; internodes of very different lengths, the lowest greatly elongated ..... 3 Leaf blades 45–65 × 0.7–1.5 mm, involute; oral setae adundant, fine, white ........................................................................................ M. involutus Leaf blades 10–30 × 1.5–3.5 mm, flat; oral setae few, coarse, brown ............................................................................................ M. steyermarkii Leaf blades narrow at the base, tapering gradually into the sheath, not obtuse, cordate, or clasping; spikelets 2.8–3.8 mm long ......... M. virgatus Leaf blades obtuse, cordate, or clasping at the base; spikelets 4–13 mm long ......................................................................................................... 4 Inflorescence a compact, ovoid, often long-exserted panicle 4–10 × 1.5– 2.5 cm ................................................................................. M. neblinaensis Inflorescence an open, ovoid to narrowly pyramidal panicle 10–70 × 2– 15 cm ...................................................................................................... 5 Leaf blades 15–20 times as long as wide ........................................... M. sp. A Leaf blades 4–12 times as long as wide .................................................... 6 Inflorescence 15–30 times as long as wide, the branches 1.5–2 cm long; upper glume abruptly awned .................................................. M. cardonae Inflorescence 1.5–15 times as long as wide, the branches 2–20 cm long;

160

P OACEAE

7(6). 7. 8(7). 8. 9(8). 9. 10(7). 10. 11(10). 11. 12(10). 12. 13(12). 13.

upper glume in most species acute or tapering gradually into an awn ................................................................................................................ 7 Leaf blades 6–15 cm wide .......................................................................... 8 Leaf blades 1–4.5(–5.5) cm wide ............................................................. 10 Spikelets 4–5.3 mm long ....................................................... M. longiramosus Spikelets 6–12 mm long ............................................................................. 9 Inflorescence branches unbranched and densely flowered to the base, 4– 8 cm long ............................................................................. M. grandifolius Inflorescence branches rebranched and lacking spikelets near the base, lax, loosely flowered, 6–12 cm long ..................................... M. paludicolus Inflorescence pyramidal, with 40–100 stiff, divergent to spreading branches 2–6 cm long .......................................................................... 11 Inflorescence ovoid, with 4–25 lax, loosely ascending to divergent branches 8–30 cm long ........................................................................ 12 Spikelets 5.2–7 mm long, the upper glume with an abrupt awn 2–3.5 mm long; inflorescence branches 2–3.5 cm long ....................... M. churunensis Spikelets 7.5–13 mm long, the upper glume unawned or rarely subaristate; inflorescence branches 3–6 cm long ................. M. distantiflorus Spikelets 7.5–13 mm long .............................................................. M. exsertus Spikelets 4–7.2 mm long .......................................................................... 13 Spikelets 4–5.3 mm long, with 2 well-developed florets, the lower noticeably larger than the upper ............................................... M. longiramosus Spikelets 6.7–7.2 mm long, with 2 well-developed, equal florets .............. .................................................................................................... M. simplex

Myriocladus cardonae Swallen, Fieldiana, Bot. 28: 35. 1951. Shrub 1–4 m tall; leaf blades (10–)20–34 × (1.5–)2.5–4.5(–5.5) cm, clasping basally, the margins with distinctive clusters of tiny prickles; inflorescence 30–60 × 2–3 cm, with scattered divergent branches; spikelets 7–9 (–10.5) mm long, glabrous. Tepui summits, along streams, rocky shrublands, wet savannas, Bonnetia forests, 1700–2200 m; Bolívar (Auyán-tepui, Macizo del Chimantá). Endemic. ◆Fig. 117. Myriocladus churunensis Swallen, Acta Bot. Venez. 2: 132. 1967. Shrub 1–3 m tall; leaf blades 10–30 × 3– 5.5 cm, cordate basally; inflorescence 30–60 × 5–7 cm, with numerous divergent branches; spikelets often curved downward. Tepui summits, forming thickets along streams, in open areas, Bonnetia forests, 1600–1800 m; Bolívar (Auyán-tepui, Uei-tepui). Endemic.

Myriocladus purpureus Swallen, Mem. New York Bot. Gard. 9: 397. 1957. Myriocladus variabilis Swallen, Mem. New York Bot. Gard. 9: 396. 1957. Myriocladus wurdackii Swallen, Mem. New York Bot. Gard. 9: 398. 1957. Shrub 1–4 m tall; leaf blades 10–25(–40) × 2.3–4.5(–5.5) cm, often cuneate basally; inflorescence 25–50 × 4–10 cm, the branches rather densely flowered; spikelets 7.5–13 mm long. Streamlet banks, cliff bases, crevices, dwarf forests on tepui summits, 1700–2500 m; Bolívar (Aparamán-tepui, Aprada-tepui, Auyán-tepui, Carrao-tepui, Kamarkawaraitepui, Macizo del Chimantá, Ptari-tepui). Endemic. Myriocladus distantiflorus is closely related to M. exsertus. Both species have fine, matted, cottony oral setae; often lateral shoots with much smaller leaf blades only 1– 4 × 0.2–1 cm; and blade margins with clusters of tiny prickles.

Myriocladus distantiflorus Swallen, Mem. New York Bot. Gard. 9: 248, fig. 4d. 1957. Myriocladus confertus Swallen, Mem. New York Bot. Gard. 9: 397. 1957.

Myriocladus exsertus Swallen, Mem. New York Bot. Gard. 9: 242, fig. 3c. 1957. Shrub 0.5–3 m tall; leaf blades 6–27 × 1.2– 5.3 cm; inflorescence 15–35 × 4–10 cm, the

Myriocladus 161

branches 10–30 cm long, loosely ascending and not densely flowered. Shrubby areas along streamlets on tepui summits, 1700– 2700 m; Amazonas (Cerro Aracamuni, Cerro Duida, Cerro Huachamacari, Cerro Marahuaka, Cerro Yutajé). Endemic. Myriocladus grandifolius Swallen, Mem. New York Bot. Gard. 9: 245, fig. 3h. 1957. Myriocladus paraquensis Swallen, Mem. New York Bot. Gard. 9: 246, fig. 3i. 1957. Shrub 1–5 m tall; leaf blades 15–43 × 6–15 cm; inflorescence 35–70 × 6–12 cm; spikelets 8–11 mm long, the upper glume not prominently awned. Stream sides, savannas, open forests on tepui summits, (700–)1200–2200 m; Amazonas (Cerro Aracamuni, Cerro Duida, Cerro Sipapo). Brazil (Amazonas: Serra Aracá). Myriocladus involutus Judz. & Davidse, Novon 1: 83. 1991. Shrub ca. 1 m tall, forming dense colonies; oral setae fine, white, numerous; spikelets 7– 8 mm long. Tepui summits, understory of Chimantaea communities, 2100–2300 m; Bolívar (Macizo del Chimatá [Apacarátepui]). Endemic. ◆Fig. 116. Myriocladus longiramosus Swallen, Mem. New York Bot. Gard. 9: 243, fig. 3e. 1957. Myriocladus affinis Swallen, Mem. New York Bot. Gard. 9: 244, fig. 3f. 1957. Myriocladus paruensis Swallen, Mem. New York Bot. Gard. 9: 244, fig. 3g. 1957. Shrub 0.5–5 m tall; leaf blades (5–)15–50 × (1–)3–5.5(–8) cm, often strongly clasping basally; inflorescence 25–60 × 4–12 cm, pyramidal, the branches 5–12 cm long; spikelets awnless. Open rocky sandstone, thickets, woodland borders on tepui summits, (1000–) 1600–2200 m; Bolívar (Cerro Guaiquinima, Cerro Jaua), Amazonas (Cerro Coro Coro, Cerro Guanay, Cerro Parú, Cerro Yaví, Cerro Yutajé). Endemic. Myriocladus neblinaensis Swallen, Mem. New York Bot. Gard. 9: 240, fig. 1. 1957. Shrub 0.3–1.5 m tall; leaf blades 4.5–8 × 1.5–3 cm; peduncle to 15 cm long; spikelets 8–12 mm long, the upper glume 6–10 mm long, awnless. Moist valleys, gaps in humid forests, Heliamphora savannas on tepui slopes and summits, 1300–2200 m; Amazo-

nas (Cerro Aracamuni, Sierra de la Neblina). Adjacent Brazil. Myriocladus paludicolus Swallen, Mem. New York Bot. Gard. 9: 246, 248, fig. 4b. 1957. Shrub 2–5(–8) m tall; leaf blades 25–50 × 6–13 cm, often clasping basally; inflorescence 40–70 × 7–15 cm; spikelets 6–12 mm long, (3–)5–12-flowered, the upper glume with a recurved awn 2–6 mm long. Marshy savannas, Bonnetia forests, Euterpe swamps on tepui summits, 1700–2200(–2700) m; Amazonas (Sierra de la Neblina). Adjacent Brazil. ◆Fig. 119. Myriocladus simplex Swallen, Mem. New York Bot. Gard. 9: 242, fig. 2. 1957. Shrub 0.5–3 m tall; leaf blades 4–6 × 0.6–1 cm; inflorescence 17–27 × 4–6 cm, the 4–10 slender branches 8–12 cm long, loosely flowered, naked near the base; spikelets awnless. Along streams in savannas on tepui slopes, ca. 1300 m; Amazonas (Cerro Yutajé). Endemic. ◆Fig. 118. Myriocladus steyermarkii Swallen, Mem. New York Bot. Gard. 9: 395. 1957. Myriocladus gracilis Swallen, Mem. New York Bot. Gard. 9: 393, fig. 74. 1957. Shrub 0.3–1.5 m tall, forming dense thickets, culms much branched above; inflorescences both terminal and lateral, each subtended by a leaf; spikelets (5–)6–7 mm long, awnless. Stream banks and tepui summit meadows, 1900–2500 m; Bolívar (Macizo del Chimantá). Endemic. Myriocladus virgatus Swallen, Fieldiana, Bot. 28: 34, fig. 4. 1951. Myriocladus maguirei Swallen, Mem. New York Bot. Gard. 9: 239, fig. 3b. 1957. Shrub 1–5 m tall; leaf blades 35–100 × 1.5–4 cm, often splitting longitudinally, the midrib prominent on the lower surface; inflorescence 40–110 × 4–8 cm, with 100–275 branches 1.5–4 cm long bearing spikelets on their lower sides; spikelets with upper glume abruptly awned. Bare sandstone, stream sides, gaps in forests of tepui slopes and summits, (700–)1000–2000 m; Bolívar (Cerro Guaiquinima, Cerro Guanacoco, Cerro Sarisariñama, Serranía Marutaní), Amazonas (Cerro Aracamuni, Cerro Aratitiyope, Cerro Autana, Cerro Duida, Cerro Huachamacari,

162

P OACEAE

Cerro Marahuaka, Cerro Parú, Cerro Sipapo, Cerro Yutajé, Río Coro Coro). Brazil (Amazonas: Serra Aracá). Myriocladus sp. A Shrub 0.5–2.5 m tall; leaf blades 15–20 × 0.8–1 cm, cuneate basally; oral setae prominent, pale; both collections sterile. Thickets and shrublands on tepuis, 1700–2000 m; Bolívar (Cerro Jaua), Amazonas (Cerro Duida). Endemic.

Fig. 116. Myriocladus involutus

Myriocladus 163

Fig. 117. Myriocladus cardonae

Fig. 118. Myriocladus simplex

164

P OACEAE

Fig. 119. Myriocladus paludicolus

Neurolepis 165

56. NEUROLEPIS Meisn., Pl. Vasc. Gen., Tab. Diagn. 1: 426; 2: 325. 1843. [Subfamily Bambusoideae, Tribe Bambuseae] Planotia Munro, Trans. Linn. Soc. London, Bot. 26: 70. 1868, nom. illeg., superfl. (substituted for Platonia Kunth 1829, non Raf. 1810, nom. rejic., non Mart. 1832.) by Emmet J. Judziewicz Reed-like bamboos from sympodial rhizomes. Culms stout, unbranched above the base, solid but pithy, not lignified, the nodes not bearing buds. Leaf blades monomorphic or the basal leaves somewhat different from the upper, or often the leaves all restricted to a basal cluster; sheaths narrow, persistent; oral setae and outer ligules absent; inner ligules membranous, often prominent, often backed by a lacerate membrane that resembles another ligule; blades often very large (1–5 m × 10–30 cm), very thick and leathery, often long-attenuate at base, narrowed into a stout, sulcate pseudopetiole, persistent or deciduous. Inflorescence terminal, an open to strongly contracted panicle, often large, with a stout rachis and branches. Spikelets with glumes usually small, unequal, awned or not; 1 or 2 lowest florets sterile, reduced to empty lemmas; terminal floret functional, larger than the sterile florets, the lemma awned or not; stamens typically 3; stigmas 2. Caryopsis oblong, rarely seen. Costa Rica, Panama, Colombia, Venezuela, Trinidad-Tobago, northwestern Ecuador, Peru, Brazil, Bolivia; ca. 7–12 species, ca. 6 in Venezuela, 5 of these in the flora area. All flora area species belong to the difficult Neurolepis mollis Swallen complex. The presence or absence or pulvini at the base of the inflorescence branches, given considerable weight by Soderstrom (1969) in his tentative key to the genus, does not appear to be a reliable character. Dwarf-flowering forms sometimes occur, and the key does not account for these. Key to the Species of Neurolepis 1. 1. 2(1). 2.

3(2). 3. 4(3). 4.

Spikelets with lemmas minutely scabrid-puberulent throughout ... N. sp. A Spikelets with lemmas glabrous ............................................................... 2 Functional lemma awned, the awn 1–2 mm long; inflorescence 10– 20 times as long as wide; spikelets 6–9 mm long ..................... N. angusta Functional lemma awnless or subaristate, the awn if present < 1 mm long; spikelets 4–6 mm long; inflorescence 4–8 (–20) times as long as wide ........................................................................................................ 3 Glumes with awns ca. 0.5 mm long; leaf blades ca. 50 × 3.5 cm ............... ........................................................................................... N. diversiglumis Glumes awnless; leaf blades 100–250 × 3–12 cm ..................................... 4 Spikelets 5.3–6 mm long, appressed to the branches; leaf blades 3–5 cm wide ......................................................................................... N. glomerata Spikelets 4.5–5.5 mm long, spreading in all directions from the branches; leaf blades 6–12 cm wide ............................................................. N. pittieri

Neurolepis angusta Swallen, Mem. New York Bot. Gard. 9: 249. 1957. Erect bamboo 1–2 m tall; leaves all basal in a cluster 2 m tall, the blades 40–65 × 2–6.5

cm; inflorescence 40–80 × 2–5 cm. Gaps in montane forests, 2000–2600 m; Bolívar (Aprada-tepui, Caraurín-tepui, Macizo del Chimantá). Colombia (Santander).

166

P OACEAE

Fig. 120. Neurolepis glomerata

Olyra

Neurolepis diversiglumis Soderstr., Mem. New York Bot. Gard. 18: 16, figs. 2, 3. 1969. Erect bamboo 1 m tall; inflorescence ca. 40 × 5 cm, with numerous, densely flowered branches ascending at 45° angles; spikelets 5–5.5 mm long. Frequent along open slopes at base of cliffs on tepui peak, 2600–2800 m; Amazonas (Pico Phelps on Sierra de la Neblina). Endemic. Neurolepis glomerata Swallen, Mem. New York Bot. Gard. 9: 399. 1957. Neurolepis densiflora Swallen, Mem. New York Bot. Gard. 9: 399. 1957. Neurolepis nigra Swallen, Mem. New York Bot. Gard. 9: 400. 1957. Erect bamboo; leaves all basal in a cluster 2 m tall, inflorescences to 10 m tall; leaf blades 55–125 cm long; inflorescence 60–80 × 4–20 cm, the spreading lowest branches to 11 cm long; spikelets with functional lemma acute or with an awn < 1 mm long. Open slopes, Bonnetia forests, and Chimantaea community on tepuis, 2000–2300 m; Bolívar

167

(Macizo del Chimantá). Andes; Colombia (Santander). ◆Fig. 120. Neurolepis pittieri McClure, J. Wash. Acad. Sci. 32: 181, fig. 8. 1942. Erect bamboo; leaves all basal in a cluster to 3 m tall, the inflorescences taller; leaf blades 100–250 m long; inflorescence 125 × 10 cm; spikelets with functional lemmas acute or with an awn to 0.5 mm long. Forests, 1800–2800 m; Bolívar (Cerro Jaua), Amazonas (Sierra de la Neblina, Sierra de Maigualida). Venezuelan Coastal Cordillera; Costa Rica, Panama, northern Colombia. Interpreted in the very widest sense, the oldest name for this taxon is Neurolepis mollis Swallen, J. Wash. Acad. Sci., 21: 14. 1931. Neurolepis sp. A Erect bamboo 2–3 m tall; leaves mostly basal, the blades 125 × 4 cm; inflorescence 75 × 0.7 cm, the lowermost branches to 3 cm long, strictly appressed; spikelets (immature) 3–3.5 mm long, awnless. Montane forests, 1000–1400 m; Bolívar (Cerro Venamo).

57. OLYRA L., Syst. Nat. ed. 10, 1261. 1759. [Subfamily Bambusoideae, Tribe Olyreae] by Emmet J. Judziewicz Monoecious, cespitose herbs of perennial or monocarpic habit. Culms erect or scandent, weak, hollow. Leaves with membranous ligules; pseudopetioles present, short; blades usually broad. Inflorescence of 1–many panicles from the uppermost nodes; panicles usually open, pyramidal, or occasionally digitate or spicate; pistillate spikelets large, usually borne on clavate pedicels; staminate spikelets usually small, borne on filiform pedicels. Spikelets 1-flowered. Staminate spikelets linear to lanceolate, hyaline, early deciduous; glumes nearly always absent; lemma 3–9veined, acuminate to aristate; palea nearly as long as lemma; stamens 3. Pistillate spikelets with floret disarticulating from the persistent glumes, or sometimes the glumes falling first, or the spikelet falling entire; glumes acuminate to long-awned, equal to unequal, many-veined, membranous, longer than the floret; floret lanceolate to ovate, indurate, white, becoming dark at maturity, glabrous to pubescent, smooth to pitted; lemma with margins usually covering the edges of the palea; stigmas 2, plumose. Neotropics, tropical Africa; 24 species, 8 in Venezuela, all in the flora area. Key to the Species of Olyra 1. 1. 2(1). 2.

Pistillate florets pitted or granular ........................................................... 2 Pistillate florets smooth ............................................................................. 5 Pistillate florets 2.5–3 mm long ................................................ O. micrantha Pistillate florets 5–9 mm long ................................................................... 3

168

3(2). 3. 4(3). 4. 5(1). 5. 6(5). 6. 7(6). 7.

P OACEAE

Pistillate and staminate spikelets on separate, spicate, conjugate branches ................................................................................... O. longifolia Pistillate and staminate spikelets mixed in an open, pyramidal panicle ................................................................................................................ 4 Pistillate glumes 8–12 mm long, ovate ........................................ O. ecaudata Pistillate glumes 13–20 mm long, narrowly lanceolate ............. O. standleyi Pistillate florets pubescent with silky hairs ............................. O. ciliatifolia Pistillate florets glabrous .......................................................................... 6 Staminate spikelets distinctly longer than the pistillate spikelets ................................................................................................. O. wurdackii Staminate spikelets much shorter than the pistillate spikelets ............. 7 Pistillate glumes with awns 25–35 mm long; pistillate florets 7–8 mm long ............................................................................................. O. caudata Pistillate glumes with awns 5–15(–20) mm long; pistillate florets 5–6 mm long ............................................................................................. O. latifolia

Olyra caudata Trin., Linnaea 10: 292. 1836. Cespitose perennial 1–2 m tall; leaf blades 20–30 × 5–8 cm, rounded basally; pistillate glumes with long, scissors-like awns. Wet forests, ca. 200 m; Bolívar (Las Pavas at Salto Pará). Apure, Barinas, Monagas, Táchira; Costa Rica, Panama, Colombia, Trinidad-Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Amazonian Brazil, Bolivia. ◆Fig. 121. Olyra ciliatifolia Raddi, Agrost. Bras. 19. 1823. —Paja carrizo. Cespitose perennial 0.5–1 m tall; leaf blades 8–13 × 2–4 cm, obliquely truncate basally; pistillate spikelets 20–30 mm long; glumes with delicate, flexuous awns; florets 7–8 mm long. Moist, often disturbed forests, 100–200 m; northern Bolívar. Barinas, Guárico, Portuguesa, Sucre; Colombia, TrinidadTobago, Guyana, French Guiana, Brazil, Bolivia, Paraguay, Argentina. ◆Fig. 123. Olyra ecaudata Döll in Mart., Fl. Bras. 2(2): 326. 1877. Cespitose perennial or often monocarpic; sterile, leafy culms 1–2 m tall, the leafless flowering ones 2–3 m long, weak, scandent, or rarely inflorescences borne on leafy culms; leaf blades 20–30 × 4–5 cm, slightly subcordate basally; pistillate spikelets with glumes awnless; florets 6–8 mm long. Wet forests, 200–800 m; Bolívar (Amaruay-tepui, Río Nichare), Amazonas (Caño Baboto on upper Río Cuao). Zulia; Nicaragua, Costa Rica, Panama, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia.

Olyra latifolia L., Syst. Nat. ed. 10, 1261. 1759. —Carrizo, Pitillo. Olyra cordifolia Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 198. 1815 [1816]. Cespitose perennial, erect at first, becoming scandent with age, the culms to 6 m long; leaf blades 10–35 × 2–9 cm, rounded basally; pistillate spikelets 15–23 mm long. Moist to wet forests, often in gaps and second growth, 50–900 m; Delta Amacuro (Serranía de Imataca), Bolívar (widespread), Amazonas (widespread). Common and widespread throughout rest of Venezuela and tropical America and Africa. Olyra longifolia Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 198. 1815 [1816]. —Caricillo, Paja perro de agua. Cespitose perennial 1–2 m tall; leaf blades 15–30 × 2.5–5 cm, slightly subcordate basally; branches of inflorescence 4–8 cm long; pistillate spikelets 15–22 mm long, the slender glumes purple-tipped. Forest and river margins, often in rocky or seasonally flooded areas, near sea level to 500 m; widespread in Delta Amacuro, Bolívar, and Amazonas. Apure, Guárico; Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Amazonian Brazil, Bolivia. ◆Fig. 122. Olyra micrantha Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 199. 1815 [1816]. —Pitillo. Cespitose perennial 1–2 m tall; leaf blades 20–35 × 6–12 cm, rounded basally; inflores-

Olyra

Fig. 121. Olyra caudata

169

170

P OACEAE

Fig. 122. Olyra longifolia

Fig. 123. Olyra ciliatifolia

Olyra 171

Fig. 124. Olyra micrantha

172

P OACEAE

cence copiously flowered, the upper portion pistillate, the lower staminate; pistillate spikelet pedicels filiform, the spikelets falling entire. Gallery or rocky upland forests, in gaps or dense shade, often common on the lower slopes of tepuis, 100–1000 m; Bolívar (El Paují, Gran Sabana, Río Paragua, Río Parguaza), Amazonas (Cerro Aracamuni, Cerro Aratitiyope, Cerro Duida, Cerro Huachamacari, Río Orinoco basin). Anzoátegui, Apure, Falcón, Lara, Mérida, Táchira, Zulia; Colombia, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia, Paraguay, Argentina. ◆Fig. 124. Olyra micrantha is now usually recognized as Parodiolyra micrantha (Kunth) Davidse & Zuloaga (Novon 9: 590. 1999). Olyra standleyi Hitchc., Proc. Biol. Soc. Wash. 40: 86. 1927. Olyra fasciculata auct. non Trin. 1834.

Cespitose perennial (or monocarpic) 1–2 m tall; leaf blades 20–26 × 4–5.3 cm, slightly subcordate basally; pistillate spikelets 13–20 mm long; florets ca. 7 mm long. Riparian lowland forests, upper montane forests, 100– 1900 m; Amazonas (Caño San Miguel, Serranía Uasadi). Distrito Federal, Guárico, Lara, Zulia; Costa Rica, Panama, Colombia. Outside of the flora area Olyra standleyi is usually a species of cloud forests and is found mostly above 1000 m. Olyra wurdackii Swallen, Phytologia 14: 85. 1966. Cespitose perennial 1–4 m tall; leaf blades ca. 25 × 5 cm; inflorescence 30 cm long; pistillate spikelets 7–7.5 mm long; staminate spikelets 9–11 mm long. Open rock on granitic outcrops, 300–500 m; Amazonas (Río Siapa). Brazil (Amazonas).

58. OPLISMENUS P. Beauv., Fl. Oware 2: 14. 1810, nom. cons. [Subfamily Panicoideae, Tribe Paniceae] by Emmet J. Judziewicz Plants creeping, the culms decumbent and rooting at nodes, forming large vegetative colonies. Leaves with ligules membranous-ciliate; blades linear to ovate. Inflorescence a pyramidal panicle of distant, ascending racemes; racemes short, secund, the rachis triquetrous, bearing paired spikelets from 2 sides. Spikelets falling entire, somewhat laterally compressed, 2-flowered, the bracts (especially the glumes) short to long-awned from a bifid apex; lower glume distinctly shorter than the spikelet, 3–7-veined; upper glume shorter than to nearly as long as the spikelet, 5–7-veined, long-awned; lower floret sterile, the lemma as long as the spikelet, 5–9veined, usually short-awned, the palea absent; upper floret concealed by the lower floret, the lemma narrowly elliptic to lanceolate, indurate, shining, the margins strongly inrolled over the palea. Worldwide in tropical and warm-temperate area; 5–15 species, 2 in Venezuela, both in the flora area. Key to the Species of Oplismenus 1. 1.

Awn of upper glume slender, antrorsely scabrous; racemes densely whitehairy ....................................................................................... O. burmannii Awn of upper glume stout, smooth, viscid; racemes sparingly pubescent .................................................................................................... O. hirtellus

Oplismenus burmannii (Retz.) P. Beauv., Ess. Agrostogr. 54, 169. 1812. —Panicum burmannii Retz., Observ. Bot. 3: 10. 1791 [1783]. —Mitë. Sprawling, mat-forming plant, culms to 40 cm tall; leaf blades 3.5–7 cm × 8–13 mm;

inflorescence 3–7 cm long, of 3–7 racemes, each ca. 1 cm long; spikelets ca. 3.5 mm long. Trail sides in forests, ca. 100 m; Bolívar (Moitaco, Río Maniapure). Anzoátegui, Apure, Aragua, Carabobo, Cojedes, Falcón, Guárico, Lara, Miranda, Portuguesa, Táchira, Zulia;

Oplismenus 173

Fig. 125. Oplismenus burmannii

Fig. 126. Oplismenus hirtellus

174

P OACEAE

Mexico, Central America, Greater Antilles, Colombia, Ecuador, Peru, Brazil, Bolivia, Paleotropics. ◆Fig. 125. Oplismenus hirtellus (L.) P. Beauv., Ess. Agrostogr. 54, 168, 170. 1812. —Panicum hirtellum L., Syst. Nat. ed. 10, 870. 1759. Sprawling, mat-forming plant, the culms to 50 cm long; leaf blades 6–13 cm × 7–25 mm; inflorescence 4–13 cm long, of 3–7

racemes, each 1–2(–3) cm long; spikelets ca. 3 mm long, with awns 5–10 mm long. Pastures, forest edges, lajas, near sea level to 1000(–1700) m; Delta Amacuro (near Los Castillos, Tucupita), Bolívar (Altiplanicie de Nuria, Gran Sabana, San Martín de Turumbán southwest of Tumeremo), Amazonas (west of Cerro Yutajé). Widespread elsewhere in Venezuela and tropical and subtropical regions worldwide. ◆Fig. 126.

59. ORTHOCLADA P. Beauv., Ess. Agrostogr. 69. 1812. [Subfamily Arundinoideae, Tribe Centotheceae] by Gerrit Davidse Cespitose perennials. Leaf sheaths and blades with hook-shaped microhairs; ligule membranous; blades on prominent pseudopetioles, elliptic, conspicuously cross-veined, bearing hooked microhairs. Inflorescence a large and diffuse panicle; pedicels long, filiform. Spikelets bisexual, 2- or 3-flowered, laterally compressed and keeled, disarticulating below glumes and second floret, the 2 lower florets bisexual, the uppermost sterile and rudimentary; glumes subequal, the lower 2veined, the upper 5-veined; lemmas 5–7-veined, acuminate or awn-tipped; rachilla slender, elongate, held by the keels of the palea. Neotropics, tropical Africa; 2 species, 1 in Venezuela. Orthoclada laxa (Rich.) P. Beauv., Ess. Agrostogr. 70, 149. 1812. —Aira laxa Rich., Actes Soc. Hist. Nat. Paris 1: 106. 1792. Panicum rariflorum Lam., Encycl. 4: 746. 1798. —Orthoclada rariflora (Lam.) P. Beauv., Ess. Agrostogr. 70, 168, 170, t. 14, fig. 9. 1812. Orthoclada rariflora var. lanceolata Döll in Mart., Fl. Bras. 2(3): 118. 1878. Orthoclada rariflora var. sesquiflora Döll

in Mart., Fl. Bras. 2(3): 118, fig. 35. 1878. Broad-leaved perennial, with prominent pseudopetioles and very diffuse inflorescences. Forests, near sea level to 500 m; Delta Amacuro (vicinity of Epaña, vicinity of Tucupita), widespread in Bolívar and Amazonas. Barinas, Portuguesa, Táchira, Zulia; Mexico, Guatemala, Honduras, Nicaragua, Costa Rica, Panama, Caribbean, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 127.

60. ORYZA L., Sp. Pl. 333. 1753. [Subfamily Bambusoideae, Tribe Oryzeae] by Emmet J. Judziewicz Annuals to perennials. Leaves with ligules membranous; blades narrow. Inflorescence a terminal panicle with few branches, the spikelets often overlapping. Spikelets bisexual, 1-flowered, elliptic, asymmetrically apiculate, usually strongly laterally compressed, deciduous or persistent; glumes not evident or reduced to a minute ridge or cupule at the summit of the pedicel; sterile lemmas 2, glume-like, unequal to subequal, greenish, rigid, smooth, lanceolate, much shorter than to less commonly as long as the spikelet; lemma and palea ± equal, the lemma keeled, 5veined, often strongly antrorsely hispid-ciliate on the margins and veins, in texture regularly granular-papillate, long-awned or not; palea keeled, 3-veined, hispid-ciliate on the keel and lateral veins; stamens 1–6.

Orthoclada 175

Fig. 127. Orthoclada laxa

176

P OACEAE

Worldwide in tropical and warm-temperate areas; ca. 20 species, 3 in Venezuela, all in the flora area. Key to the Species of Oryza 1. 1. 2(1). 2.

Plants annual; spikelets persistent, broadly elliptic, 3–3.5 mm wide, often awnless; cultivated rice ...........................................................O. sativa Plants perennial; spikelets readily deciduous, generally narrowly elliptic, 1.8–2.5 mm wide, usually awned .......................................................... 2 Fertile lemma without an oblique, purplish, bearded constriction at the summit of the body .................................................................... O. latifolia Fertile lemma with an oblique, purplish, bearded constriction separating the body from the awn ............................................................. O. rufipogon

Oryza latifolia Desv., J. Bot. Agric. 1: 77. 1813. Oryza alta Swallen, Publ. Carnegie Inst. Wash. 461: 156. 1936. Perennial 1–4 m tall; leaf blades 1.5–4 cm wide; body of spikelet 6–10 × 2–2.5 mm. Stream banks, marshes, near sea level to 200 m; Delta Amacuro (Caño Joba-Suburu east of Caño Sacupana, Río Toro), Bolívar (Ciudad Bolívar, Río Macoita), Amazonas (upper Río Orinoco). Anzoátegui, Apure, Carabobo, Falcón, Guárico, Portuguesa, Yaracuy; Mexico, West Indies, Central America, tropical South America. ◆Fig. 128. The epidermis on the culms of Oryza latifolia is used by the Yanomami for cutting hair. Oryza rufipogon Griff., Ic. Pl. Asiat. 3: 5, pl. 144, fig. 2. 1851. Oryza perennis Moench, Methodus 197.

1794, nom. confus. Perennial 1–2 m tall; body of spikelet 8–9 × 1.8–2.5 mm. Swamps, lagoons, Mauritia palm swamps, near sea level to 200 m; Delta Amacuro (Sacupana, Tucupita), Bolívar (Altiplanicie de Nuria, Ciudad Piar, Serranía de los Pijiguaos). Apure, Barinas, Cojedes, Guárico, Portuguesa; naturalized and seemingly native throughout tropical America and tropical southeastern Asia. ◆Fig. 129. Oryza sativa L., Sp. Pl. 333. 1753. —Arisi, Arroz. Annual 0.5–1 m tall; body of spikelet 10 × 3–3.5 mm. Occasionally escaped from cultivation; Delta Amacuro (Caño Araguao), Bolívar (Río Parguaza), Amazonas (Río Cataniapo). Carabobo, Sucre; widely cultivated in tropical America and occasionally escaping but not persisting, native to tropical southeastern Asia.

61. OTACHYRIUM Nees in Mart., Fl. Bras. Enum. Pl. 2: 271. 1829. [Subfamily Panicoideae, Tribe Paniceae] by Emmet J. Judziewicz Annuals or perennials. Inflorescence terminal, a pyramidal panicle, the spikelets borne singly or in pairs on capillary pedicels. Leaves with ligules membranousciliate; blades linear. Spikelets obliquely set on pedicels, orbicular in outline, often widely gaping at maturity, falling entire, 2-flowered; glumes subequal, small, rounded, membranous, not strongly veined, the upper glume often somewhat inflated and forming a pocket for the development of the upper floret; lower floret staminate, membranous to coriaceous, the lemma narrowly oblong, 3-veined, sulcate, nearly as long as the spikelet, the palea as long as the spikelet, membranous, the keels widely spaced, often winged, the margins beyond the keels often winged and forming a partial tube around the staminate flower; upper floret bisexual,

Oryza 177

Fig. 128. Oryza latifolia

Fig. 129. Oryza rufipogon

178

P OACEAE

ovate-lanceolate, plano-convex, slightly beaked, papery to indurate, pale when young, often brown when mature, smooth, shining, the lemma only loosely embracing the palea. Colombia, Venezuela, Trinidad-Tobago, Guyana, Suriname, French Guiana, Brazil, Bolivia, Paraguay, Argentina; 7 species, 2 in Venezuela, both in the flora area. Key to the Species of Otachyrium 1. 1.

Spikelets 3.3–5.5 mm long .................................................... O. grandiflorum Spikelets 1.8–2.7 mm long ........................................................... O. versicolor

Fig. 130. Otachyrium versicolor

Panicum 179

Otachyrium grandiflorum Send. & Soderstr., Smithsonian Contr. Bot. 57: 7. 1984. Cespitose perennial 30–100 cm tall; leaf blades 10–28 cm × 1–6 mm, involute to less commonly flat; inflorescence 3–13 cm long, the few branches to 4 cm long; lower palea when expanded 3.5–5 mm wide, light purplish, the spikelet resembling a petaliferous flower. White-sand savannas, ca. 100 m; Amazonas (Caño Caname, Caño Yagua). Brazil (Amazonas, Goiás, Roraima). Otachyrium versicolor (Döll) Henrard, Blumea 4: 511. 1941. —Panicum versi-

color Döll in Mart., Fl. Bras. 2(2): 254. 1877. Rhizomatous, cespitose perennial 50– 100 cm tall; leaf blades 13–25 × 0.5–1 cm; inflorescence 6–25 cm long, with numerous branches; lower palea when expanded to 2.5 mm wide. Seasonally wet savannas, Mauritia palm swamps, marshes, often forming extensive stands, 100–900 m; Bolívar (Río Orinoco basin and tributaries), Amazonas (Río Orinoco basin and tributaries). Anzoátegui, Apure, Guárico, Monagas; Colombia, Trinidad-Tobago, Guyana, Brazil, Bolivia, Paraguay, Argentina. ◆ Fig. 130.

62. PANICUM L., Sp. Pl. 55. 1753. [Subfamily Panicoideae, Tribe Paniceae] by Fernando O. Zuloaga Annuals or perennials, stoloniferous, decumbent and rooting at the lower nodes, to erect and cespitose; internodes hollow to solid; aerenchyma present or absent. Ligules membranous, membranous-ciliate, or ciliate; blades linear to ovatelanceolate, flat or inrolled. Inflorescence an open to contracted panicle, in some species branches racemose; spikelets usually disarticulating entire from the pedicels. Spikelets dorsally compressed, 2-flowered; lower glume commonly present, 1/6–4/5 the length of the spikelet, 1–3(–5)-veined; stipe conspicuous or not between the lower and upper glume; upper glume 3–14-veined, subequal with lower lemma; lower palea present or absent; lower flower pistillate or absent; upper floret indurate, muticous, usually pale, glabrous or with 2-celled microhairs, or rarely with long macrohairs, smooth or rarely rugose, simple or compound papillae present or absent; lemma 5–7-veined, palea 2-veined; lodicules 2, truncate; stamens usually 3; stigmas 2, plumose. Caryopsis with a punctiform to oblong hilum; embryo 1/2 the length of the caryopsis. Cosmopolitan (mostly lowland habitats); ca. 500 species, ca. 70 in Venezuela, 53 of these in the flora area. Key to the Species of Panicum 1. 1. 2(1). 2. 3(2). 3. 4(3).

Spikelets on racemose branches, secund .................................................. 2 Spikelets in open or contracted panicles, not in secund, racemose branches ................................................................................................. 7 Lower lemma with 2 crateriform glands on the middle portion ................ ................................................................................................ P. pulchellum Lower lemma without crateriform glands on the middle portion ........... 3 Secondary branches of panicle absent, main axis pilose; ligules usually absent .......................................................................................... P. pilosum Secondary branches of panicle present, main axis glabrous; ligules membranous ................................................................................................... 4 Upper floret 1/2 the length of the spikelet, short-stipitate, glabrous and without papillae ................................................................... P. stoloniferum

180

P OACEAE

4. 5(4). 5. 6(5). 6. 7(1). 7. 8(7). 8. 9(7). 9. 10(9). 10.

11(10). 11. 12(11). 12. 13(9). 13. 14(13). 14. 15(14). 15. 16(15). 16. 17(16). 17. 18(15). 18. 19(18). 19. 20(14). 20. 21(20). 21.

Upper floret as long as the spikelet, not stipitate, with prickly hairs at the apex or glabrous, usually papillose ................................................ 5 Blades clasping the stem, cordate basally ................................. P. hylaeicum Blades not clasping the stem, rounded to subcordate basally ................. 6 Blades linear-lanceolate, narrowed at the base; upper floret glabrous, not papillose; caryopsis black ....................................................... P. scabridum Blades lanceolate, subcordate at the base; upper floret with prickly hairs at the apex, papillose; caryopsis brownish ........................ P. polygonatum Panicles contracted, 2–25-flowered; blades filiform ................................. 8 Panicles open, usually > 25-flowered; blades ovate-lanceolate to linearlanceolate, not filiform .......................................................................... 9 Panicles with 2–10 spikelets; spikelets (1.8–)2–2.3 mm long, with a conspicuous internode between the lower and upper glume ....... P. caricoides Panicles with 10–25 spikelets; spikelets 1.1–1.7 mm long, without an internode between the lower and upper glume ............................ P. stenodes Upper floret transversely rugose ............................................................ 10 Upper floret smooth ................................................................................. 13 Culms erect, 1–3 m tall; ligules membranous-ciliate; spikelets narrowly elliptic, 3–3.8 mm long; lower flower pistillate ...................... P. maximum Culms decumbent and rooting at the lower nodes; ligules membranous; spikelets elliptic to obovate, 0.9–2.5 mm long; lower flower absent .............................................................................................................. 11 Spikelets 0.9–1.5 mm long; blades 2–7 cm long ......................... P. trichoides Spikelets 1.6–2.5 mm long; blades 6–14 cm long ................................... 12 Spikelets congested on upper portion of the branches ................... P. sellowii Spikelets regularly disposed on the branches .......................... P. millegrana All branches of the panicle whorled ............................................. P. mertensii Branches of the panicle alternate to opposite, not whorled ................... 14 Blades pungent; upper floret glabrous .................................................... 15 Blades not pungent (if pungent upper floret with 2-celled microhairs) .... 20 Culms decumbent, rooting and branching at the lower nodes .............. 16 Culms erect .............................................................................................. 18 Spikelets 3.9–4.4 mm long ....................................................... P. steyermarkii Spikelets 2.7–3.5 mm long ....................................................................... 17 Ligules absent, ligular region differentiated; spikelets 3.1–3.5 mm long ................................................................................................... P. fontanale Ligules membranous-ciliate, small, 0.3–0.7 mm long, ligular region not differentiated; spikelets 2.7–3.1 mm long ............................. P. sipapoense Ligules membranous-ciliate, 0.7–1 mm long ............................... P. jauanum Ligules absent .......................................................................................... 19 Spikelets (1.7–)2.2–2.8 mm long ................................................... P. chnoodes Spikelets (2.8–)3.1–3.7 mm long ............................................... P. eligulatum Inflorescences with both chasmogamous and cleistogamous flowers; lower palea absent ............................................................................... 21 Inflorescences with chasmogamous flowers only; lower palea present or absent ................................................................................................... 22 Spikelets 2.1–2.4(–2.8) mm long; blades asymmetric, 1.5–6 cm long ............................................................................................... P. pantrichum Spikelets 3.3–4.5 mm long; blades symmetric, 3–14 cm long ..... P. cordovense

Panicum 181

22(20). Upper floret bearded at apex and base; upper glume and lower lemma 3-veined, lower glume absent to minute ................................ P. discrepans 22. Upper floret glabrous or pilose only at base or apex; upper glume and lower lemma 5–9-veined, lower glume present, 1/6–4/5 the length of the spikelet ................................................................................................. 23 23(22). Lower glume 1/6–1/3 the length of the spikelet ......................................... 24 23. Lower glume 1/3–4/5 the length of the spikelet ......................................... 25 24(23). Plants perennial, aquatic, the culms succulent; spikelets lanceolate, 4.5– 5.6 mm long; upper floret 3/4 the length of the spikelet ..... P. elephantipes 24. Plants annual, not aquatic, the culms not succulent; spikelets ovate, 2.4– 3 mm long; upper floret as long as the spikelet .......... P. dichotomiflorum 25(23). Culms conspicuously branching, bamboo-like, with one branching above each node; plants with strong rootstocks ........................................... 26 25. Culms not branching as above; plants without strong rootstocks ......... 27 26(25). Spikelets glabrous; upper glume and lower lemma 7–9-veined; upper floret nonstipitate; panicle 20–40 cm long ........................ P. tricholaenoides 26. Spikelets pilose; upper glume and lower lemma 5-veined; upper floret stipitate; panicle ca. 20 cm long .............................................. P. deciduum 27(25). Upper floret stipitate ............................................................................... 28 27. Upper floret nonstipitate ......................................................................... 31 28(27). Spikelets 2.1–3.5 × 1–1.2 mm; upper floret 1.5–2.2 mm long; panicles terminal and axillary from the upper nodes, forming a compound inflorescence ..................................................................................................... 29 28. Spikelets 4.4–9 × 1.2–2.5 mm; upper floret 2.5–3.2 mm long; panicles strictly terminal ................................................................................... 30 29(28). Plants annual; spikelets obovate, 2.1–2.6 mm long, glabrous .... P. cayennense 29. Plants perennial; spikelets ovate, 3–3.5 mm long, sparsely hirsute ........................................................................................................ P. rudgei 30(28). Spikelets 4.4–5.7 mm long; leaf sheaths papillose-pilose with glass-like hairs; upper floret with stipe glabrous basally ......................... P. ligulare 30. Spikelets 5.5–9 mm long; leaf sheaths without glass-like hairs; upper floret with stipe pilose basally .................................................. P. cervicatum 31(27). Upper floret conspicuously pilose at the base with flattened hairs; panicle falling entire at maturity; spikelets 6–8.5 mm long ................ P. olyroides 31. Upper floret glabrous; panicle not falling at maturity; spikelets 1.5–4 mm long ....................................................................................................... 32 32(31). Sheaths papillose-pilose with glass-like hairs............................. P. hirsutum 32. Sheaths pilose or glabrous, without glass-like hairs ............................. 33 33(32). Upper glume and lower lemma 7–9-veined; ligules shortly membranous at base, ciliate at the apex; upper floret pilose or glabrous at the apex, otherwise glabrous ............................................................................... 34 33. Upper glume and lower lemma 5-veined; ligules membranous; upper floret covered with 2-celled microhairs ................................................... 35 34(33). Spikelets ovate, gaping at maturity; plants 2–3 m tall .................... P. altum 34. Spikelets broadly elliptic to elliptic, not gaping at maturity; plants 0.2– 1 m tall ............................................................................... P. hispidifolium 35(33). Aquatic plants 1–3 m tall; blades 50–70 cm long; panicles 30–70 cm long ....................................................................................................... P. grande 35. Terrestrial plants to 1 m tall; blades 1–21 cm long; panicles 1.5–18 cm

182

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long ....................................................................................................... 36 36(35). Plants 1–2 m tall; blades 10–21 cm long, cordate and clasping at the base; inflorescences 20–28 cm long .............................................................. 37 36. Plants 0.05–0.8 m tall; blades 0.7–14 cm long, not cordate and clasping at the base; inflorescences 3–18 cm long .......... ...................................... 38 37(36). Spikelets 1.8–2.4 mm long; upper floret with macrohairs; blades 6– 8.5 mm wide ............................................................................ P. tepuianum 37. Spikelets 1.1–1.7 mm long; upper floret without macrohairs; blades 8– 15 mm wide .............................................................................. P. nervosum 38(36). Spikelets obliquely set on the pedicels, papillose-pilose with long hairs ....................................................................................................... P. hirtum 38. Spikelets straight (not obliquely set on the pedicels), glabrous to shortpilose, not papillose-pilose................................................................... 39 39(38). Spikelets pear-shaped, lower and upper glume separated by a long to short stipe ............................................................................................ 40 39. Spikelets elliptic to obovate or globose, not pear-shaped, lower and upper glume without a stipe .......................................................................... 43 40(39). Branches of the panicle falling at maturity ............................. P. pyrularium 40. Spikelets falling without the branches at maturity ............................... 41 41(40). Spikelets ca. 2.3 mm long; upper floret stipitate; lower palea absent .................................................................................................. P. ichunense 41. Spikelets 1.1–1.6 mm long; upper floret nonstipitate; lower palea present or absent ............................................................................................... 42 42(41). Blades linear-lanceolate, rounded at base, 1.8–4 cm long .............. P. arctum 42. Blades lanceolate, cordate to subcordate at base, 1.6–4 cm long ..... P. rivale 43(39). Lower palea and flower absent ............................................................... 44 43. Lower palea and flower present .............................................................. 47 44(43). Spikelets 2.3 mm long; blades 6–12 × 0.6–1 cm; ligules membranousciliate ................................................................................... P. haenkeanum 44. Spikelets 0.9–1.9 mm long; blades 0.7–4 × 0.1–1 cm; ligules membranous .............................................................................................................. 45 45(44). Blades linear-lanceolate, 0.1–0.2 cm wide; spikelets obovate ................... ................................................................................................ P. polycomum 45. Blades lanceolate, 0.2–1 cm wide; spikelets elliptic ............................... 46 46(45). Blades subcordate to cordate at base, stem-clasping, 0.2–0.6(–1) cm wide; plants 5–20 cm tall ..................................................................... P. pandum 46. Blades narrowed at base, not stem-clasping, 0.2–0.3 cm wide; plants 15– 25 cm tall ................................................................................. P. fonticolum 47(43). Spikelets (1.1–)1.2–2.3 mm long ............................................................. 48 47. Spikelets 0.9–1.1(–1.5) mm long ............................................................. 51 48(47). Blades linear-lanceolate, 0.1–0.2 cm wide; plants cespitose, short-rhizomatous; panicles 3–5 cm long ........................................... P. orinocanum 48. Blades lanceolate, 0.2–1.5 cm wide; plants decumbent, rooting at the lower nodes, occasionally long-rhizomatous (cespitose in P. cyanescens but blades lanceolate, 0.3–1.5 cm wide, and panicles 7–18 cm long); panicles (1–)4–18 cm long ................................................................... 49 49(48). Plants annual; spikelets hispid; lower floret absent ................. P. yavitaense 49. Plants perennial; spikelets glabrous, lower floret staminate ................ 50 50(49). Plants 6–30(–40) cm tall; blades 0.7–3.3(–4) × 0.2–0.7 cm, often rather

Panicum 183

50. 51(47). 51. 52(51). 52. 53(52). 53. 54(53). 54. 55(51). 55. 56(55). 56.

distant and reflexed on the trailing culms; panicle (1–)4–6 cm long ............................................................................................... P. parvifolium Plants (20–)30–85 cm tall; blades 2–14 × 0.3–1.5 cm, not distant and not reflexed on the trailing culms; panicle 5–18 cm long ........... P. cyanescens Blades lanceolate, 0.2–0.8(–1.1) cm wide, often bluish-glaucous; plants decumbent, rooting and branching at the lower nodes ...................... 52 Blades linear-lanceolate to lanceolate, 0.1–0.4 cm wide; plants cespitose, culms occasionally decumbent ............................................................ 55 Spikelets 1–1.8 mm long; blades reflexed on the trailing culms; panicle (1–)4–6 cm long ..................................................................... P. parvifolium Spikelets 0.9–1.5 mm long; blades not reflexed on the trailing culms; panicle 6–17 cm long ........................................................................... 53 Plants annual; spikelets hispid .................................................. P. yavitaense Plants perennial; spikelets glabrous, rarely hispid ............................... 54 Plants cespitose with erect culms; panicles 1.3–5 cm long, blades 0.2– 0.4 cm wide ................................................................................. P. petrense Plants decumbent, rooting and branching at the base, panicles 4.5–10 cm long, blades 0.3–1 cm wide ................................................ P. granuliferum Culms freely branched at the upper nodes; lower flower absent, lower palea present or absent ......................................................... P. polycomum Culms not conspicuously branched at the upper nodes; lower flower pistillate, lower palea present ................................................................. 56 Plants annual; blades 1.3–2 cm long; panicle 4–6 cm long ........... P. petrense Plants perennial; blades 2–10 cm long; panicle 8–12 cm long ................... .............................................................................................. P. micranthum

Panicum altum Hitchc. & Chase, Contr. U.S. Natl. Herb. 17: 488, fig. 57. 1915. Panicum vigoratum Swallen, Mem. New York Bot. Gard. 9: 257. 1957. Cespitose perennial, 2–3 m tall; ligules membranous-ciliate; blades linear, 25–45 cm long, flat; spikelets 2.8–3.8 mm long, lower glume 3/4 the length of the spikelet. Sand beaches, borders of swamps, river beaches, 50–200 m; Delta Amacuro (mouth of Caño Güiniquina), Amazonas (Río Guaviarito on Cerro Guanay). Belize, Nicaragua, Panama, Colombia, Trinidad-Tobago, Guyana, Suriname, French Guiana, northern Brazil. Panicum arctum Swallen, Bull. Torrey Bot. Club 75: 87. 1948. Slender annual; spikelets obovate or pearshaped, 1–1.2 mm long, with a small stipe between the lower and upper glume. Savannas, ca. 700 m; Bolívar (Cerro Guaiquinima). Guyana. Panicum caricoides Nees in Trin., Gram. Panic. 148. 1826. Panicum stenodoides F.T. Hubb., Proc.

Amer. Acad. Arts 49: 497. 1913. Tufted perennial, culms 10–30 cm tall; blades linear-lanceolate; panicles contracted, 0.5–1.2 cm long, 2–10-flowered; spikelets with the lower glume separated from the upper glume by a conspicuous internode. Savannas in wet areas, streambanks, sandy soils, 300–1300 m; Bolívar (Gran Sabana), Amazonas (Cerro Duida, Río Cunucunuma, Simarawochi on Sierra Parima). Anzoátegui, Apure; Mexico, Belize, Panama, Colombia, Guyana, Suriname, French Guiana, Brazil, Bolivia. ◆Fig. 131. Panicum cayennense Lam., Tabl. Encycl. 1: 173. 1791. Cespitose annual; panicles several from the upper nodes and forming a compound inflorescence; spikelets obovate, 2.1–2.6 mm long, upper floret shortly stipitate. Open sandy areas in savannas, 50–200 m; Delta Amacuro (Isla Tórtola), Amazonas (near Puerto Ayacucho, Santa Bárbara del Orinoco). Anzoátegui, Apure, Barinas, Guárico, Monagas; Mexico, Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Brazil, Bolivia.

184

P OACEAE

Panicum cervicatum Chase, J. Wash. Acad. Sci. 32: 164, fig. 10. 1942. Panicum vinaceum Swallen, Fieldiana, Bot. 28: 27. 1951. Cespitose perennial, culms 40–75 cm tall; spikelets gaping, 5.5–9 mm long, obliquely set on pedicels, upper glume and lower lemma 7–11-veined, upper floret stipitate. Open savannas, near sea level to 800 m; Bolívar (southern Gran Sabana), Amazonas (Isla Ratón, near Puerto Ayacucho). Brazil, Bolivia, Paraguay. ◆Fig. 133. Panicum chnoodes Trin., Gram. Panic. 211. 1826. Panicum curvifolium Swallen, Bull. Torrey Bot. Club 58: 316. 1931. Panicum tatei Swallen, Bull. Torrey Bot. Club 58: 312, fig. 2. 1931. Panicum tropidoblephare Tutin, J. Bot. 72: 339, fig. 9. 1934. Panicum cowanii Swallen, Mem. New York Bot. Gard. 9: 262. 1957. Panicum inversum Swallen, Mem. New York Bot. Gard. 9: 262. 1957. Panicum kavanayense Swallen, Mem. New York Bot. Gard. 9: 264. 1957. Panicum maguirei Swallen, Mem. New York Bot. Gard. 9: 264. 1957. Panicum vannum Swallen, Mem. New York Bot. Gard. 9: 261. 1957. Panicum churunense Swallen, Acta Bot. Venez. 2(5–8): 133. 1967. Cespitose perennial 0.3–1.1 m tall; ligules absent; blades linear, 15–30 cm long, pungent; spikelets elliptic, glabrous (1.7–)2.2–2.8 mm long. Savannas, rocky mountainous areas, on sandy soils, 700–1800 m; widespread at high altitudes in Bolívar and Amazonas. Guyana, Brazil (Goiás, Minas Gerais). ◆Fig. 136. Panicum cordovense E. Fourn., Mex. Pl. 2: 26. 1886. Decumbent perennial, culms scandent on neighboring vegetation; blades lanceolate, symmetric at base; spikelets pilose to glabrous, 3.3–4.5 mm long; lower palea absent. Wet forest margins, ca. 800 m; Bolívar (Cerro Ceitá north of Santa Elena de Uairén). Falcón; Mexico, Central America, Colombia, Guyana, Suriname, French Guiana, Brazil, Bolivia, northern Argentina. ◆Fig. 137. Panicum cyanescens Nees ex Trin., Gram. Panic. 202. 1826. Panicum savannarum Soderstr., Mem.

New York Bot. Gard. 12(3): 2. 1965. Cespitose perennial, with or without scaly rhizomes; culms erect, 30–85 cm tall; spikelets 1.1–1.8 mm long, obovate, glabrous to rarely hirsute; upper glume and lower lemma 5-veined. Savannas, sandy, dry or wet soils, 100–2000 m; widespread in Bolívar and Amazonas. Anzoátegui, Apure, Aragua, Guárico, Zulia; Mexico, Central America, Colombia, Trinidad-Tobago, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia. ◆Fig. 139. Panicum deciduum Swallen, Mem. New York Bot. Gard. 9: 257. 1957. Cespitose perennial with freely branching culms to 4 m tall; spikelets pilose, upper glume and lower lemma 5-veined; upper floret stipitate. Tepui slopes, 800–1000 m; Amazonas (Cerro Yapacana). Endemic. ◆Fig. 142. Panicum dichotomiflorum Michx., Fl. Bor.-Amer. 1: 48. 1803. Annual, culms decumbent, rooting and branching at the lower nodes; spikelets ovate, 2.4–3 mm long, lower glume 1/4–1/3 the length of the spikelet, upper glume and lower lemma 7–9-veined. River banks, in sandy, humid soils, also in disturbed places, ca. 50 m; Delta Amacuro (Caño Araguaito). Anzoátegui, Apure, Guárico; widespread from Mexico to Argentina. ◆Fig. 145. Panicum discrepans Döll in Mart., Fl. Bras. 2(2): 252. 1877. Tufted or decumbent perennial, culms 20– 40 cm tall; ligule ciliate; spikelets 1.1–1.3 mm long, plano-convex, lower glume absent to minute, upper glume and lower lemma 3veined; upper floret gibbous, bearded at apex and base. Edges of seasonal ponds in lowland savannas, ca. 100 m; Bolívar (Maripa-Aripao area), Amazonas (Caño Caname, near Puerto Ayacucho). Apure, Monagas; Belize, Costa Rica, Cuba, Guyana, Suriname, French Guiana, Brazil, Bolivia. ◆Fig. 135. Panicum elephantipes Nees ex Trin., Gram. Panic. 206. 1826. Panicum fistulosum Hochst. ex Steud., Syn. Pl. Glumac. 1: 71. 1855 [1853]. Robust aquatic, culms succulent, 1–1.5 m long; spikelets lanceolate, 3.4–5.6 mm long, lower glume 1/6 the length of the spikelet. Usually forming floating mats in rivers, near sea level to 200 m; Delta Amacuro (Caño Acoima, mouth of Caño Güiniquina, Caño

Panicum 185

Macareo, Isla El Toro), Bolívar (south of El Dorado), Amazonas (Guachapito, lower Río Casiquiare). Apure, Barinas, Guárico, Zulia; Mexico, Central America, West Indies, south to Argentina. ◆Fig. 144. Panicum eligulatum N.E. Br., Trans. Linn. Soc. Bot. 6: 73. 1901. Panicum auyanense Swallen, Mem. New York Bot. Gard. 9: 263. 1957. Cespitose perennial; ligules absent; blades linear-lanceolate, 15–35 cm long, pungent; spikelets elliptic, (2.8–)3.1–3.7 mm long, glabrous. Tepui-summit savannas, 1100–2500 m; Bolívar (Auyán-tepui, Macizo del Chimantá, Roraima-tepui), Amazonas (Cerro Sipapo). Guyana. Panicum fontanale Swallen, Phytologia 14: 80. 1966. Culms decumbent and rooting at the lower nodes, 40–50 cm tall; sheaths auriculate; ligules absent; blades pungent; spikelets narrowly elliptic, 3.1–3.5 mm long, glabrous, upper glume and lower lemma 5-veined. Along waterfalls, ca. 1700 m; Bolívar (Macizo del Chimantá [Chimantá-tepui]). Endemic. Panicum fonticolum Swallen, Mem. New York Bot. Gard. 9: 260. 1957. Annual, culms decumbent, rooting at the lower nodes; ligules membranous, small, 0.4 mm long; blades lanceolate, narrowed at base; panicles 3.5–6 cm long; spikelets elliptic, 1.4–1.6 mm long, lower palea absent. Spray of waterfalls, 1500–1600 m; Amazonas (Caño Culebra on Cerro Duida). Endemic. Panicum grande Hitchc. & Chase, Contr. U.S. Natl. Herb. 17: 529, fig. 143. 1915. Robust aquatic perennial 1–3 m tall; blades 50–70 cm long; panicles many-flowered, 30–70 cm long; spikelets lanceolate, 2.3–2.8 mm long, upper glume and lower lemma 5-veined. Lowlands, coastal marshes, streams, near sea level to 300 m; Delta Amacuro (Caño Araguao, mouth of Río Cuyubini), Bolívar (La Vergareña). Widespread elsewhere in Venezuela; Mexico, Guatemala, Honduras, Costa Rica, Panama, Colombia, Guyana, French Guiana, Suriname, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 146. Panicum granuliferum Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 105. 1815 [1816]. Panicum spissifolium Swallen, Bull.

Torrey Bot. Club 75: 86. 1948. Panicum subcordatum Swallen, Mem. New York Bot. Gard. 9: 258. 1957. Panicum subinclusum Swallen, Mem. New York Bot. Gard. 9: 260. 1957. Panicum politii Swallen, Phytologia 14: 76. 1966. Perennial with decumbent culms, freely branching at the lower nodes; blades lanceolate, 3–6 mm wide; spikelets globose, 1– 1.1 mm long, hispid to more commonly glabrous, upper glume and lower lemma 5veined. Open places or forest edges on white sand, 50–200 m; Bolívar (Cerro Camarón south of Entrerios, Cerro Guaiquinima, Guayaraca, Río Parguaza, near Urimán), Amazonas (widespread). Colombia, Guyana, French Guiana, northern Brazil, Bolivia. ◆Fig. 141. Panicum haenkeanum J. Presl in C. Presl, Reliq. Haenk. 1: 304. 1830. Perennial with decumbent, rooting and branching culms; blades lanceolate 6–12 cm long; spikelets narrowly elliptic, ca. 2.3 mm long, upper glume and lower lemma 5veined. Forest edges, humid, shady places, 100–300 m; Bolívar (La Vergareña on Cerro Moro, Río Aro between Caicara and Ciudad Bolívar). Apure, Carabobo, Portuguesa, Sucre; Mexico, Belize, Honduras, Nicaragua, Costa Rica, Panama, Peru, Brazil, Bolivia. ◆Fig. 147. Panicum hirsutum Sw., Fl. Ind. Occid. 1: 173, fig. 24. 1797. Robust cespitose perennial, culms to 3 m tall; sheaths papillose-pilose with glass-like hairs; ligules membranous-ciliate, 0.2–0.7 mm long; blades lanceolate, 30–50 cm long; spikelets narrowly ovate, 1.8–2.6 mm long. River banks, near sea level to 50 m; Delta Amacuro (Isla Cocuina). Barinas, Falcón, Portuguesa, Táchira, Trujillo, Yaracuy, Zulia; widespread in southern U.S.A., Mexico, Central America, and West Indies, less abundant in South America in Colombia, Guyana, Ecuador, Peru, Brazil, Paraguay, Argentina. ◆Fig. 150. Panicum hirtum Lam., Encycl. 4: 741. 1797 [1798]. Annual, branching and rooting at the lower nodes; panicles open, axis of the branches with glands; spikelets papillose-pilose, 1.2–1.5 mm long, obliquely set on pedicels. Forest and savanna edges, wet

186

P OACEAE

places, ca. 50–500 m; Bolívar (Río Nichare, Río Paramichi at Río Paragua, San Félix), Amazonas (near Puerto Ayacucho, Río Mawarinuma). Anzoátegui, Barinas, Carabobo, Cojedes, Guárico, Lara, Portuguesa, Sucre; southern Mexico, Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Brazil. ◆Fig. 152. Panicum hispidifolium Swallen, Contr. U.S. Natl. Herb. 29: 424. 1950. Annual, culms erect; ligules membranousciliate, 2–3 mm long; blades lanceolate, 10– 45 cm long, papillose-pilose; spikelets elliptic, 3.2–3.7 mm long, upper glume and lower lemma 7–9-veined. Savannas, ca. 100 m; Bolívar (Río Chaviripa). Anzoátegui, Apure, Aragua, Calabozo, Cojedes, Guárico, Monagas, Portuguesa, Trujillo; Mexico, Central America, Colombia. ◆Fig. 143. Panicum hylaeicum Mez, Notizbl. Bot. Gart. Berlin-Dahlem 7: 75. 1917. Panicum guianense Hitchc., Contr. U.S. Natl. Herb. 22(6): 487, fig. 83. 1922. Usually scandent, robust perennial to 3 m; blades cordate and stem-clasping basally, 1–3 cm wide; spikelets 1.4–1.7 mm long, secund, on short branches which are appressed to first-order branches, pilose or glabrous, upper glume and lower lemma 5-veined. River banks, ca. 100 m; Amazonas (Puerto Ayacucho). Apure, Barinas, Guárico, Zulia; widespread from Mexico and the West Indies to Paraguay and Argentina. ◆Fig. 153. Panicum ichunense Swallen, Phytologia 14: 77. 1966. Annual, culms decumbent and rooting at the lower nodes; blades elliptic, 2.8–4.5 × 4–7 mm; spikelets narrowly pear-shaped to obovate, ca. 2.3 mm long, upper glume and lower lemma 5-veined, upper floret stipitate. Wet habitats near waterfalls, 500–2000 m; Bolívar (Cerro Ichún, Macizo del Chimantá [Chimantá-tepui]). Endemic. Panicum jauanum Davidse, Bol. Soc. Venez. Ci. Nat. 132: 272. 1976. Cespitose perennial ca. 1 m tall; ligules membranous-ciliate, 0.7–1 mm long; blades lanceolate 28–40 cm long, flat, pungent; spikelets elliptic, ca. 3.2 mm long, glabrous, upper glume and lower lemma 5-veined. Tepui savannas, 1700–1800 m; Bolívar (Cerro Jaua). Endemic. ◆Fig. 155.

Panicum ligulare Nees ex Trin., Gram. Panic. 206. 1826. Cespitose perennial 1.3–2 m tall; panicles lax; spikelets elliptic, 4.4–5.7 mm long, upper glume and lower lemma 7–9-veined, upper floret stipitate. Savannas, ca. 400 m; Bolívar (upper Río Suapure). Central Brazil. Panicum maximum Jacq., Icon. Pl. Rar. 1: 2, t. 13. 1781. —Urochloa maxima (Jacq.) R.D. Webster, Austral. Paniceae 241. 1987. Robust cespitose perennial 1–3 m tall; panicles lax, open, the lower branches whorled; spikelets narrowly elliptic, 3–3.8 mm long, upper glume and lower lemma 5veined, upper floret conspicuously rugose. Roadsides, disturbed places, near sea level to 500 m; scattered in Delta Amacuro, Bolívar, and Amazonas. Widespread throughout the Neotropics, introduced from Africa. ◆Fig. 140. Panicum mertensii Roth in Roem. & Schult., Syst. Veg. 2: 458. 1817. Perennial 1–3 m tall; panicles open with all branches whorled; spikelets obovate, 3.2– 4 mm long, glabrous, upper floret smooth. Common in water, near sea level to 200 m; Delta Amacuro (Caño Joba-Suburu, Isla Cocuina, Isla Guara), Amazonas (Río Orinoco below Caño Yapacana). Barinas, Monagas, Portuguesa; widely distributed in Mexico, Guatemala, Honduras, El Salvador, Costa Rica, Panama, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina. ◆Fig. 157. Panicum micranthum Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 105. 1815 [1816]. Panicum micranthum var. hirtum Maury, J. Bot. (Morot) 3: 162. 1889. Panicum densifolium Swallen, Fieldiana, Bot. 28: 25. 1951. Panicum mauryi Swallen, Fieldiana, Bot. 28: 26. 1951. Panicum supernum Swallen, Mem. New York Bot. Gard. 9: 256. 1957. Cespitose perennial, culms geniculate to erect, 12–40 cm tall, branching at the upper nodes; blades linear-lanceolate, to 3 mm wide; spikelets obovate, 0.9–1.2 mm long, pilose to glabrous, upper glume and lower lemma 5-veined, upper floret smooth, pilose. Sandy savannas, 100–1400 m; widespread in

Panicum 187

Bolívar and Amazonas (except basins of Río Casiquiare and Río Negro). Anzoátegui, Apure, Guárico, Monagas; Colombia, Guyana, Suriname, French Guiana, northern Brazil. ◆Fig. 138. Panicum millegrana Poir. in Lam., Encycl. suppl. 4, 278. 1816. Scandent perennial to 1.5 m tall, culms decumbent and rooting at the lower nodes; panicles lax, many-flowered, 10–40 cm long; spikelets obovate, 1.6–2.5 mm long, hirsute to glabrous, upper glume and lower lemma 5veined, upper floret slightly rugose. Humid places on forest edges, 200–1400 m; scattered in northern Bolívar. Mexico, Guatemala, Belize, Honduras, Panama, Colombia, Guyana, Suriname, French Guiana, Ecuador, Brazil, Bolivia, Paraguay, Argentina, Uruguay. ◆Fig. 134. Panicum nervosum Lam., Encycl. 4: 747. 1797 [1798]. Cespitose perennial 0.4–2 m tall; leaves strongly overlapping, 0.8–1.5 cm wide; spikelets obovate, 1.1–1.7 mm long, glabrous. Sandy savannas, 100–1300 m; Bolívar (Gran Sabana, Río Chicanán), Amazonas (scattered). Colombia, Trinidad-Tobago, Guyana, Suriname, French Guiana, northern Brazil. Panicum olyroides Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 102. 1815 [1816]. Panicum funckianum Steud., Syn. Pl. Glumac. 1: 77. 1855 [1853]. Cespitose perennial 0.5–1 m tall; panicles open, falling entire when mature; spikelets 6–8.5 mm long, upper floret with long hairs at the base of the lemma. Open, dry savannas, 100–1900 m; widely scattered in Bolívar. Anzoátegui, Aragua, Carabobo, Distrito Federal, Mérida, Miranda, Monagas, Sucre, Trujillo, Zulia; Colombia, French Guiana, Suriname, Brazil, Bolivia, Paraguay. ◆Fig. 158. Panicum orinocanum Luces, J. Wash. Acad. Sci. 32: 164, fig. 9. 1942. Cespitose, short-rhizomatous perennial 30–40 cm tall; blades linear-lanceolate, 3–4 × 0.1–0.2 cm; spikelets obovate, 1.4–1.7 mm long, pilose or glabrous, upper glume and lower lemma 5-veined. White-sand savannas, 100–200 m; common in western Amazonas. Apure; Colombia. ◆Fig. 151.

Panicum pandum Swallen, Brittonia 3: 150. 1939. Panicum graniticum Swallen, Phytologia 14: 71. 1966. Annual, culms 5–20 cm tall, decumbent to erect; blades subcordate to cordate, 0.2– 0.6(–1) cm wide; spikelets sparsely to densely pilose, 1.1–1.9 mm long, upper glume and lower lemma 5-veined, lower palea absent. Exposed rocks in wet places, 600–2200 m; Bolívar (Amaruay-tepui, Auyán-tepui, Cerro Guaiquinima, Cerro Venado, Macizo del Chimantá, Río Caruay), Amazonas (Cerro Huachamacari, Cerro Yutajé). Guyana. Panicum pantrichum Hack., Verh. K.K. Zool.-Bot. Ges. Wien 65: 72. 1915. Perennial, decumbent and rooting at the lower nodes; ligules membranous; spikelets elliptic 2.1–2.8 mm long, hispid to glabrous, usually with cleistogamous flowers, upper glume and lower lemma 5-veined. Base of tepui escarpments, ca. 2400 m; Bolívar (Ptari-tepui). Aragua, Distrito Federal, Lara, Mérida, Portuguesa; Honduras, Costa Rica, Panama, Colombia, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina. Panicum parvifolium Lam., Tabl. Encycl. 1: 173. 1791. Panicum kaietukense Tutin, J. Bot. 72: 340, fig. 11. 1934. Straggling, freely branching, creeping and rooting at the lower nodes; blades 0.7–4 cm long; spikelets globose, 1–1.8 mm long, upper glume and lower lemma 5-veined. Forming large colonies at edges of wet places, near sea level to 1100 m; Delta Amacuro (Caño Güiniquina), Bolívar (widespread), Amazonas (Río Atabapo, Río Orinoco). Anzoátegui, Apure, Guárico; widespread in Central America, West Indies, and South America, introduced in Africa. Panicum petrense Swallen, Mem. New York Bot. Gard. 9: 259. 1957. Cespitose annual, culms erect, 13–30 cm tall; blades lanceolate, 1.3–2 cm; panicles 4–6 cm long; spikelets obovate to elliptic, hispid, 0.9–1.3 mm long, upper glume and lower lemma 5-veined. Moist sand on or near granitic outcrops, 50–100 m; Amazonas (San Fernando de Atabapo north to El Burro). Endemic.

188

P OACEAE

Panicum pilosum Sw., Prodr. 22. 1788. Panicum pilisparsum G. Mey., Prim. Fl. Esseq. 57. 1818. Panicum distichum var. lancifolium Griseb., Fl. Brit. W. I. 548. 1864. —Panicum distichum var. lancifolium Griseb. ex Hitchc., Man. Gr. West Indies 267. 1936, nom illeg. —Panicum pilosum var. lancifolium (Griseb.) R. Pohl, Fieldiana, Bot. n.s. 4: 381. 1980. Panicum coenosum Döll in Mart., Fl. Bras. 2(2): 191, fig. 30. 1877. Panicum milleflorum Hitchc. & Chase, Contr. U.S. Natl. Herb. 17: 494, fig. 70. 1915. Stoloniferous perennial, culms 10–70 cm tall; ligule usually absent; axis of the panicle pilose; spikelets unilaterally disposed on first-order branches, biconvex, 1.2–1.5 mm long, upper glume and lower lemma 5veined. Usually found along forest margins or in disturbed places, 50–300 m; widespread in Delta Amacuro, Bolívar, and Amazonas. Apure, Aragua, Barinas, Falcón, Guárico, Lara, Miranda, Portuguesa, Sucre, Táchira, Zulia; Mexico, Central America, West Indies, most South American countries. Panicum polycomum Trin., Mém. Acad. Imp. Sci. St.-Pétersbourg, Sér. 6, Sci. Math., Seconde Pt. Sci. Nat. 1: 306. 1834. Panicum siccaneum Trin., Linnaea 10: 298. 1836. Panicum obovatum Döll in Mart., Fl. Bras. 2(2): 256. 1877. Panicum tamayonis Luces, Bol. Soc. Venez. Ci. Nat. 15: 24, fig. 17. 1953. Panicum gracillissimum Swallen, Mem. New York Bot. Gard. 9: 259. 1957. Annual 3–15(–30) cm tall, culms geniculate to erect, freely branching; blades linearlanceolate, 1–2 mm wide; spikelets obovate, 0.9–1.5 mm long, hirsute to glabrous, upper glume and lower lemma 5-veined, lower palea present or absent. Common in savannas in sandy soils, ca. 100–1300 m; widespread in Bolívar and Amazonas. Guárico; eastern Colombia, French Guiana, northern Brazil. ◆Fig. 149. Panicum polygonatum Schrad. in Schult., Mantissa 2: 256. 1824. Perennial, culms decumbent and rooting at the lower nodes, 15–50 cm tall; blades sub-

cordate, shortly pseudopetiolate; spikelets narrowly elliptic, 1.3–1.6 mm long, pointed at the apex, unilaterally disposed on the branches of the panicle. Forest margins, swamps, 100–200 m; Bolívar (El Dorado), Amazonas (Río Mawarinuma at base of Sierra de la Neblina). Apure, Aragua, Distrito Federal, Falcón, Lara, Mérida, Miranda, Monagas, Portuguesa, Táchira, Zulia; Mexico, Central America, West Indies, most South American countries. ◆Fig. 154. Panicum pulchellum Raddi, Agrostogr. Bras. 42. 1823. Culms decumbent, creeping and rooting at lower nodes, 10–65 cm tall; spikelets elliptic, 1.8–2.3 mm long, lower lemma bearing 2 crateriform glands, upper floret smooth, glabrous. Humid, shaded places, ca. 100 m; Amazonas (Isla Carestía in Río Orinoco). Anzoátegui, Apure, Aragua, Barinas, Distrito Federal, Lara, Mérida, Miranda, Portuguesa, Sucre, Yaracuy; Mexico, Central America, West Indies, Colombia, Ecuador, Peru, Brazil, Bolivia. Panicum pyrularium Hitchc. & Chase, Contr. U.S. Natl. Herb. 17: 508. 1915. Panicum quetameense Mez, Notizbl. Bot. Gart. Berlin-Dahlem 7: 75. 1917. Straggling annual, erect portion of culms 5–30 cm tall; spikelets pear-shaped, 1.2–1.8 mm long, hispid to glabrous, with a conspicuous internode between lower and upper glume, upper glume and lower lemma 5veined. Moist soil, wet river banks, rocks, sandstone or granitic outcrops, 50–1000 m; scattered in Bolívar and Amazonas. Aragua, Carabobo, Guárico, Portuguesa, Táchira; Panama, Colombia, Guyana, Suriname, French Guiana, Brazil (Pará), Bolivia. ◆Fig. 156. Panicum rivale Swallen, Bull. Torrey Bot. Club 75: 87. 1948. Panicum angulosum Swallen, Phytologia 14: 75. 1966. Decumbent annual, 5–25 cm tall; blades cordate to subcordate basally, 2–6 mm wide; spikelets hispid, slightly pear-shaped, 1.1– 1.6 mm long, upper glume and lower lemma 5-veined, lower palea usually absent. Savannas, 100–300 m; Bolívar (km 100 on El Dorado to Santa Elena road). Guyana, Suriname, French Guiana.

Panicum 189

Panicum rudgei Roem. & Schult., Syst. Veg. 2: 444. 1817. Cespitose perennial 30–130 cm tall; panicles forming a compound inflorescence 25–50 cm long; spikelets ovate, 3–3.5 mm long, upper glume and lower lemma 7–9veined, upper floret stipitate. Open savannas, 100–1200 m; Bolívar (Gran Sabana), Amazonas (widespread). Anzoátegui, Apure, Barinas, Monagas, Zulia; Mexico, Central America, West Indies, eastern Colombia, Suriname, French Guiana, Peru, Brazil, Bolivia. Panicum scabridum Döll in Mart., Fl. Bras. 2(2): 201. 1877. Panicum manacalensis Swallen, Phytologia 14: 77. 1966. Tufted perennial 0.5–1.1 m tall; panicles narrowly ovate, 7–20 cm long; spikelets biconvex, pointed, 1.3–1.8 mm long, upper glume and lower lemma 5-veined; upper floret dark; caryopsis black. Wet open places, ca. 50–100 m; Delta Amacuro (Isla Guara, Isla La Tórtola), Amazonas (San Carlos de Río Negro). Apure, Guárico, Monagas; Colombia (Meta), Guyana, Suriname, French Guiana, northern Brazil, Amazonian Bolivia.

Panicum stenodes Griseb., Fl. Brit. W. I. 547. 1864. Densely tufted perennial, culms erect, 20– 45(–75) cm tall; leaves filiform; panicles contracted, with 10–25 spikelets 1.1–1.7 mm long, glabrous, upper glume and lower lemma 5(–7)-veined. Sandy savannas, 50– 1200 m; Bolívar (Gran Sabana, La Vergareña, Río Cuchivero), Amazonas (basin of Caño Guanay, Cerro Duida, near Puerto Ayacucho). Anzoátegui, Apure, Carabobo, Guárico; Mexico, Guatemala, Belize, Honduras, Nicaragua, Costa Rica, Panama, West Indies, Colombia, Guyana, Suriname, French Guiana, Brazil, Bolivia. ◆Fig. 132. Panicum steyermarkii Swallen, Mem. New York Bot. Gard. 9: 402. 1957. Perennial 30–60 cm tall, culms decumbent and branching at the lower nodes; blades linear-lanceolate, 10–21 cm long, pungent; panicles lax, 5–10 cm long; spikelets narrowly elliptic, 3.9–4.4 mm long, glabrous, upper glume and lower lemma 5-veined. Stream sides on tepui summits, 1700–2000 m; Bolívar (Macizo del Chimantá). Endemic. ◆Fig. 159.

Panicum sellowii Nees in Mart. et al., Fl. Bras. Enum. Pl. 2: 153. 1829. Scandent perennial to 3 m long, erect portion of culms to 1 m tall; blades lanceolate, 6– 13 cm long, asymmetric; panicles lax, 6–22 cm long, with spikelets congested on upper portion of branches; spikelets obovate, 1.6– 2.4 mm long, pilose or glabrous, upper glume and lower lemma 5-veined, upper floret slightly rugose transversely. Forest edges, 100–500 m; scattered in northern and central Bolívar. Aragua, Distrito Federal, Falcón, Lara, Miranda, Monagas, Portuguesa, Sucre, Táchira; widespread from Mexico and the West Indies to Argentina.

Panicum stoloniferum Poir. in Lam., Encycl. Suppl. 4: 274. 1816. Stoloniferous perennial, culms 10–60 (–100) cm tall; spikelets disposed unilaterally in racemose branches, lanceolate, 2.2– 3.2 mm long, glabrous; upper floret shortstipitate, indurate, glabrous. Forest edges, common in wet and shady habitats, 100–900 m; Delta Amacuro (scattered), Bolívar (scattered across northern and central portions), Amazonas (Río Casiquiare, near San Fernando de Atabapo, Sierra Parima). Barinas, Lara, Miranda, Monagas, Sucre, Yaracuy, Zulia; Central America, West Indies, south to Argentina. ◆Fig. 161.

Panicum sipapoense Swallen, Mem. New York Bot. Gard. 9: 261. 1957. Perennial, culms decumbent and branching at the lower nodes; ligules membranousciliate, 0.3–0.7 mm long; blades lanceolate, pungent; spikelets elliptic, 2.7–3.1 mm long, glabrous. Stream sides, wet areas on tepui summits, 1400–2000 m; Bolívar (Cerro Guaiquinima), Amazonas (Cerro Sipapo). Endemic.

Panicum tepuianum Davidse & Zuloaga, Novon 1: 191. 1991. Cespitose perennial to 86 cm tall, culms erect; ligules 0.3–0.6 mm long, membranous, strongly arcuate; blades stiff, 18–21 cm long; panicle lax, open, 16–18 cm long; spikelets 1.8–2.4 mm long; upper floret indurate and sparsely pilose. Tepui-summit savannas, ca. 1400 m; Amazonas (Cerro Aracamuni). Endemic.

190

P OACEAE

Fig. 131. Panicum caricoides

Fig. 132. Panicum stenodes

Panicum 191

Fig. 133. Panicum cervicatum Fig. 134. Panicum millegrana

192

P OACEAE

Fig. 135. Panicum discrepans

Fig. 136. Panicum chnoodes

Panicum 193

Fig. 137. Panicum cordovense

Fig. 138. Panicum micranthum

194

P OACEAE

Fig. 139. Panicum cyanescens

Fig. 140. Panicum maximum

Fig. 141. Panicum granuliferum

Panicum 195

Fig. 142. Panicum deciduum

Fig. 143. Panicum hispidifolium

196

P OACEAE

Fig. 144. Panicum elephantipes

Fig. 145. Panicum dichotomiflorum

Panicum 197

Fig. 146. Panicum grande

198

P OACEAE

Fig. 147. Panicum haenkeanum

Fig. 148. Panicum yavitaense

Fig. 149. Panicum polycomum

Panicum 199

Fig. 150. Panicum hirsutum

200

P OACEAE

Fig. 151. Panicum orinocanum

Fig. 152. Panicum hirtum

Panicum 201

Fig. 154. Panicum polygonatum

Fig. 153. Panicum hylaeicum

202

P OACEAE

Fig. 155. Panicum jauanum

Panicum 203

Fig. 156. Panicum pyrularium

Fig. 157. Panicum mertensii

204

P OACEAE

Fig. 158. Panicum olyroides

Panicum 205

Fig. 159. Panicum steyermarkii

Fig. 160. Panicum trichoides

206

P OACEAE

Fig. 161. Panicum stoloniferum

Panicum 207

Fig. 162. Panicum tricholaenoides

208

P OACEAE

Panicum trichoides Sw., Prodr. 24. 1788. Annual, decumbent and rooting at the lower nodes, erect culms 10–60 cm tall; blades ovate-lanceolate, 2–7 cm long, flat, cordate; panicles lax, diffuse, 5–20 cm long; spikelets elliptic to obovate, pilose, 0.9–1.5 mm long, upper glume and lower lemma 3–5-veined. Wet or disturbed areas, 400–1100 m; Bolívar (Cerro Venamo, Santa María del Vapor). Widespread from U.S.A. to Argentina. ◆Fig. 160. Panicum tricholaenoides Steud., Syn. Pl. Glumac. 1: 68. 1855 [1853]. Rhizomatous perennial 1.5–2 m tall, culms bamboo-like, rigid; panicles open, 20– 40 cm long; spikelets gaping, 2.2–2.9 mm

long, upper glume and lower lemma 7–9veined. Sandy soils along rivers, 100–200 m; Amazonas (Caño Guanay, near Puerto Ayacucho, San Juan de Manapiare). Apure; eastern Colombia, Brazil, Bolivia, Paraguay, Argentina, Uruguay. ◆Fig. 162. Panicum yavitaense Swallen, Phytologia 14: 72. 1966. Annual, culms 7–20 cm tall; blades lanceolate, 20–50 × 2–7 mm, subcordate; panicles 6–12 cm long, lax; spikelets obovate, hispid, 1.2–1.4 mm long. Savannas, 100–300 m; Amazonas (Río Yatúa, base of Sierra de la Neblina, near Yavita). Colombia (Vaupés). ◆Fig. 148.

63. PAPPOPHORUM Schreb., Gen. Pl. 2: 787. 1791. [Subfamily Chloridoideae, Tribe Pappophoreae] by Emmet J. Judziewicz Coarse perennials. Leaves with ligule a line of hairs, the blades elongate, flat to folded. Inflorescence a contracted panicle. Spikelets 3–6-flowered, the florets disarticulating as a unit from the persistent glumes; glumes unequal to subequal, membranous, 1-veined, somewhat laterally compressed, shorter than the spikelet; lower 1–3 florets bisexual, the lemmas indurate, rounded on back, 11–15-awned, 5– 11-veined, the veins sometimes villous; uppermost 2 or 3 florets reduced in size, sterile, but the lemmas with 13–20 prominent awns. Southwestern U.S.A., Mexico, Central America, West Indies, tropical and warm-temperate South America; ca. 10 species, 2 species in Venezuela, 1 of these in the flora area. Pappophorum mucronulatum Nees in Mart., Fl. Bras. Enum. Pl. 2: 412. 1829. Coarse, cespitose perennial 0.5–1 m tall. Inflorescence 10–30 × 1–2 cm, a contracted panicle. Weedy places, 200–400 m; Bolívar (13 km east-southeast of Upata). Anzoátegui,

Carabobo, Distrito Federal, Lara, Portuguesa; Honduras, Colombia, Brazil. Pappophorum pappiferum (Lam.) Kuntze, a more robust species 1–2 m tall with larger, bushier panicles 30–50 cm long, may eventually be found in the flora area. This is the species that is illustrated. ◆Fig. 163.

64. PARATHERIA Griseb., Cat. Pl. Cub. 236. 1866. [Subfamily Panicoideae, Tribe Paniceae] by Emmet J. Judziewicz Plants of indefinite duration, perhaps annual, often bearing cleistogamous spikelets near the base. Main panicle terminal, spicate, sparingly flowered, the spikelets borne on short, erect branches; branches deciduous, callus-like and pungent at the base, laterally bearing a persistent, subsessile spikelet and prolonged past the spikelet as a hispid, flattened, sterile bristle. Spikelets lanceolate, acuminate-attenuate, somewhat dorsally compressed, 2-flowered; glumes subequal, minute, scale-like, translucent, veinless; lower floret sterile, the lemma as long as the

Paratheria 209

Fig. 163. Pappophorum pappiferum [expected in the flora area]

Fig. 164. Paratheria prostrata

210

P OACEAE

spikelet, several-veined, membranous, the palea absent; upper floret as long as spikelet, the lemma stiffly membranous, several-veined, its margins covering the edges of the palea. Neotropics, tropical Africa; 2 species, 1 in Venezuela. Paratheria prostrata Griseb., Cat. Pl. Cub. 236. 1866. Tufted plant, the culms decumbent and rooting, forming mats, the erect portions to 40 cm tall; leaf blades 5–10 cm × 3–7 mm; terminal inflorescences 5–12 cm long; branches bearing spikelets 2–3 cm long;

spikelets 7–10 mm long. Sand bars in rivers, ca. 100 m; Amazonas (Caño Caname, between Caño Cotuá and Cerro Yapacana). Apure, Guárico, Monagas; Costa Rica, Dominican Republic, Colombia, Guyana, French Guiana, Bolivia, Brazil, tropical Africa, Madagascar. ◆Fig. 164.

65. PARIANA Aubl., Hist. Pl. Guiane 876. 1775. [Subfamily Bambusoideae, Tribe Olyreae] by Emmet J. Judziewicz Monoecious, cespitose, usually rhizomatous or stoloniferous perennials. Culms erect to decumbent, monomorphic with the inflorescence terminating a leafy shoot, or dimorphic and with the inflorescence borne on a separate shoot with rudimentary leaves. Leaves with sheath summits often with prominent bristles (oral setae) and oblong, translucent areas (lunar marks); ligules membranous, usually short; pseudopetioles short; blades linear-lanceolate to ovate. Inflorescence terminal, spicate, of numerous deciduous fascicles; fascicles comprising a verticil of 4–6 pedicellate staminate spikelets surrounding and concealing a single sessile pistillate spikelet, together with the attached rachis internode. Spikelets 1-flowered. Pistillate spikelets with glumes equal, membranous, 1-veined; floret slightly shorter than the glumes, ovate, strongly indurate, faintly 3-veined, glabrous, straw-colored; stigmas 2, plumose. Staminate spikelet pedicels broad, corky, indurate, strongly flattened, shorter, equaling, or longer than the floret, the pedicels of adjacent spikelets often connate; spikelets with glumes equal, narrowly to broadly triangular, (1)2- or 3(4)-veined, erect; floret ovate to narrowly elliptic, acute to obtuse, dorsally compressed, firmly membranous, 3(–5)-veined; palea similar to lemma; stamens 6– 40; filaments connate below; anthers often showy and yellow. Costa Rica, Panama, Colombia, Venezuela, Trinidad-Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 5–30 species, 1 in Venezuela. Pariana grows in shaded, wet, lowland forests, where the numerous, often showy, yellow anthers are adaptations for insect pollination. Pariana campestris Aubl., an Amazonian species with inflorescences borne on the leafy shoots, may eventually be found in the flora area. Pariana radiciflora Sagot ex Döll in Mart., Fl. Bras. 2(2): 336. 1877. Pariana zingiberina Rich. ex Döll in Mart., Fl. Bras. 2(2): 337. 1877. Pariana vulgaris Tutin, J. Linn. Soc. Bot. 50: 353, fig. 17. 1936. Pariana trichosticha Tutin, J. Linn. Soc., Bot. 50: 356, pl. 9, fig. 20. 1936. Pariana obtusa Swallen, Mem. New York Bot. Gard. 9: 268. 1957.

Pariana pallida Swallen, Mem. New York Bot. Gard. 9: 268. 1957. Pariana violascens Swallen, Mem. New York Bot. Gard. 9: 267. 1957. Cespitose perennial 0.3–0.7(–1) m tall; culms dimorphic, the inflorescence borne on a weak, reduced-leafy shoot; leaves with oral setae usually prominent, the blades 7–20 (–28) × 2.5–5.8(–10) cm, lanceolate to ovate; inflorescence 3–8 cm long; staminate pedicels

Pariana 211

Fig. 165. Pariana radiciflora

212

P OACEAE

2–4 mm long, the floret 3–5.5 mm long. Gallery forests, 50–300 m; Amazonas (widespread). Apure, Aragua, Carabobo, Distrito Federal, Miranda, Yaracuy; Panama, Colombia, Trinidad-Tobago, Guyana, Suriname,

French Guiana, Ecuador, Peru, Amazonian Brazil, Bolivia. ◆Fig. 165. Pariana radiciflora is an exceedingly variable species. It is treated here in a broad sense.

66. PARODIOLYRA Soderstr. & Zuloaga, Smithsonian Contr. Bot. 69: 64. 1989. [Subfamily Bambusoideae, Tribe Olyreae] by Emmet J. Judziewicz Monoecious perennials. Culms scandent, arching and clambering, profusely branching from the middle and upper nodes. Leaves with ligules small, membranous-ciliate; pseudopetioles short; blades lanceolate to ovate, flat, stiff, usually asymmetrical, truncate at the base, acuminate and apiculate at the apex. Inflorescences borne from the uppermost nodes; panicles lax and diffuse, the lower branches with staminate spikelets only, the upper ones with staminate spikelets below and pistillate spikelets above, or with pistillate spikelets only; pistillate spikelet pedicels filiform. Spikelets 1-flowered, falling entire, the pistillate spikelets much larger than the staminate ones. Pistillate spikelets elliptic; glumes subequal, as long as spikelet, membranous to indurate, often somewhat inflated at maturity; rachilla internode between glumes prominent and thickened, pulviniform; lower glume 5–9-veined, embracing the 3–6-veined upper glume; floret short-stipitate, indurate, shorter than the glumes, smooth and shiny, acute to obtuse, with or without bottle-like microhairs at the apex, the lemma 5–veined; stigmas 2. Staminate spikelets hyaline, early deciduous, lanceolate, acuminate; stamens 3. Caryopsis elliptic, the linear hilum 1/2–3/4 the length of the caryopsis. Costa Rica, Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 3 species, 2 in Venezuela, both in the flora area. Recently, the widespread Olyra micrantha Kunth has been transferred to the genus and recognized as Parodiolyra micrantha (Kunth) Davidse & Zuloaga (Novon 9: 590. 1999). Key to the Species of Parodiolyra 1.

1.

Mature pistillate spikelets 2–2.5 mm long, the glumes blackish and indurate, the floret obtuse, not viscid; leaf blades 2–7 × 0.5–1.5 cm ..................................................................................................... P. lateralis Mature pistillate spikelets 3–3.5 mm long, the glumes light-colored and papery, the floret acute, viscid; leaf blades 6–12 × 2–3 cm .............................................................................................. P. luetzelburgii

Parodiolyra lateralis (Nees) Soderstr. & Zuloaga, Smithsonian Contr. Bot. 69: 66. 1989. —Panicum laterale C. Presl. ex Nees in Mart. et al., Fl. Bras. Enum. Pl. 2: 213. 1829. —Olyra lateralis (Nees) Chase, Proc. Biol. Soc. Wash. 21: 179. 1908. —Wanak.

Olyra sarmentosa Döll in Mart., Fl. Bras. 2(2): 319. 1877. Scandent or trailing herb with stiff, wiry culms to 3 m long. Savannas, forest edges, bluffs, stream sides, lower slopes of tepuis, 600–1800 m; Bolívar (widespread), Amazonas (Río Ocamo, Sierra Parima). Distrito

Parodiolyra 213

Fig. 166. Parodiolyra luetzelburgii

Fig. 167. Parodiolyra lateralis

214

P OACEAE

Federal, Zulia; Costa Rica, Panama, Colombia, Guyana, Suriname, northwestern Ecuador, Peru, Brazil, Bolivia. ◆Fig. 167. Parodiolyra luetzelburgii (Pilg.) Soderstr. & Zuloaga, Smithsonian Contr. Bot. 69: 70. 1989. —Olyra luetzelburgii Pilg., Notizbl. Bot. Gart. Berlin-Dahlem 10: 1049. 1930.

Cespitose, clambering perennial 0.3–1 m tall. Gallery forests, dry forested slopes, edges of Mauritia palm swamps and granitic outcrops, 100–1300 m; Bolívar (225 km south of Caicara, Gran Sabana, Macizo del Chimantá, Sierra Auraima), Amazonas (Cerro Aratitiyope, Río Orinoco, Río Siapa, Río Ventuari). Táchira; Colombia, Guyana, Suriname, French Guiana, Amazonian Brazil. ◆Fig. 166.

67. PASPALIDIUM Stapf in Prain, Fl. Trop. Afr. 9: 582. 1920. [Subfamily Panicoideae, Tribe Paniceae] by Emmet J. Judziewicz Terrestrial to semiaquatic annuals or perennials, often with decumbent culms that root at the nodes. Leaves with ligules membranous-ciliate; blades linear. Inflorescences paniculiform, terminal and axillary, the panicles elongate, narrow, of many erect to slightly divergent, short, densely flowered racemes, the lowest racemes remote, the racemes secund, the rachis triquetrous, bearing paired, shortpedicellate spikelets from 2 of the 3 sides, prolonged into a short, sterile bristle. Spikelets falling entire, ovate, acute, somewhat dorsally compressed; lower glume short, truncate, facing away from the rachis, 2-flowered; upper glume distinctly shorter than the spikelet, several-veined, the veins anastomosing toward the blunt apex; lower floret staminate, the lemma as long as the spikelet, several-veined, often somewhat sulcate, the palea well developed; upper floret slightly shorter than the spikelet, the lemma ovate, acute to apiculate, rugulose, pale, rigid, slightly concave on the back near the base, otherwise plano-convex, the margins strongly clasping the sides of the palea. Tropical and warm-temperate areas in the Neotropics and the Old World; 12– 40 species, 1 in Venezuela. Paspalidium geminatum (Forssk.) Stapf in Prain, Fl. Trop. Afr. 9: 583. 1920. —Panicum geminatum Forssk., Fl. Aegypt.-Arab. 18. 1775. Tufted plant, the culms decumbent and rooting at nodes, erect portions to 1 m tall; leaf blades 10–25 × 0.5–1 cm; inflorescence

15–20 cm long, of 10–20 erect to divergent racemes 1.5–3 cm long; spikelets 2–2.5 mm long. Marshes, along ditches and streams, 100–300 m; Bolívar (Altiplanicie de Nuria, Río Toro). Aragua, Falcón, Guárico, Sucre, Táchira, Zulia; Pantropics. ◆Fig. 168.

68. PASPALUM L., Syst. Nat. ed. 10, 2: 855, 1359. 1759. [Subfamily Panicoideae, Tribe Paniceae] Dimorphostachys E. Fourn., Compt. Rend. Hebd. Séances Acad. Sci. 80: 441. 1875. by Fernando O. Zuloaga and Emmet J. Judziewicz Annuals or perennials. Ligules membranous to membranous-ciliate. Blades lanceolate to filiform, involute or flattened. Inflorescences of 1–many racemes, these solitary, paired (sometimes conjugate), or racemosely arranged; rachis of racemes flattened to triquetrous, the spikelets borne in 2 rows on solitary (the raceme thus appearing 2-rowed) or paired (raceme appearing 4-rowed) pedicels, of-

Paspalidium 215

Fig. 168. Paspalidium geminatum

216

P OACEAE

ten overlapping, the upper glume and upper lemma facing toward the rachis; spikelets disarticulating entire from the pedicel. Spikelets terete, dorsally flattened, or most commonly plano-convex, 2-flowered, unwinged or (in one group of species) broadly winged; lower glume rudimentary (well developed in a few species) to more commonly absent; upper glume present (absent in a few species), as long as to somewhat shorter than the spikelet, 2–several-veined; lower floret sterile, the lemma as long as the spikelet, generally flattened, (2)3–5(–7)-veined, in a few species enclosing a palea; upper floret with lemma usually broadly rounded on back, slightly to considerably shorter than spikelet, papery to more commonly indurate, straw-colored to dark brown, smooth to pitted, papillose, or finely longitudinally striate, the margins commonly inrolled over the edges of a palea of similar texture; stamens usually 3; stigmas 2, plumose. Tropical and warm temperate regions worldwide; ca. 300 species, ca. 75 in Venezuela, 46 of these in the flora area. Key to the Species of Paspalum 1. 1. 2(1). 2. 3(2).

Rachis of racemes > 1 mm wide ................................................................ 2 Rachis of racemes < 1 mm wide .............................................................. 17 Rachis of racemes 5–7 mm wide; spikelets pilose .................................... 3 Rachis of racemes 1–3(–4) mm wide; spikelets glabrous or pilose .......... 4 Spikelets 5–7 mm long; racemes (1)2–6(–9) per inflorescence .................. ............................................................................................... P. lanciflorum 3. Spikelets 2.6–3.2 mm long; racemes 1 or 2 per inflorescence ..... P. stellatum 4(2). Raceme 1 per inflorescence, terminal ....................................................... 5 4. Racemes (1)2–many per inflorescence ...................................................... 8 5(4). Spikelets 5.2–8 mm long, cordate at the base .......................... P. pectinatum 5. Spikelets 2.1–5 mm long, not cordate at the base .................................... 6 6(5). Spikelets 3.3–5 mm long, pilose; upper anthecium shorter than the upper glume and lower lemma ......................................................... P. carinatum 6. Spikelets 2.1–3 mm long, glabrous; upper anthecium as long as the upper glume and lower lemma ........................................................................ 7 7(6). Raceme 3–5 cm long; foliage glabrous; lower glume absent; aquatic ............................................................................................. P. morichalense 7. Raceme 5–20 cm long; foliage pubescent; lower glume present; dry savannas ..................................................................................... P. pilosum 8(4). Spikelets 5–8 mm long, cordate at the base ............................................. 9 8. Spikelets 1.2–4.6 mm long, not cordate at the base ............................... 10 9(8). Lower lemma with prominent, submarginal, golden brown tubercles bearing spreading cilia, but without a tuft of cilia at the apex ................................................................................................ P. pectinatum 9. Lower lemma not bearing submarginal tubercle-based cilia, but with a tuft of several erect cilia at the apex ...................................... P. aspidiotes 10(8). Spikelets 3.7–4.6 mm long ...................................................... P. fasciculatum 10. Spikelets 1.2–3.5 mm long ....................................................................... 11 11(10). Racemes 2(3) per inflorescence ..................................................P. vaginatum 11. Racemes 3–many per inflorescence ......................................................... 12 12(11). Spikelets narrowly elliptic to lanceolate, 1.5–3 times as long as wide; plants aquatic ................................................................................ P. repens

Paspalum 217

12. 13(12).

13. 14(13). 14. 15(14). 15. 16(14).

16.

17(1). 17. 18(17). 18. 19(18). 19. 20(19). 20. 21(19). 21. 22(21). 22. 23(22). 23. 24(22). 24.

25(24).

Spikelets elliptic to broadly obovate or orbicular, 1–1.5 times as long as wide; plants terrestrial ........................................................................ 13 Spikelets (at least those in the upper portion of each raceme) biconvex, the upper and lower lemmas similar in texture, shiny, dark brown, indurate .................................................................................... P. convexum Spikelets plano-convex, the lower lemma flat, membranous, the upper lemma indurate, straw-colored to dark brown ................................... 14 Spikelets 2.3–3.2 mm long, with a fringe of hairs at the summit, drab grayish brown ...................................................................................... 15 Spikelets 1.6–2.3(-2.5) mm long, glabrous, greenish or straw-colored, often suffused with purple ..................................................................... 16 Upper floret brown at maturity; spikelets obovate, obtuse ........ P. virgatum Upper floret pale; spikelets elliptic, acute ............................. P. intermedium Racemes 60–200 in a congested panicle; rachis of racemes usually pilose; basal leaf sheaths usually inflated and nodulose, 1–3 cm wide when spread ........................................................................................... P. densum Racemes 13–50 in an open panicle; rachis of racemes glabrous, occasionally with a few pilose hairs; basal leaf sheaths not inflated and nodulose, 0.5–1.5 cm wide when spread ....................................... P. millegrana Raceme 1(2) per inflorescence ................................................................. 18 Racemes 2–many per inflorescence ......................................................... 29 Spikelets 2.6–3.6 mm long ......................................................... P. atabapense Spikelets 0.7–2.8 mm long ....................................................................... 19 Spikelets pentagonal to elliptic; lower glume absent, the midveins of the upper glume and lower lemma usually suppressed ........................... 20 Spikelets elliptic to obovate; lower glume present or absent, the midveins of the upper glume and lower lemma evident .................................... 21 Spikelets 1.4–2.1 mm long, elliptic; upper glume apiculate ... P. apiculatum Spikelets 1.2–1.3 mm long, obovate-pentagonal; upper glume acute ................................................................................................. P. arenarium Rachis of raceme 50–200 × 0.7–1.5 mm, pilose; lower lemma ± sulcate; foliage pilose ............................................................................... P. pilosum Rachis of raceme 10–70 × 0.3–1 mm, glabrous; lower lemma flat to slightly sulcate; foliage glabrous to pilose .......................................... 22 Spikelets 0.7–1.1 mm long, without capitellate hairs; upper floret strongly papillose ................................................................................. 23 Spikelets (0.9–)1.2–2.8 mm long (when as short as 0.9 mm long, then with capitellate hairs); upper floret smooth, not papillose ........................ 24 Upper glume 0.3–0.5 mm long, not covering the upper floret; spikelets 0.7–0.8 mm long ....................................................................... P. delicatum Upper glume 0.8–0.9 mm long, covering the upper floret; spikelets 0.9– 1.1 mm long ................................................................................... P. pictum Lower glume absent, upper glume and lower lemma with capitellate hairs; spikelets 0.9–1.6 mm long ....................................... P. clavuliferum Lower glume present (absent or to 0.2 mm long in P. nutans but spikelets 1.9–2.1 mm long), upper glume and lower lemma without capitellate hairs ..................................................................................................... 25 Lower glume absent or to 0.2 mm long; lower lemma flat on back; spikelets obovate ................................................................................... P. nutans

218

P OACEAE

25. 26(25). 26. 27(26). 27. 28(26). 28. 29(17). 29. 30(29). 30. 31(30). 31. 32(31). 32. 33(32). 33. 34(30). 34. 35(34). 35. 36(34). 36. 37(36). 37. 38(37). 38. 39(37). 39. 40(39). 40. 41(40). 41.

Lower glume 0.3–1 mm long (at least in some spikelets); lower lemma flat to slightly sulcate on back; spikelets elliptic to obovate .................... 26 Spikelets 1.8–2.8 mm long, elliptic ......................................................... 27 Spikelets 1.2–2 mm long, obovate or pentagonal ................................... 28 Spikelets broadly elliptic; upper glume glabrous or sparsely hairy on the margins ........................................................................................ P. altsonii Spikelets narrowly elliptic; upper glume finely appressed-pubescent all across the back ..................................................................... P. subfalcatum Spikelets pentagonal to slightly obovate, glabrous; leaf blades 5–17 mm wide ........................................................................................ P. decumbens Spikelets elliptic or obovate, with bracts fringed with silky hairs; leaf blades 3.5–6 mm wide ................................................................. P. petilum Racemes 2(3) per inflorescence ............................................................... 30 Racemes 3–many per inflorescence ......................................................... 46 Spikelets 0.5–1.5 mm long ....................................................................... 31 Spikelets 0.9–4.1 mm long ....................................................................... 34 Spikelets 0.5–0.8 mm long; rachis of racemes zigzag ..............P. parviflorum Spikelets 0.8–1.5 mm long; rachis of racemes straight to slightly flexuous .............................................................................................................. 32 Upper floret strongly papillose ......................................................... P. pictum Upper floret smooth ................................................................................. 33 Upper glume covered with glass-like, beaded hairs; plants tufted ................................................................................................. P. multicaule Upper glume not covered with glass-like hairs; plants creeping and rooting, forming mats ................................................................. P. orbiculatum Spikelets 2.4–4.1 mm long ....................................................................... 35 Spikelets 0.9–2.4 mm long ....................................................................... 36 Spikelets at least 2 times as long as wide, not spotted; plants strongly rhizomatous, stoloniferous .....................................................P. vaginatum Spikelets about as long as wide, minutely spotted; plants cespitose ............................................................................................... P. maculosum Spikelets fringed with spreading cilia ..................................... P. conjugatum Spikelets not fringed with cilia ............................................................... 37 Spikelets with both glumes absent ......................................................... 38 Spikelets with at least upper glume present .......................................... 39 Upper floret conspicuously papillose; spikelets ovate, 1.2–1.6 mm long, never purplish; ligules 5–6 mm long ....................................... P. nudatum Upper floret smooth; spikelets elliptic, 1.6–2.1 mm long, usually suffused with purple; ligules 0.3–0.6 mm long ................................... P. pulchellum Spikelets 0.9–1.4(–1.6) mm long ............................................................. 40 Spikelets 1.5–2.4 mm long ....................................................................... 42 Spikelets pentagonal; leaf blades lanceolate, 7–14 mm wide, papillosepilose at margin ...................................................................... P. arenarium Spikelets elliptic to obovate or round; leaf blades linear, 1.5–3.5 mm wide, not papillose-pilose at margin ............................................................. 41 Racemes 2 per inflorescence, divergent; spikelets paired, minutely pubes cent ...................................................................................... P. clavuliferum Racemes (2)3–9 per inflorescence, not divergent; spikelets solitary, glabrous ......................................................................................... P. hyalinum

Paspalum 219

42(39). Upper floret dark brown; racemes 1–4.5(–6) cm long ............................ 43 42. Upper floret tan, straw-colored, or greenish; racemes (3–)5–12 cm long .............................................................................................................. 44 43(42). Racemes 2 per inflorescence, conjugate; spikelets 1.4–1.8(–2.2) mm long; blades flat ...................................................................................... P. tillettii 43. Racemes 3–8 per inflorescence; spikelets 1.7–2.2 mm long; blades folded ....................................................................................... P. melanospermum 44(42). Racemes not conjugate, spikelets elliptic ............................................... 45 44. Racemes conjugate; spikelets broadly elliptic ......................... P. subciliatum 45(44). Spikelets 2.2–2.4 mm long; blades 10–20 cm long, glabrous or appressedpubescent abaxially ............................................................ P. corcovadense 45. Spikelets 1.4–2.1 mm long; blades 5–7 cm long, densely pilose ................ ............................................................................................... P. apiculatum 46(29). Spikelets nearly concealed by white, silky hairs 2 times as long as the spikelet ................................................................................ P. saccharoides 46. Spikelets glabrous or pubescent but not concealed by white, silky hairs .............................................................................................................. 47 47(46). Spikelets 2.5–4.6 mm long ....................................................................... 48 47. Spikelets 0.5–2.5 mm long ....................................................................... 55 48(47). Spikelets glabrous .................................................................................... 49 48. Spikelets pubescent ................................................................................. 51 49(48). Upper floret dark brown, shining; lower lemma crimped near margin ................................................................................................ P. plicatulum 49. Upper floret straw-colored to brown, not shining; lower lemma not crimped near margin ........................................................................... 50 50(49). Upper floret brown; racemes 15–30 per inflorescence, each 10–15 cm long, rachis glabrous ............................................................. P. conspersum 50. Upper floret pale; racemes 50–90 per inflorescence, each 2–15 cm long, rachis pilose ........................................................................ P. intermedium 51(48). Racemes 3–7 per inflorescence ................................................................ 52 51. Racemes 14–30 per inflorescence ............................................................ 53 52(51). Spikelets broadly oblanceolate, densely pubescent throughout; racemes 2–4 cm long ............................................................................... P. ammodes 52. Spikelets narrowly ovate, softly hairy on the margins only; racemes 5– 7 cm long .............................................................................. P. chaffanjonii 53(51). Spikelets lanceolate, 3.7–4.6 mm long ................................... P. fasciculatum 53. Spikelets elliptic, 2.7–3.2 mm long ......................................................... 54 54(53). Upper floret brown; racemes 15–30 per inflorescence, each 10–15 cm long, rachis glabrous ............................................................. P. conspersum 54. Upper floret pale; racemes 50–90 per inflorescence, each 2–15 cm long, rachis pilose ........................................................................ P. intermedium 55(47). Spikelets subtended by a fringe of golden hairs; upper floret dark brown and strongly papillose; both glumes absent ....................P. gardnerianum 55. Spikelets not subtended by golden hairs; upper floret not both dark brown and strongly papillose; glumes present or absent .................. 56 56(55). Spikelets 0.5–1.5 mm long ....................................................................... 57 56. Spikelets 1.5–2.5 mm long ....................................................................... 62 57(56). Leaf blades 7–25(–30) mm wide .............................................................. 58 57. Leaf blades 0.7–3.5(–6) mm wide ............................................................ 59

220

P OACEAE

58(57). Leaf blades 5–18 × 0.7–1.4 cm; spikelets pentagonal ............... P. arenarium 58. Leaf blades 18–33 × 1.5–2.5(–3) cm; spikelets obovate to broadly elliptic ............................................................................................. P. paniculatum 59(57). Upper glume and lower lemma 3–5-veined, the midvein evident; upper floret conspicuously tuberculate .................................................. P. pictum 59. Upper glume and lower lemma 2-veined, the midvein suppressed; upper floret smooth, not tuberculate ............................................................. 60 60(59). Spikelets 0.5–0.8 mm long, with a fringe of minute capitellate hairs on the margins of the upper lemma; leaf blades 0.7–2 mm wide ............... ...............................................................................................P. parviflorum 60. Spikelets 0.8–1.5 mm long, without a fringe of capitellate hairs on the margins of the upper lemma; leaf blades 2–3.5(–6) mm wide ........... 61 61(60). Upper floret straw-colored, elliptic; spikelets 1–1.4(–1.5) mm long; foliage usually with abundant, spreading, pilose hairs; plants tufted, found in dry places ................................................................................. P. hyalinum 61. Upper floret dark brown, round; spikelets 0.8–1.2(–1.5) mm long; foliage glabrous; plants creeping and rooting, forming mats in wet places .............................................................................................. P. orbiculatum 62(56). Racemes 11–200 per inflorescence .......................................................... 63 62. Racemes 2–12 per inflorescence .............................................................. 66 63(62). Racemes 60–200 per inflorescence .................................................. P. densum 63. Racemes 11–50 per inflorescence ............................................................ 64 64(63). Upper floret dark brown; lower lemma crimped near margin ................... ................................................................................................ P. plicatulum 64. Upper floret straw-colored; lower lemma not crimped near margin ..... 65 65(64). Spikelets pubescent, elliptic .................................................... P. coryphaeum 65. Spikelets glabrous, round to elliptic or obovate ....................... P. millegrana 66(62). Upper floret dark brown; lower lemma often crimped near margin ..... 67 66. Upper floret straw-colored to tan; lower lemma not crimped near margin .............................................................................................................. 69 67(66). Spikelets 2.3–2.4(–2.8) mm long, narrowly elliptic; racemes (5–)8–20 per inflorescence ........................................................................... P. plicatulum 67. Spikelets 1.6–2.2(–2.8) mm long, broadly obovate to elliptic; racemes 2– 8 per inflorescence ............................................................................... 68 68(67). Spikelets obovate, 1.8–3 mm long; upper glume usually at least sparingly appressed-pubescent................................................................ P. convexum 68. Spikelets round to elliptic, occasionally obovate, 1.7–2.2 mm long; upper glume usually completely glabrous ............................. P. melanospermum 69(66). Upper glume and lower lemma 2-veined (the midvein suppressed) ......... .................................................................................................. P. hyalinum 69. Upper glume and lower lemma 3–5-veined (the midvein evident) ............ ............................................................................................. P. corcovadense Paspalum altsonii Chase, J. Wash. Acad. Sci. 27: 144. 1937. Tufted plants; culms 35–70 cm tall; leaf blades 6–25 × 0.7–1.2 cm; racemes 1(2), 3–8 cm long; spikelets 2–2.8 mm long, glabrous to sparingly pubescent toward the margins;

lower glume present. Savannas, 100–1200 m; Bolívar (Guayaraca, between Kilómetro 88 and La Escalera, La Escalera), Amazonas (Río Mawarinuma). Guyana, Suriname. ◆Fig. 169.

Paspalum 221

Paspalum ammodes Trin., Gram. Panic. 120. 1826. Paspalum canum Sohns, Mem. New York Bot. Gard. 9: 256, fig. 6. 1957. Cespitose perennial 20–100 cm tall; leaves mostly basal, pilose; inflorescence of 2–6 racemes, each 2–4 cm long; spikelets 3.2–3.5 mm long, oblanceolate, densely pubescent. Savannas, near sea level to 1200 m; Bolívar (Río Arabopó basin, near Río Caura). Guyana, Brazil, Bolivia, Paraguay. Paspalum apiculatum Döll in Mart., Fl. Bras. 2(2): 48. 1877. Tufted perennial 30–60 cm tall; leaf blades 5–7 × 0.7–1 cm, densely pilose; inflorescences both terminal and lateral, each with 1(–4) racemes 3–8 cm long; spikelets elliptic, 1.4–2.1 mm long, densely papillosepilose to glabrous; upper glume and lower lemma 3-veined. Savannas, 100–300 m; north-central to northwestern Bolívar. Anzoátegui, Guárico, Táchira; Brazil. ◆Fig. 171. Paspalum arenarium Schrad. in Schult., Mantissa 2: 172. 1824. Tufted, stoloniferus perennial; culms 20– 70 cm tall; leaf blades 5–18 × 0.7–1.4 cm; inflorescences both terminal and lateral, each of the 1–3 racemes 4–7 cm long; spikelets 1.2–1.3 mm long, obovate-pentagonal, strawcolored with purple-based hairs. Savannas, ca. 300 m; Bolívar (Río Chicanán). Monagas; Guyana, Suriname, French Guiana, throughout Brazil. Paspalum aspidiotes Trin., Sp. Gram. 3: pl. 269. 1829–1830. Paspalum setiglume Chase, Brittonia 3: 150, fig. 1. 1939. Paspalum erectifolium Swallen, Fieldiana, Bot. 28: 22. 1951. Cespitose perennial 30–125 cm tall; foliage densely pubescent; inflorescence of (3)4– 7 ascending racemes, each 3–13 cm long; rachis 2–4 mm wide; spikelets 6–7.5 mm long; upper glume tipped with a few long-setose hairs; upper floret pilose. Savannas, 100– 1200 m; widespread in Bolívar and Amazonas. Colombia, Guyana, Brazil (Amazonas), Bolivia. ◆Fig. 173. Paspalum atabapense Davidse & Zuloaga, Novon 2: 193. 1992.

Short-rhizomatous perennial 45–75 cm tall; inflorescence a terminal raceme 3–9.5 cm long; rachis 0.5 mm wide; spikelets solitary, 2.6–3.6 mm long, pilose; upper glume shorter than the floret, 3-veined; lower lemma 3-veined; upper floret pale, papillose. White-sand savannas, ca. 100 m; Amazonas (western base of Cerro Yapacana). Endemic. Paspalum carinatum Humb. & Bonpl. ex Flüggé, Gram. Monogr., Paspalum 65. 1810. Cespitose perennial, culms 35–70 cm tall; leaf blades inrolled, densely papillose-pilose; raceme solitary, 5–12 cm long, occasionally a second raceme present; rachis 2–2.3 mm wide; spikelets 3.3–5 mm long; upper glume and lower lemma 3-veined, pilose; upper floret pilose at the apex. Sandy savannas, river banks, 50–1600 m; widespread in Bolívar and Amazonas. Apure, Monagas; Nicaragua, Trinidad and Tobago, Colombia, Guyana, Suriname, Brazil, Bolivia. ◆Fig. 174. Paspalum chaffanjonii Maury, J. Bot. 3: 159, fig. 5. 1889. Strongly rhizomatous perennial 60–80 cm tall; inflorescence terminal, of 3 or 4 racemes, each 5–7 cm long; rachis ca. 0.5 mm wide, glabrous; spikelets solitary, 2.9–3.1 mm long, acuminate, pilose toward the margins; upper floret papillose. Savannas, ca. 100 m; Bolívar (El Tigre, mouth of Río Pao). Barinas, Guárico, Portuguesa. Paspalum clavuliferum C. Wright, Anales Acad. Ci. Méd. Habana 8: 203. 1871. Delicate annual; culms 20–50 cm tall; inflorescences both terminal and lateral, 2, when paired each of conjugate racemes 2–5 cm long; spikelets 0.9–1.6 mm long, elliptic to obovate, with minute capitellate hairs. Savannas, 100–300 m; Bolívar (Altiplanicie de Nuria, San Félix). Barinas, Guárico; Mexico, Central America, Cuba, Haiti, Dominican Republic, Guyana, Colombia, Brazil, Bolivia. Paspalum conjugatum P.J. Bergius, Acta Helv. Phys.-Math. 7: 129, pl. 8. 1762. Colonial, stoloniferous plant, erect parts of culms to 100 cm tall; inflorescence a pair of divergent, conjugate racemes each 7–12 cm long; spikelets 1.2–1.8 mm long, ovate; lower lemma fringed with spreading, ciliate hairs. Pastures, forest edges, stream sides, and dis-

222

P OACEAE

turbed areas generally, 50–1000 m; widespread in Delta Amacuro, Bolívar, and Amazonas. Widespread elsewhere in Venezuela and throughout the Neotropics and Old World tropics. Paspalum conspersum Schrad. in Schult., Mantissa 2: 174. 1824. Coarse, cespitose perennial to 1.7 m tall; leaf blades 30–70 × 1.5–3 cm tall; inflorescence of 15–30 racemes, each 10–15 cm long; rachis 0.6–0.9 mm wide; spikelets 2.7–3.2 mm long, elliptic, acute. Sandy savannas, river banks, ca. 100 m; Amazonas (near Puerto Ayacucho, San Carlos de Río Negro). Mexico, Colombia, Guyana, Suriname, French Guiana, Brazil, Bolivia, Paraguay, Argentina. Paspalum convexum Humb. & Bonpl. ex Flüggé, Gram. Monogr., Paspalum 175. 1810. Tufted, often decumbent and stoloniferous plant, erect parts of culms 25–75 cm tall; inflorescence of 1–5 racemes, each 3–8.5 cm long; rachis 0.5–1.2 mm wide; spikelets 1.8–3 mm long, broadly obovate, plano-convex or sometimes biconvex (if lower floret indurate). Stream sides, 50–300(–1000) m; Bolívar (Altiplanicie de Nuria, La Vergareña, Santa Elena de Uairén), Amazonas (Puerto Ayacucho, Río Coromoto). Anzoátegui, Aragua, Barinas, Distrito Federal, Guárico, Portuguesa, Sucre; widespread in the Neotropics. ◆Fig. 176. Paspalum corcovadense Raddi, Agrostogr. Bras. 27. 1823. Cespitose perennial 30–90 cm tall; leaf blades 10–20 × 0.8–1 cm, tapering to both ends; inflorescence of 3–10 racemes, each 8– 11 cm long; spikelets elliptic, 2.2–2.4 mm long. Savannas, 900–1000 m; Bolívar (ca. 10 km southwest of Karaurín-tepui). Mexico, Belize, Honduras, Peru, Guyana, Brazil. Paspalum coryphaeum Trin., Gram. Panic. 114. 1826. Paspalum familiare Steud., Syn. Pl. Glumac. 1: 24. 1855 [1853]. Paspalum indutum Luces, J. Wash. Acad. Sci. 32: 162, fig. 6. 1942. Cespitose perennial 75–200 cm tall; leaf blades 20–38 × 1–2 cm, glabrous; inflores-

cence of 10–40 racemes, each 8–17 cm long; spikelets 1.6–2.6 mm long, elliptic, pubescent with brown-based hairs. Savannas; 100–800 m; Bolívar (Hato Divina Pastora, La Vergareña), Amazonas (Canaripó). Guárico, Falcón, Lara, Miranda, Sucre, Táchira; southern Mexico, Central America, Colombia, TrinidadTobago, Guyana, Suriname, French Guiana, Brazil. ◆Fig. 177. Paspalum decumbens Sw., Prodr. 22. 1788. Sprawling, decumbent and rooting plant, culms 20–40 cm tall; leaf blades 4–10 × 0.5– 1.7 cm; inflorescence a solitary, arcuate raceme 1.2–3.5 cm long; rachis sparingly pilose, 0.3–0.8 mm wide; spikelets 1.2–1.9 mm long, pentagonal to slightly obovate; lower glume cuff-like, the upper glume not covering the upper floret. Common on roadsides or other disturbed areas, near sea level to 500 (–1500) m; widespread in Delta Amacuro, Bolívar, and Amazonas. Widespread elsewhere in Venezuela and throughout the Neotropics. ◆Fig. 179. Paspalum delicatum Swallen, Contr. U.S. Natl. Herb. 29: 268. 1949. Delicate sprawling annual 20–30 cm tall; inflorescence a solitary arcuate raceme 1–3 (–4.5) cm long; spikelets 0.7–0.8 mm long, obovate; upper glume short and not covering the upper floret; lower lemma as long as but distinctly narrower than the upper floret. Low open forests on granitic outcrops, ca. 600 m; Amazonas (slope of Cerro Aracamuni). Apure, Guárico, Portuguesa; Colombia, French Guiana, Bolivia, Brazil. Paspalum densum Poir. in Lam., Encycl. 5: 32. 1804. Robust, cespitose perennial 1–3 m tall; leaves with basal sheaths inflated, shiny, nodulose; inflorescence of 60–200 racemes, each 4–9 cm long; rachis 1–2 mm wide, densely pilose; spikelets 1.6–2.4 mm long, orbicular, glabrous. Moist savannas, 100–300 m; Bolívar (Serranía Necuima on lower Río Caroní), Amazonas (San Juan de Manapiare). Anzoátegui, Apure, Cojedes, Guárico, Monagas, Portuguesa, Zulia; Mexico, Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Brazil, Bolivia. ◆Fig. 182.

Paspalum 223

Paspalum fasciculatum Willd. ex Flüggé, Gram. Monogr., Paspalum 69. 1810. Cespitose perennial 1–2 m tall; inflorescence of 10–30 racemes, each 7–15 cm long; spikelets 3.7–4.6 mm long, broadly lanceolate. Introduced for forage and sometimes naturalized, 50–100 m; Delta Amacuro (Isla Tortola, Tucupita), Amazonas (near San Vicente on Río Orinoco). Apure, Barinas, Falcón, Guárico, Miranda, Monagas, Zulia; widespread in the Neotropics. ◆Fig. 178. Paspalum gardnerianum Nees, Hooker’s J. Bot. Kew Gard. Misc. 2: 103. 1850. Slender tufted perennial 50–100 cm tall; inflorescence of 3–10(–12) distant racemes, each 2.5–9 cm long; spikelets 1.4–2.4 mm long, their pedicels bearing stiff golden hairs; upper glume absent; upper floret brown, papillose. Sandy savannas, 50–1100 m; Delta Amacuro (between Los Castillos and Piacoa), Bolívar (southeast of Chiguao, Gran Sabana, Quebrada Las Flores in Río Parguaza basin, lower Río Caura), Amazonas (30 km northnortheast of Puerto Ayacucho). Anzoátegui, Apure, Barinas, Falcón, Guárico, Sucre; Panama, Colombia, Guyana, Suriname, French Guiana, Brazil, Bolivia, Paraguay. ◆Fig. 172. Paspalum hyalinum Nees ex Trin., Gram. Panic. 103. 1826. Paspalum abstrusum Trin., Mém. Acad. Imp. Sci. Saint-Pétersbourg, Sér. 6, Sci. Math., Seconde Pt., Sci. Nat. 3: 135. 1834. Paspalum gossypinum Mez, Repert. Spec. Nov. Regni Veg. 15: 68. 1917. Paspalum polychaetum Mez, Bot. Jahrb. Syst. 56(Beibl. 125): 11. 1921. Tufted perennial 30–60 cm tall; leaves often with abundant spreading hairs; inflorescence of (2)3–9 racemes, each 1.5–7 cm long; spikelets solitary, 1–1.4(–1.5) mm long, elliptic, glabrous. Savannas, often where periodically flooded, 50–1300 m; widespread in Bolívar and Amazonas. Apure, Guárico; Colombia, Guyana, Suriname, French Guiana, Brazil, Peru, Bolivia, Paraguay. ◆Fig. 185. Paspalum intermedium Munro ex Morong & Britton, Ann. New York Acad. Sci. 7: 258. 1893. Cespitose perennial 1.2–2 m tall, the lower sheaths spongy, succulent; inflores-

cence of 50–90 racemes, each 2–15 cm long; rachis pilose; spikelets 2.2–3.2 mm long, elliptic; upper glume sparsely pilose. Marshy areas in savannas, ca. 800 m; Bolívar (Santa Elena de Uairén). Anzoátegui; Brazil, Bolivia, Paraguay, Argentina, Uruguay. Paspalum lanciflorum Trin., Sp. Gram. 3: pl. 286. 1829–1830. Paspalum contractum Pilg., Bot. Jahrb. Syst. 25: 709. 1898. Paspalum aureolatum Swallen, Fieldiana, Bot. 28: 22. 1951. Paspalum piligerum Swallen, Fieldiana, Bot. 28: 24. 1951. Cespitose perennial 50–80(–150) cm tall; leaves densely pubescent, the blades 17–35 × 0.3–1 cm; inflorescence of (1)2–6(–9) racemes, each 6–10 cm long; rachis 5–7 mm wide; spikelets 5–7 mm long; upper floret stipitate. Sandy and rocky savannas, 50– 1600 m; widespread in Bolívar and Amazonas. Anzoátegui, Apure; Panama, Colombia, Guyana, Suriname, Brazil, Bolivia. ◆Fig. 186. Paspalum maculosum Trin., Gram. Panic. 98. 1826. Cespitose perennial 60–90 cm tall; inflorescence a pair of divergent, conjugate racemes, each 4–16 cm long; spikelets 2.4–3 mm long, broadly elliptic, glabrous, usually minutely spotted. Savannas, 100–1000 m; Bolívar (Hato Divina Pastora, La Vergareña, Santa Elena de Uairén), Amazonas (near El Burro). Apure, Guárico; Colombia, Guyana, Suriname, Brazil, Bolivia, Paraguay, Argentina, Uruguay. ◆Fig. 188. Paspalum melanospermum Desv. ex Poir. in Lam., Encycl. suppl. 4: 315. 1816. Tufted annual, the culms decumbent below, erect parts 30–70 cm tall; inflorescence of 3–8 racemes, each 1–6 cm long; rachis 0.4– 0.9 mm wide; spikelets 1.7–2.2 mm long, elliptic. Savannas and disturbed areas such as gardens and Amerindian villages, river edges on sandy soils, 100–500 m; widespread in Delta Amacuro, Bolívar, and Amazonas. Aragua, Barinas, Distrito Federal, Mérida, Miranda, Táchira, Yaracuy, Zulia; Mexico, Panama, Lesser Antilles, Colombia, Guyana, Suriname, French Guiana, northeastern Brazil, Bolivia. ◆Fig. 175.

224

P OACEAE

Paspalum millegrana Schrad. in Schult., Mantissa 2: 175. 1824. Cespitose perennial 45–150 cm tall; sheaths spongy, succulent, leaf blades 30–60 × 0.7–1.3 cm, coarse; inflorescence of 13–50 racemes, each 5–15 cm long; rachis 1–1.5 mm wide; spikelets 1.8–2.5 mm long, orbicular, glabrous; upper glume and lower lemma papyraceous, 3-veined. Roadsides, moist savannas, near sea level to 100 m; Delta Amacuro (near Capure), Bolívar (Río Cuchivero, Santa María del Vapor in Municipio Piar), Amazonas (20 km southeast of Puerto Ayacucho). Anzoátegui, Apure, Barinas, Cojedes, Guárico, Miranda, Monagas, Portuguesa, Sucre, Zulia; southern Mexico, Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Brazil. ◆Fig. 183. Paspalum morichalense Davidse, Zuloaga & Filgueiras, Novon 5: 234. 1995. Creeping or floating aquatic perennial; culms rooting at the lower nodes, 13–25 cm tall; inflorescence a terminal raceme 3–5 cm long, occasionally 2 racemes present; rachis 1.8–3.5 mm wide, glabrous; spikelets solitary, 2.2–2.9 mm long, elliptic, glabrous. Mauritia palm swamps in open savannas, 100–200 m; Bolívar, Amazonas (base of Los Pijiguaos). Guárico; Colombia, Brazil, Suriname, Bolivia. Paspalum multicaule Poir. in Lam., Encycl. suppl. 4: 309. 1816. Tufted annual 10–40 cm tall; inflorescence of (1)2 slender racemes, each 1–4.5 cm long; rachis 0.5–0.8 mm wide; spikelets 0.8–1.3 mm long, covered with minute, beaded, glass-like hairs. Sandy savannas, 100–500 m; widespread in Bolívar and Amazonas. Anzoátegui, Guárico, Sucre; southern Mexico, Central America, Cuba, Trinidad-Tobago, Colombia, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia, Paraguay. Paspalum nudatum Luces, J. Wash. Acad. Sci. 32: 163, fig. 7. 1942. Slender, tufted plant to 60 cm tall; inflorescence of conjugate racemes, each 8–11 cm long; spikelets 1.2–1.6 mm long, ovate, glabrous; glumes absent; lower lemma as long and nearly as broad as upper floret. Seasonally flooded savannas, near sea level to 500

m; Bolívar (29 km southeast of Caicara, Caño Azul, Hato de Becerra). Apure, Cojedes, Guárico; Guyana, Suriname, Brazil (Mato Grosso), Bolivia. ◆Fig. 191. Paspalum nutans Lam., Tabl. Encycl. 1: 175. 1791. Creeping and rooting plant 30–70 cm tall; inflorescence of 1(–4) raceme(s) 3–6 cm long; spikelets 1.6–2.1 mm long, obovate, glabrous or sparingly pubescent; lower glume absent or to 2 mm long. Savannas, stream sides, 50– 500 m; Delta Amacuro (Sacupana), Bolívar (Altiplanicie de Nuria, between Kilómetro 88 and La Escalera, Represa Guri). Apure, Aragua, Falcón, Miranda, Nueva Esparta, Táchira, Zulia; Mexico, Guatemala, Belize, Honduras, El Salvador, Nicaragua, Costa Rica, Panama, Lesser Antilles, Colombia, Guyana, Suriname, French Guiana, Ecuador, Brazil. Paspalum orbiculatum Poir. in Lam., Encycl. 5: 32. 1804. Creeping, mat-forming plant, erect culms 10–15 cm tall; inflorescence of 3–9 racemes, each 1–4 cm long; spikelets 0.8–1.2(–1.5) mm long, orbicular to elliptic, pilose or glabrous; upper glume and lower lemma 2-veined, thin. Stream banks and muddy ground, 50– 500 m; Delta Amacuro (Sacupana), Bolívar (La Vergareña, Moitaco, Río Acanán, Río Caura, Río Suapure, Río Toro), Amazonas (Culebra, Isla San Sebastián on Río Casiquiare). Apure, Cojedes, Guárico, Miranda, Portuguesa; widespread in the Neotropics. ◆Fig. 190. Paspalum paniculatum L., Syst. Nat. ed. 10, 2: 855. 1759. Cespitose perennial 65–150 cm tall; leaf blades 18–33 × 1.5–2.5(–3) cm, softly pilose; inflorescence of 11–50 racemes, the lowest 5– 8 cm long; spikelets paired, 1.2–1.4 mm long, obovate to broadly elliptic, pubescent, brownish. Weedy places, near sea level to 900 m; Delta Amacuro (Ibaruma), Bolívar (La Vergareña, Puerto Ordaz, Río Ambutuir north of Urimán, Santa Elena). Widespread elsewhere in Venezuela and throughout the Neotropics. Paspalum parviflorum Rhode ex Flüggé, Gram. Monogr., Paspalum 98. 1810. Tufted annual 10–40 cm tall; leaves with spreading hairs; inflorescence of 1–4 ra-

Paspalum 225

cemes, each 1–3 cm long; rachis zigzag, pilose; spikelets 0.5–0.8 mm long, elliptic, pilose. Dry savannas, rock outcrops, 50–800 m; Bolívar (Caño Azul, La Vergareña), Amazonas (near Galipero). Costa Rica, Panama, Puerto Rico, Colombia, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia. ◆Fig. 192. Paspalum pectinatum Nees ex Trin., Sp. Gram. 1: pl. 117. 1828. Cespitose perennial 40–100 cm tall; leaves coarse, densely pubescent; inflorescence of (1)2 or 3 racemes, each 4–9 cm long; rachis 2.2–2.7 mm wide; spikelets 6–8 mm long, cordate; upper glume nearly flat, the lower lemma narrower than the upper glume, with tuberculate hairs; upper floret pilose. Savannas, edges of Mauritia palm swamps, 100– 1000 m; common in eastern Bolívar, Amazonas (east of Caño Parucito, La Esmeralda). Apure, Barinas; southern Mexico, Central America, Colombia, Guyana, Suriname, French Guiana, Peru, throughout Brazil, Bolivia, Paraguay. ◆Fig. 187. Paspalum petilum Chase, J. Wash. Acad. Sci. 27: 145. 1937. Paspalum dispar var. marahuacense H. Rodr., Ernstia 52: 1. 1989. Tufted, sprawling and rooting plant, culms 15–40 cm tall; leaf blades 60–110 × 3.5–6 mm, tapering to both ends; inflorescence of a solitary raceme 1–2.5(–4) cm long; spikelets 1.4–1.7(–2) mm long, elliptic or obovate; lower glume cuff-like to 1/2 or more the length of the spikelet, subulate; bracts all fringed with hairs, more so toward the margins. Stream banks, wet rocks, 100–1500 m; widespread in Bolívar and Amazonas. Guyana, Brazil (Amazonas). ◆Fig. 180. Paspalum pictum Ekman, Ark. Bot. 10(17): 11, t. 6, fig. 7. 1911. Delicate annual 40–60 cm tall; inflorescence of 2–7 racemes, each 3–6 cm long; spikelets 0.9–1.1 mm long, obovate-pyriform, glabrous; lower glume absent; upper glume narrowly triangular, nearly as long as the spikelet. Savannas, near sea level to 600 m; Amazonas (Caño Guaviarito, Río Manapiare, upper Río Parucito). Anzoátegui, Guárico, Monagas, Portuguesa; Costa Rica, Colombia, Guyana, Brazil, Bolivia. ◆Fig. 184.

Paspalum pilosum Lam., Tabl. Encycl. 1: 175. 1791. Cespitose perennial 60–100 cm tall; leaf blades 15–35 × 0.5–1 cm, pilose; inflorescence a solitary raceme, occasionally 2, 5–20 cm long; rachis 0.7–1.5 mm wide; spikelets 2.2–3 mm long, elliptic, glabrous, dimorphic with respect to the size of the lower glume; lower lemma slightly sulcate. Savannas, river banks, forest edges, 50–1400 m; Bolívar (El Pilón, Hato Divina Pastora, Las Patillas, Ptari-tepui), Amazonas (Capuana, Sierra Parima, Solano). Anzoátegui, Aragua, Distrito Federal, Falcón, Guárico, Mérida, Monagas, Táchira, Trujillo; southern Mexico, Central America, Colombia, Trinidad-Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 181. Paspalum plicatulum Michx., Fl. Bor.Amer. 1: 45. 1803. Cespitose perennial 50–150 cm tall; leaf blades 25–45 × 0.8–1.5 cm; inflorescence of (5–)8–20 racemes, each 5–12 cm long; rachis 0.8–1 mm wide; spikelets 2.3–2.4(–2.8) mm long, elliptic; lower lemma with a depressed central area and crimped margins. Wet savannas, 50–800 m; widespread in Bolívar and Amazonas. Widespread in Venezuela and throughout tropical and warm temperate America. Paspalum pulchellum Kunth, Mém. Mus. Hist. Nat. 2: 68. 1815. Cespitose perennial 50–90 cm tall; inflorescence of 2 racemes, each 1.5–9 cm long; rachis ca. 0.7 mm wide, flexuous; spikelets 1.6–2.1 mm long, elliptic, glabrous; glumes absent; lower lemma flat, purple, 3-veined; upper floret shiny, straw-colored. Savannas, 50–1300 m; widespread in Delta Amacuro, Bolívar, and Amazonas. Anzoátegui, Apure, Guárico, Monagas; southern Mexico, Central America, Greater Antilles, Colombia, Guyana, Suriname, French Guiana, Brazil. ◆Fig. 189. Paspalum repens P.J. Bergius, Acta Helv. Phys.-Math. 7: 129, pl. 7. 1762 [1772]. Floating or creeping aquatic perennial 30–60 cm tall; inflorescence of 20–100 racemes, the lowest 4–7 cm; rachis 1.5–2.3 mm wide and prolonged as a bristle; spikelets solitary, 1.2–2.2 mm long, lanceolate-

226

P OACEAE

elliptic; upper glume and lower lemma 2veined; upper floret smooth, shining. Shallow water, muddy shores, near sea level to 200 m; widespread in the flora area along Río Orinoco and major tributaries. Widespread elsewhere in Venezuela and throughout tropical and subtropical America. ◆Fig. 194. Paspalum saccharoides Nees ex Trin., Sp. Gram. 1: pl. 107. 1828. Perennial, culms robust, 1–2 m long; inflorescence of 30–50 racemes, each 15–25 cm long; rachis flexuous, 0.2–0.5 mm wide; spikelets 2.5–2.8 mm long, lanceolate; upper glume with long, silky hairs toward the margins; upper floret smooth. Roadsides, forest edges, 500–1400 m; Bolívar (La Escalera). Aragua, Barinas, Distrito Federal, Lara, Miranda, Monagas, Nueva Esparta, Portuguesa, Sucre, Táchira, Trujillo, Yaracuy; Central America, Lesser Antilles, Colombia, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 193. Paspalum stellatum Humb. & Bonpl. ex Flüggé, Gram. Monogr., Paspalum 62. 1810. Paspalum wagenerianum Schltdl., Linnaea 26: 15. 1854. Slender perennial 30–50 cm tall; inflorescence of 1 or 2 racemes, each 3–10 cm long; rachis 5–7 mm wide; spikelets 2.6–3.2 mm long, ovate, the callus heavily bearded; upper glume and lower lemma fringed with long white hairs; upper floret stipitate, smooth. Savannas, 800–1000 m; Bolívar (Hato Divina Pastora). Aragua, Barinas, Carabobo, Cojedes, Distrito Federal, Lara, Miranda, Portuguesa; Mexico, Central America, Hispaniola, Colombia, Brazil, Bolivia, Paraguay, Argentina, Uruguay. Paspalum subciliatum Chase, J. Wash. Acad. Sci. 17: 144, fig. 1. 1927. Tufted, short-rhizomatous perennial 20– 70 cm tall; inflorescence of conjugate racemes, each 2–8 cm long; spikelets 2–2.8 mm long, elliptic; upper glume pubescent, upper glume and lower lemma 3–5-veined; upper floret minutely papillose. Lowland savannas, ca. 100 m; Bolívar (El Tigre). Anzoátegui, Guárico; Panama, Guyana, French Guiana, Brazil.

Paspalum subfalcatum (Döll) Tutin, J. Bot. 72: 338. 1934. —Panicum subfalcatum Döll in Mart., Fl. Bras. 2(2): 181. 1877. Short-rhizomatous perennial 20–40 cm tall; leaf blades lanceolate, 65–95 × 3–5 mm; inflorescences terminal and lateral from the upper nodes; racemes 1(2), 2–7.5 cm long; spikelets 1.8–2.3 mm long, long-elliptic, sparsely pilose; lower glume dimorphic. Sandy river shores, ca. 100 m; Amazonas (Río Baría at mouth of Río Yatúa). Brazil (Amazonas). ◆Fig. 170. Paspalum tillettii Davidse & Zuloaga, Novon 2: 195. 1992. Cespitose perennial 20–60 cm tall; inflorescence terminal, racemes 2, each 1.5–6 cm long; rachis flexuous, 0.2–0.5 mm wide; spikelets 1.4–1.8(–2.2) mm long, dark, sparsely pilose with capitellate hairs; lower glume occasionally present; upper floret brownish, smooth. White-sand savannas, 100–200 m; Amazonas (Caño Caname, lower Río Pasimoni). Amazonian Colombia. Paspalum vaginatum Sw., Prodr. 21. 1788. Rhizomatous or stoloniferous perennial 25–50 cm tall; inflorescence of 2 conjugate racemes, each 2.5–7 cm long; rachis 0.5–2 mm wide; spikelets 2.6–3.5 mm long, glabrous. Brackish places near ocean, near sea level to 50 m; Delta Amacuro (mouth of Caño Güiniquina). Anzoátegui, Falcón, Sucre; Central America, Greater Antilles, most of South America, Old World tropics. Paspalum virgatum L., Syst. Nat. ed. 10, 2: 855. 1759. Coarse, cespitose perennial 1–2 m tall; leaf blades 35–90 × 1–3.5 cm; inflorescence of 7–30 racemes, the lowest 6–15 cm long; rachis 1.2–1.9 mm wide, scabrous and sparsely pilose; spikelets 2.3–3.2 mm long, broadly obovate, brownish; upper glume fringed with hairs; lower lemma glabrous to puberulent. Common in disturbed, open places and roadsides, also in moist savannas, swampy places, near sea level to 1000 m; widespread in Delta Amacuro, Bolívar, and Amazonas. Aragua, Barinas, Falcón, Guárico, Mérida, Miranda, Monagas, Portuguesa, Táchira, Yaracuy, Zulia; widespread in the Neotropics. ◆Fig. 195.

Paspalum 227

Fig. 169. Paspalum altsonii

Fig. 170. Paspalum subfalcatum

228

P OACEAE

Fig. 171. Paspalum apiculatum

Fig. 172. Paspalum gardnerianum

Paspalum 229

Fig. 173. Paspalum aspidiotes

Fig. 174. Paspalum carinatum

230

P OACEAE

Fig. 175. Paspalum melanospermum

Fig. 176. Paspalum convexum

Paspalum 231

Fig. 177. Paspalum coryphaeum

Fig. 178. Paspalum fasciculatum

232

P OACEAE

Fig. 180. Paspalum petilum

Fig. 179. Paspalum decumbens

Fig. 181. Paspalum pilosum

Paspalum 233

Fig. 182. Paspalum densum

Fig. 183. Paspalum millegrana

234

P OACEAE

Fig. 184. Paspalum pictum

Fig. 185. Paspalum hyalinum

Paspalum 235

Fig. 186. Paspalum lanciflorum

Fig. 187. Paspalum pectinatum

236

P OACEAE

Fig. 188. Paspalum maculosum

Fig. 189. Paspalum pulchellum

Fig. 190. Paspalum orbiculatum

Paspalum 237

Fig. 191. Paspalum nudatum

Fig. 192. Paspalum parviflorum

238

P OACEAE

Fig. 193. Paspalum saccharoides

Fig. 194. Paspalum repens

Paspalum 239

Fig. 195. Paspalum virgatum

240

P OACEAE

69. PENNISETUM Rich. in Pers., Syn. Pl. 1: 72. 1805. [Subfamily Panicoideae, Tribe Paniceae] by Emmet J. Judziewicz Annuals to robust perennials. Leaves with ligules membranous-ciliate; blades linear. Inflorescence densely spicate, of deciduous fascicles, each subtended by numerous, antrorsely scabrous, usually filiform bristles that enclose and partially conceal 1–several persistent spikelets. Spikelets sessile to pedicellate within the fascicles, lanceolate, dorsally compressed, glabrous; lower glume absent or to half as long as spikelet, hyaline, 2-flowered; upper glume nearly as long as the spikelet, several-veined; lower floret sterile or staminate, as long as the spikelet, the lemma 5–7-veined, the palea usually well developed; upper floret shorter than to nearly as long as the spikelet, hyaline to coriaceous. Worldwide in tropical and warm-temperate regions; ca. 80 species, ca. 5 in Venezuela, 2 of these in the flora area. Key to the Species of Pennisetum 1. 1.

Spikelets 1 in each fascicle, 3.2–4 mm long, subtended by ciliate bristles .............................................................................................. P. polystachion Spikelets 2–4 in each fascicle, each 4.5–6 mm long, subtended by eciliate bristles .................................................................................... P. purpureum

Pennisetum polystachion (L.) Schult., Mantissa 2: 146. 1824. —Panicum polystachion L., Syst. Nat. ed. 10, 870. 1759. —Tson. Cenchrus setosus Sw., Prodr. 26. 1788. —Pennisetum setosum (Sw.) Rich. in Pers., Syn. Pl. 1: 72. 1805. Coarse, rhizomatous perennial 1–3 m tall; inflorescence 10–20 cm long, the rachis glabrous. Savannas, disturbed open places, 100–1700 m; Bolívar (Gran Sabana, Río Orinoco basin), Amazonas (Río Orinoco, Río Samariapo). Anzoátegui, Apure, Aragua, Barinas, Carabobo, Guárico, Lara, Portuguesa;

U.S.A. (Florida), Mexico, Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina, throughout the paleotropics. ◆Fig. 196. Pennisetum purpureum Schumach., Beskr. Guin. Pl. 44. 1827. Robust perennial 2–5 m tall; inflorescence 13–25 cm long, the rachis pubescent. Old fields, ca. 100 m; Amazonas (Río Autana). Lara, Mérida, Miranda, Portuguesa, Táchira; widespread in Pantropics, native of Africa.

70. PHARUS P. Browne, Civ. Nat. Hist. Jamaica 344, pl. 38, fig. 3. 1756. [Subfamily Bambusoidae, Tribe Phareae] by Emmet J. Judziewicz Monoecious perennials (some species at least facultatively monocarpic). Culms erect to decumbent and rooting at the nodes, solid, producing stout prop roots. Leaves with ligules short, membranous; pseudopetioles elongate, conspicuous, twisted 180° at the summit and thus inverting the blade; blades linear to ovate, the veins diverging obliquely from the midrib. Inflorescence a terminal, open panicle, the spreading branches covered with minute hooked hairs and deciduous from the rachis; spikelets 1-flowered, borne on short branchlets, the sexes paired or some of the pistillate ones solitary. Pistillate spikelets larger than the staminate ones, subsessile, elongate; glumes unequal to subequal, lanceolate, persistent, several-

Pennisetum 241

Fig. 196. Pennisetum polystachion

242

P OACEAE

veined, purplish or less commonly green; floret indurate, cylindrical, linear to sigmoid, variously covered with minute, hooked hairs, longer than and disarticulating from the glumes, the lemma 7-veined, with scroll-like margins that embrace the palea; staminodes 6, minute; lodicules absent; stigmas 3, hispid. Staminate spikelets membranous, elliptic, persistent, borne on long pedicels appressed to the pistillate spikelet; lodicules 3, minute; stamens 6. Caryopsis elongate, with a small, basal embryo and a long linear hilum extending the full length of the fruit. Mexico, Central America, West Indies, tropical and subtropical South America; 7 species, 5 in Venezuela, all in the flora area. A forest understory grass, Pharus is adapted for animal dispersal, as evidenced by the hooked hairs on the branches of the inflorescence and pistillate florets. Key to the Species of Pharus 1. 1. 2(1). 2. 3(2). 3. 4(2).

4.

Pistillate glumes green; branches of the inflorescence detaching in verticils of 3 ........................................................................................ P. virescens Pistillate glumes purple; branches of the inflorescence detaching singly, not in verticils ........................................................................................ 2 Pistillate florets strongly sigmoid or at least falcate in the upper portion, tapering gradually to the pungent apex ............................................... 3 Pistillate florets straight throughout, abruptly short-beaked at the apex ................................................................................................................ 4 Pistillate glumes 2/3–3/4 as long as the slightly falcate floret ....... P. latifolius Pistillate glumes about 1/2 as long as the strongly sigmoid floret ...... P. mezii Leaf blades narrowly elliptic, acute at both base and apex, symmetrical, 10–20 × 1.7–3 cm; plants cespitose to weakly rhizomatous ............................................................................................... P. lappulaceus Leaf blades lanceolate, rounded basally, acuminate apically, asymmetrical, 9–12 × 1.5–2.5 cm; plants strongly rhizomatous and forming large clones, never cespitose ............................................................ P. parvifolius

Pharus lappulaceus Aubl., Hist. Pl. Guiane 859. 1775. Pharus glaber Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 196. 1815 [1816]. Cespitose perennial 30–100 cm tall; axis of inflorescence terminating in a staminate spikelet-tipped bristle; pistillate spikelets 7– 11 mm long. Moist to wet forests, 200–700 m; Bolívar (Serranía de Imataca, Upata), Amazonas (Piedra Cocuy). Anzoátegui, Aragua, Falcón, Mérida, Miranda, Monagas, Sucre, Táchira, Zulia; Mexico, Central America, West Indies, tropical and warm-temperate South America (south to Uruguay). ◆Fig. 197. Pharus latifolius L., Syst. Nat. ed. 10, 1269. 1759. —Pitillo. Cespitose perennial 50–100 cm tall; leaf blades 15–28 × (3–)5–8.5 cm, ovate, acumi-

nate; pistillate florets 13–18 mm long, bearded with hooked hairs only near the apex. Gaps in wet, often disturbed forests, near sea level to 500 m; Delta Amacuro (southeast of Piacoa, Serranía de Imataca), Bolívar (widespread), Amazonas (Río Baría, Río Orinoco). Widespread in wet lowlands of Venezuela; southern Mexico, Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 198. Pharus mezii Prodoehl, Bot. Arch. 1: 250. 1922. Cespitose perennial ca. 70 cm tall; leaf blades ca. 15 × 5 cm, obovate, acuminate; pistillate florets 11–13 mm long, bearded with hooked hairs only near the apex. Dry forests, ca. 100 m; Bolívar (between Río Parguaza

Pharus 243

Fig. 197. Pharus lappulaceus

Fig. 198. Pharus latifolius

244

P OACEAE

Fig. 199. Pharus virescens

Piresia 245

and the road from El Burro to Puerto Ayacucho). Cojedes, Portuguesa; southern Mexico, Guatemala, Nicaragua, Costa Rica, Panama, Colombia, Ecuador. Pharus parvifolius Nash, Bull. Torrey Bot. Club 35: 301. 1908. Southern Mexico, Central America, Greater Antilles, tropical South America; 2 subspecies, 1 in Venezuela. P. parvifolius subsp. parvifolius Clone-forming stoloniferous perennial (or monocarpic) with erect culms 25–40 cm tall; pistillate spikelets 10–13 mm long. Forests, ca. 200 m; Delta Amacuro (Serranía de Imataca). Barinas, Monagas, Yaracuy; Guatemala, Belize, Nicaragua, Panama, Greater

Antilles, Trinidad-Tobago, Guyana, Suriname, French Guiana, Brazil, Ecuador. Pharus virescens Döll in Mart., Fl. Bras. 2(2): 21. 1871. Clone-forming stoloniferous perennial (or monocarpic) with erect culms 50–100 cm tall; leaf blades 20–30 × 4–6 cm, often oblanceolate; pistillate spikelets 11–15 mm long, the glumes 1/2–3/4 as long as the straight, abruptly beaked lemma. Forests, often in old gaps, 100–600 m; Bolívar (Altiplanicie de Nuria, Río Samay affluent of Río Icabaru), Amazonas (base of Cerro Duida, Río Cunucunuma). Apure; Guatemala, Costa Rica, Panama, Dominican Republic, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil. ◆Fig. 199.

71. PHRAGMITES Adans., Fam. Pl. 2: 34, 559. 1763. [Subfamily Arundinoideae, Tribe Arundineae] by Gerrit Davidse Tall, stout, perennial, rhizomatous, colonial reeds. Rhizomes vigorous, 1–2 cm thick. Culms hollow with numerous elongated nodes. Leaves cauline; ligule a ciliate membrane; blades well developed, flat; Inflorescence a large, terminal, plumose panicle. Spikelets several-flowered; glumes narrow, the first 3-veined, the second 5veined, shorter than the florets; an evident internode between the first and second glumes; disarticulation above the first floret and the base of the rachilla internode beneath each succeeding floret; rachilla internodes, except the lowermost, covered with numerous long, silky hairs; lowermost floret persistent, its lemma 5-veined, the flower staminate; other florets with bisexual flowers; lemmas slender, 3-veined, acuminate, glabrous; paleas much shorter than the lemmas; uppermost florets shorter than the lower ones. Widespread in temperate and tropical zones of the world; 5 species, 1 in Venezuela. Phragmites australis (Cav.) Trin. ex Steud., Nomencl. Bot. 2(2): 324. 1841. —Arundo australis Cav., Ann. Hist. Nat. 1: 100. 1799. Arundo phragmites L., Sp. Pl. 81. 1753. Arundo altissima Benth., Cat. Pl. Pyrénées 62. 1826.

Phragmites communis Trin., Fund. Agrost. 134. 1841. Reed 1–4 m tall, with large, silvery panicles. Margins of rivers, marshes, ditches, 50–500 m; rare, Bolívar (Represa Guri). Aragua, Falcón, Zulia; Cosmopolitan. ◆Fig. 200.

72. PIRESIA Swallen, Phytologia 11: 152. 1964. [Subfamily Bambusoideae, Tribe Olyreae] by Emmet J. Judziewicz Monoecious, cespitose perennials. Culms usually dimorphic. Vegetative culms erect, leafy, sterile, in the upper half bearing a pinnate- to palmate-appearing leaf complement; leaves with ligules short, membranous; pseudopetioles short; blades

246

P OACEAE

Fig. 200. Phragmites australis

Piresia 247

linear-lanceolate to oblong. Floriferous culms decumbent, creeping, often buried in leaf litter, with leaves reduced to bladeless sheaths or minute, reflexed blades, all nodes bearing racemiform inflorescences only partially exserted from the sheaths. Individual inflorescences bearing a terminal pistillate spikelet accompanied by several staminate spikelets below it, occasionally 1 or 2 more pistillate spikelets present below these. Spikelets 1-flowered, the pistillate ones much larger than the staminate ones. Pistillate spikelets borne on clavate pedicels; glumes as long as the spikelet, subequal, elliptic-ovate, short-apiculate, often purplish, 5-veined; floret elliptic, indurate, pubescent, nearly as long as the glumes, becoming marbled with dark spots at maturity; stigmas 2. Staminate spikelets borne on filiform pedicels, deciduous, hyaline, lacking glumes; stamens 3. Colombia, Venezuela, Trinidad-Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 4 species, 1 in Venezuela.

Fig. 201. Piresia sympodica

248

P OACEAE

Piresia sympodica (Döll) Swallen, Phytologia 11: 153. 1964. —Olyra sympodica Döll in Mart., Fl. Bras. 2(2): 322. 1877. Cespitose perennial ca. 30 cm tall; leaves 5–7 in a fan-shaped complement, the blades 5–8 × 0.8–1.1 cm, the lower surface bluish

green; pistillate spikelets 8–9 mm long. Forest understories, 200–500 m; Bolívar (28 km south of El Dorado, Río Ambutuir south of Auyán-tepui). Colombia, Trinidad-Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Amazonian Brazil. ◆Fig. 201.

73. RADDIELLA Swallen, Bull. Torrey Bot. Club. 75: 89. 1948. [Subfamily Bambusoideae, Tribe Olyreae] by Emmet J. Judziewicz Monoecious tufted perennials, or delicate, mat-forming, and of indefinite duration. Leaves small, pseudopetiolate, the blades elliptic to ovate-triangular, often strongly asymmetrical and with an apiculate apex, exhibiting sleep movements or not. Inflorescences small, few-flowered, 1–several from both terminal and axillary nodes, barely exserted from the leaf sheaths. Terminal inflorescences usually staminate, sometimes both staminate and pistillate; axillary inflorescences either all pistillate or with both pistillate and staminate spikelets; pedicels of pistillate spikelets minutely cupulate at the apices. Spikelets 1-flowered, the broad pistillate spikelets slightly shorter than the narrower staminate ones. Pistillate spikelets ovate-lanceolate, falling entire from pedicels or the glumes persistent and the floret falling; glumes about equal, membranous, as long as spikelet, acute, several-veined; floret elliptic, acute, glabrous, thinly coriaceous, straw-colored or becoming dark when mature, the margins of the lemma inrolled over the edges of the palea; stigmas 2. Staminate spikelets borne on capillary pedicels, linear to lanceolate, hyaline, early deciduous, lacking glumes; stamens 3. Panama, Colombia, Venezuela, Trinidad-Tobago, Guyana, Suriname, French Guiana, Brazil, Bolivia; 7 species, 3 in Venezuela, all in the flora area. Key to the Species of Raddiella 1.

1.

2(1). 2.

Leaf blades 10–20 × 6–10 mm, ovate-triangular, asymmetrical, apiculate, firmly membranous, glabrous, exhibiting sleep movements, folding upwards at night or under water stress; pistillate spikelets 1.8–2.7 mm long; common, often in dry places .......................................... R. esenbeckii Leaf blades 6–10 × 2.5–5.5 mm, ovate-elliptic, symmetrical, acute, delicately membranous, glabrous or puberulent, not exhibiting sleep movements; pistillate spikelets 1.3–2 mm long; rare, shaded cliffs and bases of waterfalls ................................................................................. 2 Leaf blades densely puberulent beneath .......................... R. aff. kaieteurana Leaf blades glabrous or puberulent only on the veins beneath ................. ......................................................................................... R. aff. potaroensis

Raddiella esenbeckii (Steud.) C.E. Calderón & Soderstr., Smithsonian Contr. Bot. 44: 21. 1980. —Panicum esenbeckii Steud., Syn. Pl. Glumac. 1: 90. 1855 [1854], based on Panicum laterale Nees var. ß Nees, Agrost. Brasil. 213. 1829.

—Olyra nana Döll in Mart., Fl. Bras. 2(2): 329. 1877, nom. superfl. —Raddia nana (Döll) Chase, Proc. Biol. Soc. Wash. 21: 185. 1908. —Raddiella nana (Döll) Swallen, Bull. Torrey Bot. Club 75: 89. 1948.

Raddiella 249

Fig. 202. Raddiella aff. kaieteurana

Cespitose perennial 10–30 cm tall; culms numerous; pistillate spikelets with glumes pubescent, the lemma smooth. Sandy or gravelly savannas, rock outcrops, river banks, forest margins, often on lowest slopes of tepuis, 100–1300 m; Bolívar (widespread), Amazonas (widespread). Anzoátegui, Falcón, Sucre, Táchira; Panama, Colombia, TrinidadTobago, Guyana, Suriname, French Guiana, Brazil, Bolivia. ◆Fig. 203. Raddiella aff. kaieteurana Soderstr., Mem. New York Bot. Gard. 12(3): 6. 1965. Sprawling herb ca. 10 cm tall. Moist, semishaded rock faces, dripping cliff bases, bases of waterfalls, 600–800 m; Bolívar (Amaruay-tepui). Guyana, Suriname, Brazil (Pará). ◆Fig. 202. The flora area specimen of Raddiella aff. kaieteurana has pistillate spikelets with glabrous glumes. Raddiella aff. potaroensis Soderstr., Mem. New York Bot. Gard. 12(3): 6. 1965. Raddiella maipuriensis Soderstr., Mem. New York Bot. Gard. 12(3): 7. 1965. Sprawling herb ca. 10 cm tall; both collections sterile; wet, streamside rocks in spray zone of waterfalls, 500–1000 m; Bolívar (Río Chicanán, Salto Kamá in Río Aponguao basin). Guyana. Outside the flora area, the pistillate spikelets are 1.3–2 mm long, the staminate spikelets 2.7–4 mm long.

Fig. 203. Raddiella esenbeckii

250

P OACEAE

74. REIMAROCHLOA Hitchc., Contr. U.S. Natl. Herb. 12: 198. 1909. [Subfamily Panicoideae, Tribe Paniceae] by Emmet J. Judziewicz Tufted or mat-forming perennials, or of indefinite duration, sprawling and forming mats. Leaves with ligules membranous-ciliate; blades linear. Cleistogamous inflorescences produced within lower leaf sheaths, otherwise inflorescences terminal, of several to many scattered or more commonly subdigitate, deciduous racemes; racemes stiff, often deflexed at full maturity, the rachis filiform to flattened, triquetrous, bearing 2 rows of solitary spikelets, the lowermost spikelet much

Fig. 204. Reimarochloa acuta

Rhipidocladum 251

the largest, the upper ones gradually reduced in size. Spikelets all cleistogamous, ovate to narrowly lanceolate, acute to attenuate, strongly dorsally compressed, hyaline to firmly membranous, straw-colored, glabrous to finely spreading-ciliate on margins, deciduous and falling together with the short pedicel, 2-flowered; glumes absent; lower floret sterile, as long as spikelet, the lemma 3-veined, the palea absent; upper floret slightly shorter than spikelet, thinly coriaceous, the lemma 3veined, its margins not inrolled over the edges of the well-developed palea. U.S.A. (Florida), Honduras, El Salvador, Cuba, Hispaniola, Colombia, Venezuela, Guyana, French Guiana, Peru, Brazil, Bolivia, Paraguay, Argentina; 3 species, 1 in Venezuela. Reimarochloa acuta (Flügge) Hitchc., Contr. U.S. Natl. Herb. 12: 198. 1909. —Reimaria acuta Flügge, Gram. Monogr., Paspalum 217. 1810. Agrostis brasiliensis Spreng., Novi. Provent. 45. 1818 [1819]. —Reimarochloa brasiliensis (Spreng.) Hitchc., Contr. U.S. Natl. Herb. 12: 198. 1909. Tufted, extensively stoloniferous plant, erect portions of culms 10–20 cm tall; inflorescence of 3–14 erect, divergent, or reflexed

racemes 17–25 mm long borne on a rachis to 15 mm long; lowermost spikelet in each raceme 3.8–6 mm long, the lower glume glabrous or with a few marginal, spreading cilia. Muddy or sandy shores, near sea level to 200 m; common in the flora area along Río Orinoco and major tributaries. Anzoátegui, Apure, Barinas, Guárico; Honduras, El Salvador, Cuba, Hispaniola, Colombia, Venezuela, Guyana, French Guiana, Peru, Brazil, Bolivia, Paraguay, Argentina. ◆Fig. 204.

75. RHIPIDOCLADUM McClure, Smithsonian Contr. Bot. 9: 101. 1973. [Subfamily Bambusoideae, Tribe Bambuseae] by Emmet J. Judziewicz Small to medium-sized, cespitose, woody bamboos from short, determinate rhizomes. Culms erect, hollow, thin-walled, weak, unarmed, usually arching and drooping in the upper portions; branches at midculm nodes in a fan-shaped or triangular cluster, numerous, equal, produced at some distance above the nodal line, unbranched. Culm leaves well developed or not, the blades erect, often apiculate. Foliage leaves with both inner and outer ligules present; oral setae present but usually not prominent; pseudopetioles short; blades linear to lanceolate, lacking a conspicuous midvein, borne in loose complements near the ends of the branches. Inflorescence a spike-like raceme or sparingly branched panicle, the spikelets often restricted to one side of the rachis. Spikelets terete to laterally compressed, severalflowered, the glumes and lowest (sterile) lemma persistent; glumes 2, unequal; lower (sterile) lemmas 1(2); functional florets several, the lemmas 5–7-veined, acuminate to long-awned; stamens 3; stigmas 2. Mexico, Central America, West Indies, tropical South America (except eastern Brazil); ca. 25 species, ca. 8 in Venezuela, 4 of these in the flora area. Key to the Species of Rhipidocladum 1. 1. 2(1). 2.

Spikelets 2 or 3 per inflorescence, with awns 4–6 mm long ......... R. sibilans Spikelets > 3 per inflorescence, awnless or with awns < 2 mm long ....... 2 Culms 12–15 mm diameter; foliage leaf blades 9–12 mm wide ....... R. sp. A Culms 5–10 mm diameter; foliage leaf blades 5–9 mm wide .................. 3

252

3(2). 3.

P OACEAE

Inflorescence with 10–15 stout, crowded spikelets ............. R. racemiflorum Inflorescence with 4 or 5 slender, distant spikelets .......................... R. sp. B

Rhipidocladum aff. racemiflorum (Steud.) McClure, Smithsonian Contr. Bot. 9: 106. 1973. —Arthrostylidium racemiflorum Steud., Syn. Pl. Glumac. 1: 336. 1855 [1854]. Cespitose woody bamboo forming dense thickets, 2–4 m tall but scandent and pendent to 9 m; 12–25 branches per midculm node; foliage leaf blades 50–80 × 5–9 mm; inflorescence (not seen in the flora area) ca. 5 cm long, with 10–15 crowded spikelets 15–20 mm long; lemmas often with prominent apical hair tufts. Gallery forests, bases of granitic outcrops, 100–900 m; Bolívar (Gran Sabana, Río Tirica), Amazonas (Caño Caname). Aragua, Distrito Federal, Falcón, Lara, Yaracuy, Zulia; Mexico, Central America, Trinidad-Tobago, throughout tropical and subtropical South America. Rhipidocladum sibilans Davidse, Judz. & L.G. Clark, Novon 1: 84. 1991. Cespitose, scandent woody bamboo; culms 4–8 m long, 7–15 mm diameter; 100–150 branches per midculm node; foliage leaf blades 50–70 × 1.8–3.5 mm; inflorescence of 2 or 3 spikelets; spikelets 2–4 cm long, 4–7flowered, both glumes and lemmas with awns 4–6 mm long. Gallery forests, 900– 1200 m; Bolívar (Gran Sabana). Anzoátegui, Sucre. ◆Fig. 205.

Rhipidocladum sp. A Woody bamboo; culms to 4 m tall, 15 mm diameter, smooth, weak; culm leaves 9 cm long, the blades 3.5 cm long, triangular, with an apiculus 1 cm long; 20–30 branches per midculm node; foliage leaves with delicate orange-brown oral setae 2–3 mm long, the blades 65–80 × 9–12 mm; both collections sterile. Gallery forests, ca. 100 m; Bolívar (Quebrada Las Flores in Río Parguaza basin), Amazonas (22 km south of Puerto Ayacucho). Generic placement is uncertain; sterile collections bear some resemblance to Arthrostylidium longiflorum Munro of the Venezuelan Andes. Rhipidocladum sp. B could be conspecific. Rhipidocladum sp. B Woody bamboo; culms to 5 m tall, 7 mm diameter, smooth; culm and foliage leaves not seen; ca. 50 branches per midculm node, each branch ca. 30 cm long, leafless, flowering; inflorescence with rachis slender, straight; spikelets 1.5–2.5 cm long, apparently 3-flowered; lemmas 10–14 mm long, narrowly lanceolate, awnless. Abundant at edge of granitic outcrops, ca. 300 m; Bolívar (Caño Trapichote off Río Orinoco, near Puerto Ayacucho).

76. RHYNCHELYTRUM Nees in Lindl., Intr. Nat. Syst. Bot. ed. 2, 446. 1836. [Subfamily Panicoideae, Tribe Paniceae] by Emmet J. Judziewicz Annuals or perennials. Leaves with ligules ciliate; blades linear. Inflorescence an open but rather bushy panicle, the spikelets usually concealed by long, silky hairs, falling entire from the pedicel or the upper floret falling first. Spikelets ovate to elliptic, laterally compressed, 2-flowered; lower glume short, linear; upper glume and lower lemma similar, subequal, as long as spikelet, ovate, beaked, 5-veined, keeled, often short-awned from just below the emarginate or bidentate summit, usually densely covered with ascending silky hairs; lower floret staminate, the palea well developed; upper floret distinctly shorter than the spikelet, strongly laterally compressed, narrowly elliptic, acute, smooth, shining. Tropical and South Africa (one species naturalized throughout the world in tropical and subtropical regions); 10–15 species, 1 in Venezuela.

Rhipidocladum 253

Fig. 205. Rhipidocladum sibilans

254

P OACEAE

Fig. 206. Rhynchelytrum repens

Rhytachne 255

Rhynchelytrum repens (Willd.) C.E. Hubb., Bull. Misc. Inform. (1934): 110. 1934. —Saccharum repens Willd., Sp. Pl. ed. 4, 1(1): 322. 1797. —Melinis repens (Willd.) Zizka, Biblioth. Bot. 1380: 55. 1988. Tricholaena rosea Nees, Linnaea 11: 129. 1837. —Rhynchelytrum roseum (Nees) Stapf & C.E. Hubb. in Prain, Fl. Trop. Afr. 9: 880. 1930.

Cespitose perennial 0.5–1 m tall; inflorescence 10–15 cm long, the branches filiform, flexuous; spikelets 3.5–4.5 mm long, partly concealed by reddish hairs. Dry weedy places; Delta Amacuro (Barrancas to Tucupita), Bolívar (Ciudad Bolívar, San Martín de Turumbán southwest of Tumeremo), Amazonas (Puerto Ayacucho). Naturalized in tropical and subtropical regions worldwide, native of Africa. ◆Fig. 206.

77. RHYTACHNE Desv. ex Ham., Prodr. Pl. Ind. Occid. xiv, 11. 1825. [Subfamily Panicoideae, Tribe Andropogoneae] Lepturopsis Steud., Syn. Pl. Glumac. 1: 357. 1855 [1854]. by Gerrit Davidse Perennials or rarely annuals, cespitose. Culms hollow. Leaf sheaths rounded or slightly keeled; ligule a ciliolate membrane; blades linear, involute to flat. Inflorescence a solitary, cylindrical raceme generally terminal sometimes axillary; rachis articulated, bearing paired spikelets, the 2 spikelets and rachis internode segment falling as 1 unit; internodes clavate, semicylindrical, as long as or longer than spikelets, usually glabrous, rarely pubescent. Spikelets dorsally compressed, sessile and pedicellate. Sessile spikelets bisexual, 2-flowered, lanceolate, with or without an awn; callus truncate with central peg which becomes oily at caryopsis maturity; lower glume chartaceous to crustaceous, smooth, rugose or longitudinally ribbed, 2keeled or rounded on flanks, wingless or obscurely winged, obtuse or 2-denticulate, rarely awned; upper glume membranous, navicular, 3-veined, sometimes awned; lower floret sterile or staminate, the lower lemma and palea hyaline; upper floret bisexual; upper lemma and palea hyaline; lodicules 2; stamens 3; styles 2; hilum punctate. Pedicellate spikelets usually rudimentary rarely as large as the sessile spikelets or represented by an awn; pedicel free. Neotropics, tropical Africa; 12 species, 4 in Venezuela, 3 of these in the flora area. This treatment follows the review of Clayton (Kew Bull. 32: 767–771. 1978). The perennial American species are not very distinct. There is often a great deal of variation in spikelets within the same inflorescence and there are intermediate populations between all three species recognized here. It might be more realistic to recognize these entities at an infraspecific level. The annual Rhytachne gonzalezii Davidse, which has been collected in nearby Guárico and in Suriname, is to be expected in seasonally wet areas in the Trachypogon savannas of Bolívar. Key to the Species of Rhytachne 1. 1. 2(1).

Annuals; spikelets awned, the awns 5–14 mm long (expected) ... R. gonzalezii Perennials; spikelets usually awnless, rarely with awns to 2 mm long ................................................................................................................ 2 Second internode from the base of the raceme 10–18 mm long, reducing to 6–13 mm in the middle of the raceme; sessile spikelet 3 5–9.5 mm long; lower glume of sessile spikelets smooth, especially near the base

256

2.

3(2). 3.

P OACEAE

of the raceme, to strongly and coarsely rugose, especially toward the tip of a raceme ...................................................................... R. subgibbosa Second internode from the base of the raceme 2.5–8 mm long, not greatly longer than the rest of internodes; lower glume of sessile spikelets smooth to strongly and coarsely rugose ................................................ 3 Rachis internodes 2.5–5.5 mm long; spikelets 2–6 mm long; lower glumes strongly and coarsely rugose or muricate on the keels ... R. rottboellioides Rachis internodes 5–8 mm long; spikelets 4.5–6.5 mm long; lower glumes entirely smooth or muricate only on the keels ..................... R. guianensis

Rhytachne guianensis (Hitchc.) Clayton, Kew Bull. 20: 262. 1966. —Manisuris guianensis Hitchc., Contr. U.S. Natl. Herb. 22: 510. 1922. Perennial to 90 cm tall. Wet areas in savannas, 50–1100 m; Bolívar (Guayaraca, Hato la Vergareña, Río Aícha near Los Caribes, upper Río Arabopó), Amazonas (Caño Cotúa). Apure, Guárico; Colombia, Guyana, Suriname, Brazil, Bolivia. Rhytachne rottboellioides Desv. ex Ham., Prodr. Pl. Ind. Occid. 12. 1825. Rottboellia loricata Trin., Mém. Acad. Imp. Sci. St. Pétersbourg, Sér. 6, Sci. Math. 2(3): 250. 1832. —Manisuris loricata (Trin.) Kuntze, Revis. Gen. Pl. 2: 780. 1891. —Coelorachis loricata (Trin.) Nash, N. Amer. Fl. 17(1): 85. 1909. Perennial to 90 cm tall. Wet areas in sa-

vannas, rocky river banks, 500–1300 m; Bolívar (Gran Sabana). Apure; Caribbean, Brazil, Bolivia, Paraguay. ◆Fig. 207. Rhytachne subgibbosa (Winkl. ex Hack.) Clayton, Kew Bull. 20: 261. 1966. —Rottboellia loricata subsp. subgibbosa Winkl. ex Hack. in Mart., Fl. Bras. 2(3): 311, t. 71, fig. 2. 1883. —Mnesithea subgibbosa (Winkl. ex Hack.) de Koning & Sosef, Blumea 31(2): 292. 1986. Rottboellia loricata subsp. glaberrima Hack. in Mart., Fl. Bras. 2(4): 311, pl. 71, fig. 3. 1883. Perennial to 160 cm tall. Mauritia palm swamps, marshy areas in savannas, 50–800 m; Bolívar (Hato Divina Pastora, Hato la Vergareña), Amazonas (vicinity of Puerto Ayacucho). Guárico; Mexico, Brazil, Bolivia, Paraguay, Argentina.

78. SACCHARUM L., Sp. Pl. 54. 1753. [Subfamily Panicoideae, Tribe Andropogoneae] Erianthus Michx., Fl. Bor.-Amer. 1: 54. 1803. by Emmet J. Judziewicz Coarse, cespitose perennials. Inflorescence large, a cylindrical to ovate, plumy panicle of numerous ascending racemes; rachis internodes filiform to weakly clavate, falling together with the persistent sessile spikelet, the pedicellate spikelet often early-deciduous. Spikelets similar, both with the lower floret sterile and the upper bisexual; callus heavily bearded; glumes firmly membranous, glabrous to villous, as long as spikelet; lower glume flat to weakly concave or convex on back, with 2 evident submarginal veins, the midvein usually suppressed, the margins inflexed over the keeled, 1–3-veined upper glume and florets; florets hyaline; lower floret nearly as long as the spikelet, lacking a palea; upper floret about half as long as the spikelet, the lemma awned or not, the palea rudimentary but present. Tropical and warm-temperate regions worldwide; ca. 35 species, 2 in Venezuela, both in the flora area. Saccharum officinarum L. (caño de azucar, sugar cane), with sweet pith, is cultivated locally in the flora area, including gardens of Amerindian tribes.

Rhytachne 257

Fig. 207. Rhytachne rottboellioides

258

P OACEAE

Fig. 208. Saccharum trinii

Sacciolepis 259

Key to the Species of Saccharum 1. 1.

Spikelets 4–5 mm long, the glumes glabrous ............................... S. asperum Spikelets 6–8 mm long, the glumes sparingly to densely villous, at least at margins ....................................................................................... S. trinii

Saccharum asperum (Nees) Steud., Syn. Pl. Glumac. 1: 407. 1855 [1854]. —Erianthus asper Nees in Mart. et al., Fl. Bras. Enum. Pl. 2: 315. 1829. Stout cespitose perennial 2–3 m × 1–1.5 cm, the nodes bearded; leaf blades 50–75 × 1.5–3.5 cm; inflorescence 17–25 × 7–12 cm, ovate, often pinkish or purplish, less commonly white. Savannas, 1200–1800 m; Bolívar (Gran Sabana, La Escalera). Apure; Brazil, Bolivia, Paraguay, Argentina, Uruguay. Saccharum trinii (Hack.) Renvoize, Kew Bull. 39: 184. 1984. —Saccharum giganteum Trin., Mém. Acad. Imp. Sci.

St.-Pétersbourg, Sér. 6, Sci. Math. 2: 311. 1832, non (Walter) Pers. 1805. —Erianthus saccharoides var. ß trinii Hack. in Mart., Fl. Bras. 2(3): 258. 1883. —Erianthus trinii (Hack.) Hack. in A. DC., Monogr. Phan. 6: 135. 1889. Stout cespitose perennial 1.5–3 m × 1–2 cm, the nodes bearded; leaf blades 45–75 × 1– 2 cm; inflorescence 25–35 × 4–7 cm, cylindrical, white. Mauritia palm swamps, wet savannas, road sides, 500–1300 m; Bolívar (southeastern base of Auyán-tepui, Gran Sabana, Hato Divina Pastora, 20 km east of Icabarú). Apure, Sucre, Táchira; Colombia, Guyana, Suriname, Brazil, Bolivia, Paraguay, Argentina, Uruguay. ◆Fig. 208.

79. SACCIOLEPIS Nash in Britton, Man. Fl. N. States 89. 1901. [Subfamily Panicoideae, Tribe Paniceae] by Emmet J. Judziewicz Terrestrial to aquatic annuals or perennials. Culms hollow, not aerenchymatous. Leaves with ligules membranous, sometimes ciliolate; blades linear. Inflorescence a loosely contracted to densely spicate panicle, the branches strongly appressed to and partially fused with the rachis; pedicels filiform and minutely cupulate; spikelets disarticulating from summit of pedicel or upper floret falling first. Spikelets ovate to lanceolate, usually gibbous and strongly veined, 2-flowered; lower glume present, several-veined; upper glume and lower lemma as long as spikelet, many-veined, one or both gibbous or saccate at the base (or neither saccate in a few species), usually firmly membranous; lower palea present or absent; upper floret often much shorter than the spikelet, elliptic, pale, shining, glabrous or rarely spiculate near the apex, coriaceous. Worldwide in warm-temperate and tropical regions; ca. 30 species, 5 in Venezuela, all in the flora area. Key to the Species of Sacciolepis 1.

1.

2(1).

Spikelets terete, the upper glume and lower lemma delicately membranous, not saccate; lower glume 2/3 as long as to nearly as long as the spikelet ............................................................................... S. angustissima Spikelets somewhat laterally compressed, the upper glume and lower lemma firmly membranous, ± saccate; lower glume about 1/2 as long as the spikelet ............................................................................................. 2 Spikelets 1.3–2 mm long ............................................................................ 3

260

2. 3(2). 3.

4(2).

4.

P OACEAE

Spikelets 2.5–3.7 mm long ......................................................................... 4 Upper glume and lower lemma membranous, green, strongly veined throughout; spikelets elliptic ...................................................... S. myuros Upper glume and lower lemma indurate, straw-colored, conspicuously veined only near the apex, the lower 1/2 veinless; spikelets deltoid ............................................................................................ S. otachyrioides Inflorescence loosely spicate, the lower branches 2–3 cm long; spikelets 3–3.7 mm long, glabrous, the upper glume very strongly saccate ....................................................................................................... S. striata Inflorescence densely spicate, the lower branches < 1 cm long; spikelets 2.5–3.2 mm long, hispid near the apex or uncommonly glabrous, the upper glume moderately saccate ............................................. S. vilvoides

Sacciolepis angustissima (Hochst. ex Steud.) Kuhlm., Comiss. Linhas Telegr. Estratég. Mato Grosso Amazonas, publ. 67, annexo 5, Bot. 11: 92. 1922. —Panicum angustissimum Hochst. ex Steud., Syn. Pl. Glumac. 1: 66. 1855 [1853]. Sacciolepis pungens Swallen, Phytologia 14: 85. 1966. Tufted and decumbent plant, erect portions of culms to 70 cm tall; inflorescence 4– 13 cm long; spikelets 1.3–1.8 mm long, glabrous to pubescent. Wet savannas, Mauritia palm swamps, pond margins, 100–1100 m; Bolívar (Cerro Gavilán, Hato Divina Pastora, La Vergareña, Río Arabapó, Río Marirupa, Río Uruyén, Urimán), Amazonas (Río Orinoco, Río Parucito). Apure, Guárico; Colombia, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia. ◆Fig. 209. Sacciolepis myuros (Lam.) Chase, Proc. Biol. Soc. Wash. 21: 7. 1908. —Panicum myuros Lam., Tabl. Encycl. 1: 172. 1791. Tufted, decumbent plant, the culms 20–90 cm tall; inflorescence (3–)6–20(–32) cm long; spikelets 1.3–2 mm long. Wet savannas, stream banks, marshes, Mauritia palm swamps, near sea level to 400 m; Bolívar (Altiplanicie de Nuria, along Río Orinoco and tributaries), Amazonas (Puerto Ayacucho). Apure, Barinas, Carabobo, Guárico, Monagas, Portuguesa, Táchira; southern Mexico, Central America, Greater Antilles, Colombia, Tri-

nidad-Tobago, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia, Paraguay. Sacciolepis otachyrioides Judz., Syst. Bot. 15: 418. 1990. Tufted, decumbent plant, the culms 30–40 cm tall; inflorescence 5–20 cm long; spikelets 1.5–1.8 mm long. Wet savannas, ca. 100 m; Amazonas (Río Parucito). Apure, Guárico; Amazonian Colombia, Guyana, Brazil, Bolivia. Sacciolepis striata (L.) Nash, Bull. Torrey Bot. Club 30: 383. 1903. —Holcus striatus L., Sp. Pl. 1048. 1753. Sprawling, decumbent and rooting plant, the culms 40–75 cm tall; inflorescence 10–17 cm long. Wet areas, 100–300 m; Delta Amacuro (Serranía de Imataca). Southeastern U.S.A., Mexico, Honduras, Panama, Greater Antilles, Guyana, Suriname, French Guiana, Brazil (Amapá). ◆Fig. 211. Sacciolepis vilvoides (Trin.) Chase, Proc. Biol. Soc. Wash. 21: 7. 1908. —Panicum vilvoides Trin., Gram. Panic. 171. 1826. Tufted, decumbent plant, the culms 30–60 cm tall; inflorescence 12–25 cm long. Wet savannas, near sea level to 50 m; Delta Amacuro (Río Amacuro). Apure, Guárico; Cuba, Guyana, Suriname, French Guiana, Brazil, Bolivia, Paraguay, Argentina, Uruguay. ◆Fig. 210.

80. SCHIZACHYRIUM Nees in Mart. et al., Fl. Bras. Enum. Pl. 2: 331. 1829. [Subfamily Panicoideae, Tribe Andropogoneae] by Emmet J. Judziewicz Annual or perennial savanna grasses. Inflorescences few to usually numerous, both axillary and terminal; individual inflorescences each bearing a solitary

Sacciolepis 261

Fig. 209. Sacciolepis angustissima

Fig. 210. Sacciolepis vilvoides

Fig. 211. Sacciolepis striata

262

P OACEAE

raceme, the subtending spathaceous bract poorly to well developed; racemes of several to many pairs of spikelets, these falling attached to the ± thickened adjacent rachis internode; rachis internodes narrowed at base, usually swollen at the cupulate apex, the margins bearing several irregular lobes; rachis terminating in a triad of 1 sessile and 2 pedicellate spikelets. Sessile spikelets with glumes subequal, as long as and concealing the florets; lower glume membranous, dorsally compressed, nearly flat to strongly convex, entire to bidentate, 2–many-veined, the midvein never prominent, the marginal veins often conspicuous, the margins usually enfolding the upper glume and florets; upper glume membranous to hyaline, 1(3)-veined, strongly keeled; lower floret sterile, represented by a hyaline lemma, the palea absent; upper floret bisexual, the lemma hyaline, strongly bilobed, usually awned, the awn arising from between the teeth on the lemma, strongly geniculate below, kinked shortly after its emergence from the spikelet, the palea absent. Pedicel of pedicellate spikelet similar to rachis internode but more slender; pedicellate spikelet sterile, similar to sessile spikelet or rudimentary and greatly reduced in size. Tropical to temperate areas in New World and Old World; ca. 60 species, 4 in Venezuela, all in the flora area. Key to the Species of Schizachyrium 1.

1.

2(1).

2. 3(2).

3.

Racemes flexuous, the sessile spikelet not squeezed between the divergent internodes and pedicels; racemes numerous (over 100) in a cylindrical panicle 20–35 cm long ............................................. S. condensatum Racemes stiff and straight; sessile spikelet appressed to and/or squeezed between the straight internodes and pedicels; racemes < 25, in a loose panicle < 20 cm long .............................................................................. 2 Delicate annuals or biennials with creeping or decumbent culms; leaf blades 2.5–6(–9) cm long, obtuse or abruptly acute, sometimes bifid; sessile spikelets 3–4 mm long .............................................. S. brevifolium Perennials with erect culms; leaf blades (7–)10–27 cm long, acuminate to acute, never bifid; sessile spikelets 4–8 mm long ................................ 3 Lower glume of sessile spikelet 0.3–0.7 mm wide, inrolled, strongly convex on back, covered with spreading hairs or uncommonly glabrous, straw-colored to purplish, bicarinate, the veins evident only near the apex; sessile spikelets 5–8 mm long; leaf blades V-shaped to flat, 3– 5(–8) mm wide ..................................................................... S. sanguineum Lower glume of sessile spikelet 0.8–1 mm wide, flat on the back or only weakly convex, glabrous, greenish, evidently several-veined throughout; sessile spikelets 4–5.5 mm long; leaf blades V-shaped to involute, 1–2 mm wide .............................................................................. S. tenerum

Schizachyrium brevifolium (Sw.) Nees ex Büse in Miq., Pl. Jungh. 359. 1854. —Andropogon brevifolius Sw., Prodr. 26. 1788. Delicate, straggling annual, the culms branched and decumbent, erect portions 20– 35 cm tall; sessile spikelets 2–2.5 mm long. Mauritia palm swamps, savannas, 100–300

m; Bolívar (Altiplanicie de Nuria, Caño El Garzón, La Vergareña, Serranía de Cerbatana), Amazonas (near Puerto Ayacucho). Widespread elsewhere in Venezuela and tropical regions worldwide. ◆Fig. 213. Schizachyrium condensatum (Kunth) Nees in Mart. et al., Fl. Bras. Enum. Pl.

Schizachyrium 263

2: 333. 1829. —Andropogon condensatus Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 188. 1815 [1816]. —Wana’ ipun kawëmën. Andropogon microstachyum Desv. in Ham., Prodr. Pl. Ind. Occid. 8. 1825. —Schizachyrium microstachyum (Desv.) Roseng. et al., Bol. Univ. Republ. Fac. Agron. Montevideo 103: 35. 1968.

Fig. 212. Schizachyrium condensatum

Cespitose perennial 0.5–1.2 m tall; leaf blades 7–20 × 0.5–0.8 cm; racemes 2–3 cm long; sessile spikelets 3–5 mm long. Savannas, 100–1200 m; Bolívar (widespread), Amazonas (widespread). Widespread elsewhere in Venezuela and in tropical regions worldwide. ◆Fig. 212.

Fig. 213. Schizachyrium brevifolium

264

P OACEAE

Fig. 214. Schizachyrium tenerum

Fig. 215. Schizachyrium sanguineum

Setaria 265

Schizachyrium sanguineum (Retz.) Alston, Handb. Fl. Ceylon 6: 334. 1931. —Rottboellia sanguinea Retz., Observ. Bot. 3: 25. 1791 [1783]. Schizachyrium hirtiflorum Nees in Mart. et al., Fl. Bras. Enum. Pl. 2: 334. 1829. —Andropogon hirtiflorus (Nees) Kunth, Révis. Gramin. 1(suppl.): 39. 1830. Schizachyrium semiberbe Nees in Mart. et al., Fl. Bras. Enum. Pl. 2: 336. 1829. —Andropogon semiberbis (Nees) Kunth, Révis. Gramin. 1(suppl.): 39. 1830. Andropogon riedelii Trin., Mém. Acad. Imp. Sci. St.-Pétersbourg, Sér. 6, Sci. Math. 2: 263. 1832. —Schizachyrium riedelii (Trin.) A. Camus, Ann. Soc. Linn. Lyon 70: 88. 1924. Cespitose perennial 0.5–2 m tall; leaf blades 12–25 cm long; racemes 5–12 cm long; sessile spikelets 5–8 mm long. Savannas,

thickets, 50–1400 m; Bolívar (widespread), Amazonas (widespread). Widespread elsewhere in Venezuela and tropical regions worldwide. ◆Fig. 215. Schizachyrium tenerum Nees in Mart. et al., Fl. Bras. Enum. Pl. 2: 336. 1829. —Andropogon tener (Nees) Kunth, Révis. Gramin. 2: 565, t. 197. 1829 [1832]. Slender, tufted perennial 0.5–1 m tall; leaf blades 7–18 cm long; racemes 3–5 cm long; sessile spikelets 4–5.5 mm long. Savannas, laja margins, 100–1400 m; Bolívar (Gran Sabana, base of Roraima-tepui), Amazonas (Puerto Ayacucho, upper Río Ventuari). Anzoátegui, Aragua, Distrito Federal, Lara, Monagas, Sucre; southern U.S.A., Mexico, Central America, Colombia, Guyana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina, Uruguay. ◆Fig. 214.

81. SETARIA P. Beauv., Ess. Agrostogr. 51, 178. 1812, nom. cons. [Subfamily Panicoideae, Tribe Paniceae] by Emmet J. Judziewicz Annuals or perennials. Leaves with ligules membranous-ciliate; blades linear, flat, parallel-veined or in some species broad, plicate, pseudopetiolate, and obliquely veined. Inflorescence an open to more commonly spicate panicle; spikelets or spikelet clusters subtended by (0)1–many, usually scabrous bristles, the spikelets falling entire from the persistent bristles. Spikelets usually dorsally compressed, plano-convex, 2-flowered; lower glume usually < 1/2 as long as the spikelet, 1–5veined; upper glume 1/2 as long as to about as long as the spikelet, broadly rounded at apex, 5–11-veined; lower floret sterile, staminate, or bisexual, about as long as the spikelet, the lemma membranous to somewhat indurate, 5–7-veined, the palea often well developed; upper floret somewhat shorter than spikelet, typically elliptic and corrugated, granular, or rugose, uncommonly smooth and shining, the margins covering the edges of the palea. Worldwide in temperate and tropical areas; 100–125 species, ca. 10 in Venezuela, 6 of these in the flora area. Key to the Species of Setaria 1. 1. 2(1). 2. 3(2). 3. 4(3).

Spikelets 3.7–4.5 mm long; leaf blades pleated, obliquely veined, 30–50 (–100) mm wide ...................................................................... S. poiretiana Spikelets 1.3–2.8 mm long; leaf blades flat, 3–35 mm wide .................... 2 Spikelets each subtended by 5–10 bristles ................................ S. parviflora Spikelets each subtended by 1 or 0 bristles .............................................. 3 Bristles subtending spikelets both antrorsely and retrorsely barbed ..... 4 Bristles subtending spikelets antrorsely barbed only .............................. 5 Plant annual; leaf blades 3–4 mm wide; spikelets 1.3–1.6 mm long .............................................................................................. S. tenacissima

266

4. 5(3). 5.

P OACEAE

Plant perennial; leaf blades 8–20 mm wide; spikelets 2.2–2.8 mm long ......................................................................................................... S. tenax Inflorescence 5–10(–20) cm long, linear, with bristles 5–10 mm long; leaf blades 3–5(–10) mm wide ................................................. S. macrostachya Inflorescence 23–35 cm long, narrowly elliptic, with bristles 10–20 mm long; leaf blades 20–35 mm wide ............................................. S. vulpiseta

Setaria macrostachya Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 110. 1815 [1816]. Tufted perennial 50–100 cm tall; spikelets ca. 2 mm long. Open areas, 100–500 m; Bolívar (Altiplanicie de Nuria, La Vergareña). Anzoátegui, Aragua, Distrito Federal, Falcón, Lara, Miranda, Zulia; southwestern U.S.A., southern Mexico, West Indies, Trinidad-Tobago. Setaria parviflora (Poir.) Kerguélen, Lejeunea n.s. 120: 161. 1987. —Cenchrus parviflorus Poir. in Lam., Tabl. Encycl. 6: 52. 1804. —Paja cepillo. Panicum geniculatum Willd., Enum. Pl. 1031. 1809, non Lam. 1798. —Setaria geniculata (Willd.) P. Beauv., Ess. Agrostogr. 51, 169, 178. 1812, nom. illeg. Setaria gracilis Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 109. 1815 [1816]. Tufted perennial 15–75 cm tall, culms also decumbent and rooting; inflorescence 2–10 cm long, yellowish orange; spikelets 1.8–2.8 mm long, subtended by 5–10 antrorsely scabrous bristles. Pastures and other disturbed open areas, savannas, 100–900 m; widespread in Delta Amacuro, Bolívar, and Amazonas. Widespread elsewhere in Venezuela; eastern U.S.A., Mexico, Central America, West Indies, tropical and temperate South America, widely naturalized throughout the Old World. Setaria poiretiana (Schult.) Kunth, Révis. Gramin. 1: 47. 1829. —Panicum poiretianum Schult., Mantissa 2: 229. 1824. —Pegadera. Cespitose perennial 75–200 cm tall; leaf blades 25–60 cm long; inflorescence 30–60 cm long, loosely contracted, the lowest branches to 6 cm long. River sides, near sea level to 1000 m; Delta Amacuro (Tucupita),

Bolívar (20 km east of La Paragua, Río Icutu). Anzoátegui, Aragua, Distrito Federal, Miranda, Sucre, Zulia; southern Mexico, Central America, Lesser Antilles, Colombia, Trinidad-Tobago, Guyana, Suriname, French Guiana, Brazil, Bolivia. Setaria tenacissima Schrad. ex Schult., Mantissa 2: 279. 1824. Tufted annual 50 cm tall; inflorescence 3– 5 cm long, densely spicate. Open areas, ca. 1000 m; Bolívar (Santa Elena de Uairén). Aragua; Mexico, Guatemala, Belize, Honduras, El Salvador, Costa Rica, Panama, Greater Antilles, Colombia, Trinidad-Tobago, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 217. Setaria tenax (Rich.) Desv., Mem. Soc. Agric. Angers 1: 182. 1831. —Panicum tenax Rich., Actes Soc. Hist. Nat. Paris 1: 106. 1792. —Paja imán, Wana’ ipun. Cespitose perennial 50–100 cm tall; leaf blades 17–30 cm long; inflorescence 7–20 cm long, densely spicate. Savannas, often on rocky soil, 50–500 m; Bolívar (occasional in Río Orinoco basin), Amazonas (Puerto Ayacucho). Widespread elsewhere in Venezuela; Neotropics. ◆Fig. 216. Setaria vulpiseta (Lam.) Roem. & Schult., Syst. Veg. 2: 495. 1817. —Panicum vulpisetum Lam., Encycl. 4: 735. 1798. Cespitose perennial 50–150 cm tall; spikelets 2–2.2 mm long. Open areas, 100–500 m; Bolívar (between Encrucijada and El Pao, headwaters of Río Saca, San Félix, Santa María del Vapor). Anzoátegui, Aragua, Barinas, Carabobo, Distrito Federal, Miranda, Portuguesa, Sucre, Yaracuy, Zulia; Neotropics. ◆Fig. 218. Setaria magna Griseb., a closely related species with smooth upper florets, may eventually be found in the flora area.

Setaria 267

Fig. 216. Setaria tenax

Fig. 217. Setaria tenacissima

268

P OACEAE

Fig. 218. Setaria vulpiseta

Sorghum 269

82. SORGHASTRUM Nash in Britton, Man. Fl. N. States 71. 1901. [Subfamily Panicoideae, Tribe Andropogoneae] by Emmet J. Judziewicz Annuals or more commonly robust perennials. Inflorescence paniculate, often bushy, of numerous raceme-like branches, these sparingly rebranched; branches with nodes bearing a perfect sessile spikelet and a cupulate, hairy, sterile pedicel (2 at the apex of the branches); disarticulation at the base of each internode, the sessile spikelet falling with its sterile pedicel and the attached internode above it. Sessile spikelets ovate-lanceolate, dorsally compressed, the callus bearded, obtuse to pungent; glumes as long as spikelet, coriaceous, concealing the much smaller hyaline florets; lower glume 7–9-veined, pubescent, somewhat enfolding the 5veined, glabrous upper glume; lower floret sterile, the lemma veinless, bifid, awnless, ciliate above; upper floret bisexual, the lemma ciliate, 3-veined, bifid, short- to long-awned from between the teeth, the awn straight to geniculate. Tropical and temperate regions worldwide; 17 species, 3 in Venezuela, 1 of these in the flora area. Sorghastrum setosum (Griseb.) Hitchc., Contr. U.S. Natl. Herb. 12: 195. 1909. —Andropogon setosus Griseb., Cat. Pl. Cub. 235. 1866. Sorghum parviflorum Desv. in Ham., Prodr. Pl. Ind. Occid. 12. 1825. —Sorghastrum parviflorum (Desv.) Hitchc. & Chase, Contr. U.S. Natl. Herb. 18: 287. 1917. Cespitose perennial 1–2 m tall; leaves often bluish glaucous, the blades 25–50 × 0.4– 0.8 cm; inflorescence 15–35 × 3–5 cm; sessile

spikelets 3.5–4.5 mm, the upper floret awnless or with awn to 4 mm long. Savannas, Mauritia palm swamps, river banks, sometimes dominant, 100–900 m; Bolívar (16 km north of El Manteco, Gran Sabana), Amazonas (Isla Carestía in Río Orinoco, Guapachana, Puerto Ayacucho, San Juan de Manapiare). Apure, Barinas, Guárico, Monagas, Portuguesa; Mexico, Central America, Greater Antilles, Colombia, Guyana, French Guiana, Brazil, Bolivia, Paraguay, Argentina, Uruguay. ◆Fig. 219.

83. SORGHUM Moench, Methodus 207. 1794, nom. cons. [Subfamily Panicoideae, Tribe Andropogoneae] by Emmet J. Judziewicz Cespitose to rhizomatous annuals or perennials. Foliage often coarse, maizelike. Inflorescence an open to contracted panicle of numerous, divergent racemes; racemes with spikelets paired, the sessile spikelet bisexual, the pedicellate spikelet staminate or sterile; apex of racemes bearing a triad of 1 sessile and 2 pedicellate spikelets; rachis internodes slender, disarticulating at maturity along with the persistent spikelets or the sessile spikelet falling first. Sessile spikelet plump, terete, the glumes lanceolate-ovate to nearly round, glabrous to ± pubescent, shiny, coriaceous, conspicuously 5–9-veined adaxially, the veins not evident abaxially except near the apex; margins of lower glume inflexed over the edges of the upper glume; florets much shorter than the spikelet, with well-developed, hyaline lemmas; lower floret sterile, lacking a palea; upper floret bisexual, the lemma bilobed, a delicate, geniculate awn often arising from the sinus, the palea rudimentary. Pedicellate spikelet narrowly lanceolate, membranous, awnless; glumes strongly 5–7-veined, the lower glume often flat or slightly sulcate on the back; upper floret staminate. Old World tropics and subtropics; ca. 20 species, 2 cultivated and naturalized worldwide, including Venezuela; 1 of these in the flora area.

270

P OACEAE

Fig. 219. Sorghastrum setosum

Sorghum 271

Fig. 220. Sorghum bicolor

272

P OACEAE

Sorghum bicolor (L.) Moench, Methodus 207. 1794. —Holcus bicolor L., Mant. Pl. 2: 301. 1771. Holcus sorghum L., Sp. Pl. 1047. 1753. —Andropogon sorghum (L.) Brot., Fl. Lusit. 1: 88. 1804. —Sorghum vulgare Pers., Syn. Pl. 1: 101. 1805. Stout, maize-like annuals, the culms 1–3 m tall; inflorescence 15–40 cm, pyramidal,

open, loosely flowered; sessile spikelets 3–7 mm long, ovate to roundish. Weedy places, 50–100 m; Amazonas (Puerto Ayacucho, Río Ventuari). Widespread elsewhere in Venezuela and throughout tropical and subtropical regions, native of subtropical Africa. ◆Fig. 220. Sorghum bicolor is cultivated worldwide for grain, molasses, and forage.

Fig. 221. Spartina alterniflora

Sporobolus 273

84. SPARTINA Schreb., Gen. Pl. 1: 43. 1789. [Subfamily Chloridoideae, Tribe Cynodonteae] by Emmet J. Judziewicz Cespitose to more commonly strongly rhizomatous perennials. Culms often over 1 m tall. Foliage coarse, the ligules ciliate, the blades linear, elongate, flat or inrolled. Inflorescence terminal, a panicle consisting of few to numerous ascending spikes; spikes densely flowered, the strongly overlapping spikelets borne in rows along 2 sides of the triquetrous rachis. Spikelets 1-flowered, strongly laterally compressed, firmly membranous, awnless, disarticulating entire; glumes lanceolate, unequal, the lower 1-veined, short, the upper 3–5-veined, as long as or longer than the floret; floret with lemma slightly shorter than the palea, 1–3-veined, the palea membranous, obscurely 2-keeled; stamens 3; stigmas 2, plumose. Near the shores of the Atlantic Ocean in North America, Mexico, Central America, West Indies, South America, and Europe, also inland in North America; 14 species, 1 in Venezuela. Spartina alterniflora Loisel., Fl. Gall. 719. 1807. Spartina brasiliensis Raddi, Agrostogr. Bras. 21. 1823. Coarse, strongly rhizomatous perennial, the culms to 1 m tall; inflorescence 10–25 cm long, of few, ascending racemes 4–8 cm long;

spikelets 10–15 mm long, straw-colored. Tidal mud flats, muddy lagoons, and estuaries near the ocean; Delta Amacuro (Caño Mariusa). Eastern Canada, eastern U.S.A., Trinidad-Tobago, Guyana, Suriname, French Guiana, eastern Brazil, eastern Argentina. ◆Fig. 221.

85. SPOROBOLUS R. Br., Prodr. 169. 1810. [Subfamily Chloridoideae, Tribe Cynodonteae] Agrosticula Raddi, Fl. Bras. Enum. Pl. 33. 1823. Cryptostachys Steud., Syn. Pl. Glumac. 1: 181. 1855 [1854]. Diachyrium Griseb., Abh. Königl. Ges. Wiss. Göttingen 19: 257, t. 2, fig. 8. 1874. by Gerrit Davidse Annuals or perennials, cespitose or rhizomatous. Sheaths rounded or keeled; ligule a row of hairs; blades flat to convolute, rarely cylindrical. Inflorescence a spicate to open panicle. Spikelets small, rounded or laterally compressed, 1-flowered, bisexual; disarticulating above the glumes or remaining intact until extrusion of the seed; glumes equal or unequal, membranous or hyaline, 1-veined or veinless, sometimes with faint lateral veins, the lower shorter than the spikelet, the upper shorter to as long as the spikelet; lemma 1-veined, membranous, awnless; palea nearly as long as the lemma, often splitting between the veins as the caryopsis matures; lodicules 2; stamens 1–3; styles 2. Caryopsis utricle-like with the mature pericarp usually gelatinizing when wetted and extruding the naked seed; embryo ca. 1/2 as long as the utricle; hilum punctate. Tropics and subtropics, a few species in temperate areas; ca. 160 species, 8 in Venezuela, 3 of these in the flora area. Not yet collected in the flora area are two weedy species and a coastal dune species. The weedy Sporobolus pyramidatus (Lam.) Hitchc. should be expected in Delta Amacuro in areas with saline soils. Important diagnostic characters are: panicle branches whorled; spikelets 1.4–1.7 mm long; base of plant decumbent,

274

P OACEAE

green to straw-colored; inflorescences 3–7 cm long; leaf blades 2–7 cm long. The weedy S. indicus (L.) R. Br. is very similar to S. jacquemontii but has more appressed panicle branches and slightly larger spikelets; it could be expected in weedy, open areas at middle elevations, such as the Gran Sabana. The coastal S. virginicus (L.) Kunth is to be expected from the northern part of Delta Amacuro. Important diagnostic characters are: panicle branches solitary or paired; lower glume 1.6–1.8 mm long; plants strongly rhizomatous; inflorescence 3–10 cm long, narrow and compact; spikelets straw-colored. Key to the Species of Sporobolus 1. 1. 2(1). 2.

Spikelets 0.7–0.9(–10) mm long .............................................. S. tenuissimus Spikelets 1.3–4 mm long. ........................................................................... 2 Lowest panicle branches whorled ................................................. S. cubensis Lowest panicle branches solitary or paired ........................... S. jacquemontii

Sporobolus cubensis Hitchc., Contr. U.S. Natl. Herb. 12: 237. 1909. Perennial 50–70 cm tall, with deeply buried, bulbous bases. Savannas, 50–900 m; widespread in Bolívar and Amazonas. Anzoátegui, Apure, Aragua, Barinas, Falcón, Guárico, Mérida, Monagas, Portuguesa, Sucre; Mexico, Belize, Honduras, Nicaragua, Costa Rica, Panama, Caribbean, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 222. Sporobolus jacquemontii Kunth, Révis. Gram. 2: 427, t. 127. 1831. —Vilfa jacquemontii (Kunth) Trin., Mém. Acad. Imp. Sci. St.-Pétersbourg, Sér. 6, Sci. Math., Seconde Pt. Sci. Nat. 6, 4(1–2): 92. 1840. —Sporobolus pyramidalis var. jacquemontii (Kunth) Jovet & Guédès, Taxon 22: 163. 1973. Sporobolus pyramidalis P. Beauv., Fl. Oware 2: 36, t. 80. 1816. —Vilfa pyramidalis (P. Beauv.) Trin. ex Steud., Nomencl. Bot. ed. 2, 2: 768. 1843. —Sporobolus indicus var. pyramidalis (P. Beauv.) Veldkamp, Blumea 35(2): 439. 1991. Weedy perennial 40–110 cm tall. Roadsides and other open secondary vegetation, 50–600 m; widespread in Bolívar and Amazonas. Anzoátegui, Apure, Aragua, Barinas, Carabobo, Cojedes, Distrito Federal, Falcón, Guárico, Lara, Mérida, Miranda, Monagas, Nueva Esparta, Portuguesa, Sucre, Táchira, Trujillo, Yaracuy, Zulia; U.S.A., Mexico, Caribbean, Guatemala, Belize, Honduras, Nica-

ragua, Costa Rica, Panama, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. Sporobolus tenuissimus (Mart. ex Schrank) Kuntze, Revis. Gen. Pl. 3(2): 369. 1898. —Panicum tenuissimum Mart. ex Schrank, Denkschr. Königl.Baier. Bot. Ges. Regensburg 2: 26. 1822. Agrosticula muralis Raddi, Agrostogr. Bras. 33, t. 1, fig. 2. 1823. —Sporobolus muralis (Raddi) Hitchc. & Chase, Contr. U.S. Natl. Herb. 18: 368. 1917. Vilfa minutiflora Trin., Gram. Unifl. Sesquifl. 158. 1824. —Sporobolus minutiflorus (Trin.) Link, Hort. Berol. 1: 88. 1827. —Agrostis minutiflora (Trin.) Desf. ex Steud., Nomencl. Bot. 1: 41. 1840. Sporobolus effusus Desv., Mem. Soc. Agric. Angers 1: 164. 1831. Weedy annual 15–90 cm tall with delicate, open panicles. Roadsides and other open secondary vegetation, 100–200 m; Bolívar (San Martín de Turumbán southwest of Tumeremo). Aragua, Cojedes, Distrito Federal, Falcón, Guárico, Miranda, Monagas, Sucre, Yaracuy; U.S.A., Mexico, Belize, Honduras, El Salvador, Costa Rica, Panama, Caribbean, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Paraguay. ◆Fig. 223. This is apparently a recent introduction (1979) into the flora area. This is not surprising since Sporobolus tenuissimus is in the process of spreading throughout tropical America.

Sporobolus 275

Fig. 222. Sporobolus cubensis

Fig. 223. Sporobolus tenuissimus

276

P OACEAE

86. STEINCHISMA Raf., Bull. Bot., Geneva 1: 220. 1830. [Subfamily Panicoideae, Tribe Paniceae] by Fernando O. Zuloaga Cespitose perennial, short- or long-rhizomatose; internodes hollow. Ligules membranous, shortly ciliate or laciniate at the apex; blades lanceolate to filiform, flat or inrolled. Inflorescence an open to contracted panicle; spikelets disarticulating entire from the pedicels. Spikelets ellipsoid to long-ellipsoid, dorsally compressed, 2-flowered, glabrous; lower glume 3-veined, 1/3–1/2 the length of the spikelet; upper glume and lower lemma 3–5(–7)-veined, subequal; lower flower staminate or neuter; lower palea conspicuous, expanded at maturity; upper anthecium ovoid to ellipsoid, with verrucose papillae regularly distributed over the lemma and palea, and with or without prickle hairs at the apex of the lemma; lodicules 2, truncate; stamens 2 or 3; stigmas 2, plumose. Caryopsis with an oblong hilum; embryo 1/2 or less the length of the caryopsis. Southern U.S.A., Mexico, Central America, West Indies, most of South America; 7 species, 3 in Venezuela, all in the flora area. Key to the Species of Steinchisma 1. 1. 2(1). 2.

Upper flower with 2 stamens; spikelets 1–1.7 mm long ..................... S. laxa Upper flower with 3 stamens; spikelets 1.8–3 mm long .......................... 2 Panicles contracted, spiciform; blades narrowly lanceolate, 5–7 mm wide ................................................................................................... S. decipiens Panicles open; blades filiform, 2–5(–10) mm wide .................. S. stenophylla

Steinchisma decipiens (Nees ex Trin.) W.V. Br., Mem. Torrey Bot. Club 23: 20. 1977. —Panicum decipiens Nees ex Trin., Gram. Panic. 227. 1826. Rhizomatous perennial, 15–60 cm tall; panicles contracted, spiciform; spikelets glabrous, with upper glume and lower lemma 5veined, lower palea expanded at maturity, upper floret papillose. In wet, open places, on sandy soils, ca. 800 m; Bolívar (Hato Divina Pastora north of Santa Elena de Uairén). Colombia, northern Brazil, Bolivia, Paraguay, Argentina, Uruguay. Steinchisma laxa (Sw.) Zuloaga, Amer. J. Bot. 90: 817. 2003. —Panicum laxum Sw., Prodr. 23. 1788. Panicum luticola Hitchc., Contr. U.S. Natl. Herb. 22: 485, fig. 82. 1922. Panicum caroniense Luces, Bol. Soc. Venez. Ci. Nat. 15: 26, fig. 12. 1953. Perennial, culms decumbent and rooting at the lower nodes, 15–80 cm tall; spikelets unilaterally disposed on the branches, ellip-

tic, 1–1.7 mm long, pilose or glabrous, lower palea conspicuous, upper glume and lower lemma 5-veined. Common in wet and disturbed places, 100–900 m; scattered in Bolívar and Amazonas. Anzoátegui, Apure, Barinas, Cojedes, Falcón, Guárico, Lara, Miranda, Monagas, Portuguesa, Sucre, Táchira, Yaracuy, Zulia; Mexico, Central America, West Indies, most of South America. Steinchisma stenophylla (Hack.) Zuloaga & Morrone, Ann. Missouri Bot. Gard. 85: 651. 1998. —Panicum stenophyllum Hack., Oesterr. Bot. Z. 51: 371. 1901. Short-rhizomatous perennial, culms erect, 30 cm tall, branching at the upper nodes; blades filiform, 2–5(–10) mm wide; spikelets narrowly elliptic, 2.4–3 mm long, lower palea expanded at maturity. River banks, in sandy soils, 100–1100 m; Amazonas (Cerro Parú, Río Coro Coro, Río Cunucunuma). Suriname, northern Brazil. ◆Fig. 224.

Steirachne 277

Fig. 224. Steinchisma stenophylla

Fig. 225. Steirachne barbata

278

P OACEAE

87. STEIRACHNE Ekman, Ark. Bot. 10(17): 35. 1911. [Subfamily Chloridoideae, Tribe Cynodonteae] by Emmet J. Judziewicz Cespitose perennials. Culms with bases sometimes slightly cormose; cleistogamous spikelets often present within the lower leaf sheaths near the plant base. Leaves with ligules minute, ciliolate; blades linear. Inflorescence (besides cleistogamous ones) a terminal, open panicle. Spikelets laterally compressed, several- to many-flowered, the florets not strongly overlapping and thus the rachilla internodes usually visible, disarticulating above the persistent glumes and between the florets; glumes subulate, narrowly lanceolate, unequal, 1-veined, shorter than the lowest floret; florets with lemmas lanceolate, attenuate, 3-veined, the lateral veins parallel to the midvein; paleas shorter than the lemmas; stamens 2. Venezuela, Guyana, northeastern Brazil; 2 species, 1 in Venezuela. Steirachne barbata (Trin.) Renvoize, Kew Bull. 39: 184. 1984. —Eragrostis barbata Trin., Mém. Acad. Imp. Sci. St.Pétersbourg, Sér. 6, Sci. Math., Seconde Pt. Sci. Nat. 4(2): 76. 1836. Tufted perennial 30–60 cm tall; inflores-

cence 4–10 cm long; spikelets 6–10 mm long, purplish, elliptic, 7–13-flowered, the florets 3–4 mm long. Open, seasonally flooded savannas, 50–100 m; Bolívar (18 km southeast of Caicara). Apure, Guárico; Guyana, northeastern Brazil. ◆Fig. 225.

88. STEYERMARKOCHLOA Davidse & R.P. Ellis, Ann. Missouri Bot. Gard. 71: 995, fig. 1–27. 1984 [1985]. [Subfamily Arundinoideae, Tribe Steyermarkochloeae] by Gerrit Davidse Perennial, with dimorphic culms and leaves. Developed leaf solitary on each vegetative culm; sheath solid, cylindrical; blade flattened; ligule not differentiated. Inflorescence spicate, cylindrical with staminate and/or bisexual spikelets lowermost and pistillate spikelets uppermost. Spikelets solitary, 3-flowered, usually unisexual, sometimes bisexual, dorsally compressed; disarticulation below the glumes; glumes 2; uppermost floret rudimentary; lower floret of the pistillate spikelet sterile; palea of the pistillate floret spongy, curved, (5–)7–11-veined, longer than the lemma; lodicules 0; stigmas 2; style 1; caryopsis fusiform, the hilum linear. Colombia, Venezuela, Brazil; 1 species. Steyermarkochloa angustifolia (Spreng.) Judz., Ann. Missouri Bot. Gard. 77: 204. 1990. —Pariana angustifolia Spreng., Syst. Veg. 2: 609. 1825. Steyermarkochloa unifolia Davidse & R.P. Ellis, Ann. Missouri Bot. Gard. 71: 995, fig. 1–27. 1984 [1985]. Perennial 1–3 m tall. Margins of streams

in seasonally water-logged soils or flooded white-sand savannas, 100–200 m; Amazonas (Caño Caname, Caño Pimichín, Caño San Miguel, Caño Ucata southeast of Síquita, Caño Yagua, base of Cerro Yapacana, Río Atacavi, Río Atabapo, Río Guasacavi, Río Temi). Colombia (Guaínia), Brazil (Amazonas). ◆Fig. 226.

Steyermarkochloa 279

Fig. 226. Steyermarkochloa angustifolia

280

P OACEAE

89. STREPTOGYNA P. Beauv., Ess. Agrostogr. 80. 1812. [Subfamily Bambusoideae, Tribe Streptogyneae] by Emmet J. Judziewicz Rhizomatous or cespitose perennials. Culms solid, unbranched. Leaves with inner ligules membranous; outer ligules short, indurate; blades deciduous above the outer ligule, linear to lanceolate, narrowed into a short pseudopetiole. Inflorescences terminal spike-like panicles, the rachis triquetrous, 2 sides alternately bearing the spikelet pedicels. Spikelets short-pedicellate, linear-lanceolate, severalflowered, greenish, the lower florets functional, the upper ones sterile, disarticulating between the functional florets, each of which falls attached to the extended, curved, hook-like rachilla segment above it; glumes membranous, persistent, manyveined, unequal, the upper convolute and enclosing the lowest floret at the base; lemmas slender, elongate, long-awned, inrolled, indurate, the base extended beyond the attachment of the floret into a sharp, oblique callus; paleas deeply sulcate; lodicules 3; stamens 2; stigmas 2 or 3, hispid, at maturity becoming hardened and persistent, intertwined at maturity with the stigmas of other florets in the same and adjacent spikelets. Caryopsis linear. Tropical America, tropical Africa, southern India, Sri Lanka; 2 species, 1 in Venezuela. The needle-like spikelets are adapted for external animal dispersal. Streptogyna americana C.E. Hubb. in Hook., Icon. Pl. 36: 5, t. 3572, fig. 1. 1956. —Kabadi-wochi, Paja cortadera. Cespitose perennial 0.5–1 m tall; leaves mostly basal, the blades 40–60 × 1–1.5 cm; inflorescence 20–30 cm long, arching; spikelets 20–25 mm long, 4–6-flowered. Moist to wet forests, 100–200 m; Delta Amacuro

(Piacoa, Río Cuyubini), Bolívar (28 km south of El Dorado, southwest of El Manteco, Río Aro, Río Asa, Río Caura), Amazonas (Río Cunucunuma, Río Orinoco). Sucre; southern Mexico, Central America, Colombia, Trinidad-Tobago, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia. ◆Fig. 227.

90. STREPTOSTACHYS Desv., Nouv. Bull. Sci. Soc. Philom. Paris 2: 190. 1810. [Subfamily Panicoideae, Tribe Paniceae] by Emmet J. Judziewicz Cespitose or short-rhizomatous perennials, the culms hollow. Leaves with ligules ciliate or absent; blades ovate-lanceolate, subcordate to strongly clasping the culm. Panicles lax, ± open, the spikelets borne in loose racemes. Spikelets disarticulating below the glumes, oblong, somewhat dorsally compressed, 2-flowered; rachilla thickened and corky between the glumes; lower glume much shorter than to nearly as long as the spikelet, 1–several-veined; upper glume as long as the spikelet, 5–7-veined; lower floret usually sterile, occasionally staminate, the lemma as long as the spikelet, 3–5-veined, the palea usually rudimentary if present; upper floret elliptic, pale, indurate, shining, papillose, often also pilose, the lemma with margins enclosing the palea. Caryopsis with hilum linear. Venezuela, Trinidad-Tobago, Guyana, Suriname, French Guiana, Brazil, Paraguay; 4 species, 1 in Venezuela.

Streptogyna 281

Fig. 227. Streptogyna americana

282

P OACEAE

Streptostachys asperifolia Desv., Nouv. Bull. Sci. Soc. Philom. Paris 2: 190. 1810. —Panicum asperifolium (Desv.) Hitchc., Contr. U.S. Natl. Herb. 22: 489. 1922. —Paja macaguera. Loosely cespitose perennial 50–80 cm tall; leaf blades 15–25 × 1.8–3.5 cm, clasping basally; inflorescence 10–23 cm long, with few, stiff, divergent branches to 5 cm long; spikelets 3.8–4.5 mm long, narrowly obovate, pubescent, the base toroidally prolonged around the summit of the pedicel. Forest edges, edges of Mauritia palm swamps, near sea level to 300 m; Bolívar (Río Orinoco basin west to Río Caura). Anzoátegui, Monagas, Sucre; Trinidad-Tobago, Guyana, Suriname, French Guiana, northeastern Brazil, Bolivia. ◆Fig. 228.

Fig. 228. Streptostachys asperifolia

91. THRASYA Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 120. 1815 [1816]. [Subfamily Panicoideae, Tribe Paniceae] by Emmet J. Judziewicz Annuals or perennials. Leaves with ligules membranous; blades linear. Inflorescences 1–several from the uppermost nodes, each of 1–several straight to arcuate raceme(s); racemes with rachis flattened or V-shaped, the membranous margins often embracing the spikelets; spikelet pedicels paired, borne in 1 or 2 rows, the spike-

Thrasya 283

lets arranged so that the lower lemmas face each other. Spikelets disarticulating below the glumes, elliptic to lanceolate, ± terete, 2-flowered; lower glume absent to 1/2 as long as the spikelet; upper glume 1/3 to nearly as long as spikelet (rarely absent), (1–)3–9-veined, in some species reticulate in the upper portion and with the veins excurrent as mucros; lower floret staminate or sterile; lower lemma membranous and flat to more typically indurate and sulcate, if sulcate then often splitting down the obscure midvein; lower palea absent to well developed; upper floret bisexual, slightly shorter than spikelet; lemma ovate to lanceolate, coriaceous, whitish, usually finely papillose, uncommonly pitted, glabrous or sometimes pubescent at the apex or on back; palea of similar texture, enfolded by the margins of the lemma; stamens 2 or 3; stigmas 2, pilose. Southern Mexico, Central America, Colombia, Venezuela, Trindad-Tobago, Guyana, Suriname, French Guiana, Brazil, Bolivia, Paraguay, Argentina; ca. 25 species, 10 in Venezuela, 6 of these in the flora area. Key to the Species of Thrasya 1. 1. 2(1). 2. 3(2). 3. 4(3). 4. 5(4).

5.

Spikelets 0.9–1.1 mm long; both glumes absent; upper floret minutely pitted .......................................................................................... T. axillaris Spikelets 2.2–5.3 mm long; at least upper glume present; upper floret minutely papillose ................................................................................. 2 Rachis of raceme(s) 2.5–6 mm wide when spread .......................... T. petrosa Rachis of raceme(s) 1–2 mm wide when spread ....................................... 3 Upper glume 0.5–1 mm long, narrowly triangular, at most 1/3 the length of the upper floret; spikelets 2.5–2.7 mm long ............................. T. stricta Upper glume 1.5–4 mm long, lanceolate-ovate, > 1/2 as long as the upper floret; spikelets 2.2–4.8 mm long .......................................................... 4 Raceme(s) 15–25 cm long; lower lemma sulcate but not splitting ...................................................................................................... T. robusta Raceme(s) 3–10 cm long; lower lemma sulcate and readily splitting ...... 5 Upper glume distinctly longer than the upper floret, acute, often with stiff, divergent whitish to commonly golden, tuberculate cilia all across the back ................................................................................. T. paspaloides Upper glume distinctly shorter than to as long as the upper floret, obtuse to emarginate, glabrous or with appressed whitish hairs on both sides of the midrib ............................................................................ T. trinitensis

Thrasya axillaris (Swallen) A.G. Burm. in Görts, Fl. Guianas, Fam. 187 (Poaceae): 630. 1991. —Paspalum axillare Swallen, Bull. Torrey Bot. Club 75: 84. 1948. Delicate straggling annual, the culms to 15 cm tall; raceme solitary, divergent, 1.5–3.5 cm long; spikelets glabrous, the lower floret sterile, consisting only of a flattened lemma. Sprays of waterfalls, soil pockets on cliffs, 400–1400 m; Bolívar (Cerro Venado 20 km east of Canaima, 58 km west of Santa Elena de Uairén), Amazonas (Sierra de la Neblina). Suriname, Brazil (Amazonas). ◆Fig. 230.

Thrasya paspaloides Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 121, t. 39. 1815 [1816]. Thrasya setosa Swallen, Fieldiana, Bot. 28: 17. 1951. Cespitose perennial to 50 cm tall, forming dense colonies; raceme(s) 1(–3), each 3–10 cm long, the rachis 1–2 mm wide; spikelets 2.7–4 mm long; lower glume absent; lower floret often with a basal tuft of hairs; upper floret ca. 2 mm long, short-ciliate at the apex. Moist places in savannas and lajas, 100–200 m; Bolívar (Isla El Gallo in Río Orinoco),

284

P OACEAE

Fig. 229. Thrasya petrosa

Fig. 230. Thrasya axillaris

Thrasya 285

“setosa” variation

Fig. 231. Thrasya paspaloides

286

P OACEAE

Fig. 232. Thrasya trinitensis

Fig. 233. Thrasya stricta

Trachypogon 287

Amazonas (common along Río Orinoco). Endemic. ◆Fig. 231. Thrasya paspaloides forms a difficult complex with T. trinitensis and T. thrasyoides (Trin.) Chase. Thrasya petrosa (Trin.) Chase, Proc. Biol. Soc. Wash. 24: 115. 1911. —Panicum petrosum Trin., Sp. Gram. 3: pl. 280. 1831. —Sádu. Cespitose perennial 75–150 cm tall; raceme(s) 1(–3), each 12–23 cm long; spikelets 3.7–5.3 mm long, on a swollen, ca. 0.5 mm long, oil-bearing callus. Wet to dry savannas and lajas, sometimes dominant, often on termite mounds (ant-dispersed), 100–1100 m; Bolívar (widespread), Amazonas (widespread). Throughout the rest of Venezuela; Mexico, Guatemala, Costa Rica, Panama, Colombia, Guyana, Suriname, Peru, Brazil, Bolivia, Paraguay. ◆Fig. 229. Thrasya robusta Hitchc. & Chase, Contr. U.S. Natl. Herb. 18: 297. 1917. —Wiri-yu. Cespitose perennial to 1 m tall; leaf blades 20–27 × 1 cm; raceme(s) 1 or 2, each 15–25 cm long; rachis 1.5–2 mm wide (spread width); spikelets 3.7–4.8 mm long, elliptic, densely pubescent. Savannas, 500–1400 m;

Bolívar (between Piedra de la Virgen and La Escalera). Costa Rica, Panama, Colombia, Trinidad-Tobago, Suriname, French Guiana. Thrasya stricta A.G. Burm., Acta Bot. Venez. 14(4): 75. 1987. Plant probably perennial, decumbent, rooting, and forming dense colonies, erect portions to 50 cm tall; raceme solitary, ca. 10 cm long, the rachis 1 mm wide; lower glume absent; lower lemma readily splitting down the middle. Granitic domes, ca. 100 m; Amazonas (Laja Grande, near Puerto Ayacucho). Endemic. ◆Fig. 233. Thrasya trinitensis Mez, Repert. Spec. Nov. Regni Veg. 15: 125. 1918. Cespitose perennial 30–50 cm tall; foliage densely pubescent; raceme(s) 1(2), each 3–8 cm long, the rachis 1–2 mm wide; spikelets 2.2–3 mm long; lower glume often present, cuff-like. Savannas, marshes, granitic domes, Mauritia palm swamps, open slopes of tepuis, sometimes dominant, 100–1800 m; Bolívar (widespread), Amazonas (widespread). Anzoátegui, Apure; Belize, Honduras, El Salvador, Nicaragua, Costa Rica, Colombia, Trinidad-Tobago, Guyana, Brazil, Bolivia. ◆Fig. 232.

92. TRACHYPOGON Nees in Mart. et al., Fl. Bras. Enum. Pl. 341. 1829. [Subfamily Panicoideae, Tribe Andropogoneae] Homopogon Stapf in A. Chevalier, Mém. Soc. Bot. France 8(b): 103. 1908. by Gerrit Davidse Plants perennial, cespitose or with short rhizomes. Culms unbranched, solid. Leaf sheaths rounded; ligule a membrane adnate to the margins of the sheath; blades linear, usually flat, sometimes convolute. Inflorescences terminal, solitary, of 1 solitary raceme or several digitate racemes; rachis persistent after maturity of the spikelets, bearing the old staminate spikelets; internodes of the rachis and spikelet pedicels linear. Spikelets paired, dimorphic, 2-flowered, unequally pedicelled, one of each pair short-pedicellate, persistent, awnless, staminate, dorsally compressed; other spikelet of each pair longer pedicelled, deciduous, bisexual, awned, nearly terete. Bisexual spikelets deciduous at maturity, falling with an oblique, hairy, pointed callus formed of the apical portion of the pedicel; lower glume coriaceous, narrowly elliptical, truncate, its margins inrolled over the second glume and mostly concealing it; upper glume about as long as the lower, somewhat keeled, 3-veined, the apex clasping the base of the awn; lower floret sterile; lower lemma hyaline, 2-veined, grooved between the keels, the margins inflexed; lower palea absent; upper floret bisexual; upper lemma narrow, firm except at the hyaline base, flattened, tapering directly into the base of the awn, the awn twisted, hispid, twice geniculate, well exserted; upper palea absent; stamens 3; styles 2, separate. Staminate spikelets with

288

P OACEAE

the lower glume dorsally flattened, rounded on the back, 5–11-veined, the margins sharply inflexed and covering the margins of a membranous lower glume of about equal length; lower floret sterile; upper floret staminate; lower and upper lemma about equal, hyaline; lower and upper paleas absent; lodicules 2; anthers 3. Tropics and subtropics of America and Africa; 2–6 species, 1 or 3 in Venezuela, 1 in the flora area. Although this is a small, distinctive genus, the taxonomy is very complex with a bewildering array of intergrading forms. The perennial species are the dominants of large areas of savannas, especially in the American tropics. Polyploidy occurs on both continents, although the lowland populations of the Americas are mostly diploid (personal observation). Many species have been described on the basis of the extremes of single characters, but no satisfactory taxonomy has emerged that has correlated all morphological and cytological data. There is a certain amount of correlation between major soil types and morphologies. In very general terms densely pubescent forms occur predominantly on very deep, sandy soils, tall, broad-leaved, less-pubescent to glabrous forms on loam and clay soils, and very narrow-leaved, small plants occur on dry, rocky and gravelly soils. There seems to be a continuum of variation and there are exception to the generalizations. Much the same kind of morphologies and patterns of variation occur in the African populations of the perennial species, where there is also a very distinct annual species. In contemporary accounts, the perennial African populations are generally included in one species, the earliest name applying to any species in the genus. Since the African perennials cannot be reliably separated from the American perennials, it seems best to interpret the American populations in the same broad way, although I have tried at times to recognize more narrowly defined species. For those who prefer to recognize the American perennials as distinct from the African ones, the earliest name that applies is Trachypogon plumosus. Trachypogon spicatus (L. f.) Kuntze, Revis. Gen. Pl. 2: 794. 1891. —Stipa spicata L. f., Suppl. Pl. 111. 1781. —Andropogon spicatus Steud., Nomencl. Bot. 1: 93. 1840. Andropogon plumosus Humb. & Bonpl. ex Willd., Sp. Pl. 4: 918. 1806. —Trachypogon plumosus (Humb. & Bonpl. ex Willd.) Nees in Mart. et al., Fl. Bras. Enum. Pl. 2: 344. 1829. —Trachypogon polymorphus var. plumosus (Humb. & Bonpl. ex Willd.) Hack. in Mart., Fl. Bras. 2(4): 265. 1883. Andropogon angustifolius Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 184. 1815 [1816], non Sibth. & Sm. 1806. —Trachypogon angustifolius E. Fourn. ex Hemsl., Biol. Cent.-Amer., Bot. 3(19): 522. 1885. —Trachypogon angustifolius Nees ex Hack. in A. DC. & C. DC., Monogr. Phan. 6: 326. 1889.

Andropogon montufarii Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 184. 1815 [1816]. —Trachypogon montufarii (Kunth) Nees in Mart. et al., Fl. Bras. Enum. Pl. 2: 342. 1829. —Trachypogon polymorphus var. montufarii (Kunth) Hack. in Mart., Fl. Bras. 2(4): 264. 1883. —Trachypogon plumosus var. montufarii (Kunth) Hack., Meded. Rijks-Herb. 40: 40. 1921. Trachypogon canescens Nees in Mart. et al., Fl. Bras. Enum. Pl. 2: 343. 1829. —Andropogon canescens (Nees) Kunth, Enum. Pl. 1: 487. 1833. —Trachypogon polymorphus var. canescens (Nees) Hack. in Mart., Fl. Bras. 2(4): 263. 1883. Trachypogon mollis Nees in Mart. et al., Fl. Bras. Enum. Pl. 2: 343. 1829. —Andropogon mollis (Nees) Kunth, Révis. Gramin. 2: 561, t. 195. 1832. —Trachypogon montufarii var. mollis (Nees) Andersson, Öfvers. Förh. Kongl. Svenska

Trachypogon 289

Vetensk.-Akad. 14: 49. 1857. —Trachypogon polymorphus subvar. mollis (Nees) Hack. in Mart., Fl. Bras. 2(4): 265. 1883. —Trachypogon capensis subvar. mollis (Nees) Roberty, Boissiera 9: 150. 1960. Trachypogon ligularis Nees in Mart. et al., Fl. Bras. Enum. Pl. 2: 345. 1829. —Andropogon ligularis (Nees) Kunth, Enum. Pl. 1: 497. 1833 —Trachypogon polymorphus var. ligularis (Nees) Hack. in Mart., Fl. Bras. 2(4): 265. 1883. Heteropogon stipoides J. Presl, Reliq. Haenk. 1: 335. 1830. —Andropogon stipoides (J. Presl) Kunth, Enum. Pl. 1: 487. 1833, non Kunth 1816. —Trachypogon preslii Andersson, Öfvers. Förh. Kongl. Svenska Vetensk.-Akad. 14: 50. 1857. Heteropogon secundus J. Presl, Reliq. Haenk. 1: 335. 1830. —Andropogon secundus (J. Presl) Kunth, Enum. Pl. 1: 487. 1833, non Elliott 1821. —Trachypogon preslii f. secundus (J. Presl) Andersson, Öfvers. Förh. Kongl. Svenska Vetensk.-Akad. 14: 50. 1857. —Trachypogon polymorphus subvar. secundus (J. Presl) Hack. in A. DC. & C. DC., Monogr. Phan. 6: 326. 1889. —Trachypogon plumosus subvar. secundus (J. Presl) Hack. ex Henrard, Meded. Rijks-Herb. 40: 40. 1921. —Trachypogon plumosus var. secundus (J. Presl) Beetle, Phytologia 54: 5. 1983. Trachypogon capensis Trin., Mém. Acad. Imp. Sci. St. Pétersbourg, Sér. 6, Sci. Math. 2(4): 257. 1832. —Trachypogon polymorphus subvar. capensis (Trin.) Hack. in A. DC. & C. DC., Monogr. Phan. 6: 326. 1889. Trachypogon macroglossus Trin., Mém. Acad. Imp. Sci. St. Pétersbourg, Sér. 6, Sci. Math. 2(3): 257. 1832. —Andropogon macroglossus (Trin.) Steud., Syn. Pl. Glumac. 1: 368. 1855 [1854]. —Trachypogon polymorphus var. macroglossus (Trin.) Hack. in Mart., Fl. Bras. 2(4): 264. 1883. Heteropogon truncatus Nees, Fl. Afr. Austral. Ill. 102. 1841. —Andropogon truncatus (Nees) Steud., Syn. Pl. Glumac. 1: 368. 1855 [1854]. —Trachypogon truncatus (Nees) Andersson, Öfvers. Förh. Kongl. Svenska Vetensk.-Akad. 14: 49.

1857. —Trachypogon polymorphus var. truncatus (Nees) Hack. in A. DC. & C. DC., Monogr. Phan. 6: 327. 1889. Trachypogon dissoluta Nees, Linnaea 19(6): 695. 1847. —Andropogon dissolutus (Nees) Steud., Syn. Pl. Glumac. 1: 381. 1855 [1854]. —Trachypogon polymorphus var. dissolutus (Nees) Hack. in A. DC. & C. DC., Monogr. Phan. 6: 328. 1889. Andropogon dactyloides Steud., Syn. Pl. Glumac. 1: 381. 1855 [1854]. —Trachypogon polymorphus subvar. dactyloides (Steud.) Hack. in Mart., Fl. Bras. 2(4): 265. 1883. Andropogon vestitus Steud., Syn. Pl. Glumac. 1: 378. 1855 [1854]. Trachypogon gracilis Andersson, Öfvers. Förh. Kongl. Svenska Vetensk.-Akad. 14: 50. 1857. —Trachypogon polymorphus subvar. gracilis (Andersson) Hack. in Mart., Fl. Bras. 2(4): 265. 1883. Trachypogon gracilis var. ciliatus Andersson, Öfvers. Förh. Kongl. Svenska Vetensk.-Akad. 14: 50. 1857. Trachypogon gracilis var. hirtus Andersson, Öfvers. Förh. Kongl. Svenska Vetensk.-Akad. 14: 50. 1857. Trachypogon micans Andersson, Öfvers. Förh. Kongl. Svenska Vetensk.-Akad. 14: 47. 1857. Trachypogon montufarii var. grandiflorus Andersson, Öfvers. Förh. Kongl. Svenska Vetensk.-Akad. 14: 49. 1857. Trachypogon montufarii var. pauciflorus Andersson, Öfvers. Förh. Kongl. Svenska Vetensk.-Akad. 14: 49. 1857 Trachypogon vestitus Andersson, Öfvers. Förh. Kongl. Svenska Vetensk.-Akad. 14: 52. 1857. —Trachypogon polymorphus var. vestitus (Andersson) Hack. in Mart., Fl. Bras. 2(4): 266. 1883. Trachypogon polymorphus var. filifolius Hack. in Mart., Fl. Bras. 2(4): 264, pl. 62, fig. 1. 1883. —Trachypogon filifolius (Hack.) Hitchc., Contr. U.S. Natl. Herb. 12: 191. 1909. Trachypogon muelleri E. Fourn., Biol. Cent.-Amer., Bot. 3: 523. 1885. Trachypogon montufarii var. pilosus E. Fourn., Mexic. Pl. 2: 66. 1886. Trachypogon gouinii E. Fourn., Mexic. Pl. 2: 66. 1886. —Trachypogon polymorphus

290

P OACEAE

Fig. 234. Trachypogon spicatus

Tridens 291

var. gouinii (E. Fourn.) Hack. in A. DC. & C. DC., Monogr. Phan. 6: 327. 1889. Andropogon trichospirus Hack. in A. DC. & C. DC., Monogr. Phan. 6: 536. 1889. Trachypogon polymorphus Hack. in Mart., Fl. Bras. 2(4): 263. 1883. Trachypogon polymorphus var. karwinskyi Hack. in A. DC. & C. DC., Monogr. Phan. 6: 329. 1889. —Trachypogon karwinskyi (Hack.) Nash, N. Amer. Fl. 17(1): 97. 1909. Trachypogon polymorphus var. thollonii Franch., Bull. Soc. Hist. Nat. Autun 8: 322. 1893. —Trachypogon thollonii (Franch.) Stapf, Fl. Trop. Afr. 9(3): 402. 1919. Trachypogon polymorphus var. truncatus Hack. in A. DC. & C. DC., Monogr. Phan. 6: 327. 1889. Trachypogon polymorphus var. bolivianus Pilg., Bot. Jahrb. Syst. 27: 22. 1899. —Trachypogon montufarii var. bolivianus (Pilg.) Pilg., Notizbl. Bot. Gart. Berlin-Dahlem 11: 777. 1933. Trachypogon palmeri Nash, N. Amer. Fl. 17(2): 96. 1909. Trachypogon involutus Pilg., Wiss. Erg. Schwed. Rhod.-Kongo-Exped. 1: 196. 1915. Trachypogon durus Stapf, Fl. Trop. Afr. 9: 405. 1919.

Trachypogon glaucescens Pilg., Notizbl. Bot. Gart. Berlin-Dahlem 11: 803. 1933. Trachypogon ramosus Swallen, Mem. New York Bot. Gard. 9(3): 277. 1957. Trachypogon densus Swallen, Phytologia 14: 94. 1966. Trachypogon parviflorus Swallen, Phytologia 14: 94. 1966. Trachypogon rigidifolius Swallen, Phytologia 14: 93. 1966. Trachypogon renvoizei Cat. Guerra, Fontqueria 44: 144. 1996. Trachypogon mayaensis Wipff & S.D. Jones, Sida 18: 242, fig. 1. 1998. Perennial 50–250 cm tall. Savannas, open treeless grasslands, near sea level to 1500 m; widespread in Delta Amacuro, Bolívar, and Amazonas. Anzoátegui, Apure, Aragua, Barinas, Carabobo, Cojedes, Distrito Federal, Falcón, Guárico, Lara, Mérida, Miranda, Monagas, Portuguesa, Sucre, Táchira, Trujillo, Yaracuy, Zulia; Canada, U.S.A., Mexico, Central America, Caribbean, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Chile, Paraguay, Argentina, Uruguay, Africa. ◆Fig. 234. In the flora area, Trachypogon spicatus is dominant in most savannas, especially open treeless grasslands that are frequently burned. However, it is never dominant in the wet parts.

93. TRIDENS Roem. & Schult., Syst. Veg. 2: 34, 599. 1817. [Subfamily Chloridoideae, Tribe Cynodonteae] by Emmet J. Judziewicz Cespitose perennials. Leaves with ligules usually ciliate; blades linear. Inflorescence terminal, an open to spike-like panicle. Spikelets moderately laterally compressed, several-flowered, disarticulating above the glumes and between the florets; glumes unequal to subequal, hyaline, typically much shorter than the spikelet, typically 1-veined or the upper 3–5-veined; florets with lemmas rounded on the back, 3-veined, the veins densely hairy below, often prolonged above into short mucros or awns; paleas shorter than the lemmas, 2-keeled, the keels submarginal; stamens 3; stigmas 2. Eastern U.S.A., Mexico, Central America, West Indies, tropical and warmtemperate South America, Angola; 18 species, 2 in Venezuela, 1 of these in the flora area. Tridens flaccidus (Döll) Parodi, Revista Argent. Agron. 4: 249. 1937. —Uralepis flaccida Döll in Mart., Fl. Bras. 2(3): 95. 1878. —Triodia flaccida (Döll) Hitchc.,

Contr. U.S. Natl. Herb. 22: 457. 1922. Cespitose perennial 60–100 cm tall; leaf blades 18–28 cm × 3–9 mm; inflorescence 15– 20 × 3–7 cm, open; spikelets 6–8 mm long,

292

P OACEAE

Fig. 235. Tridens flaccidus

Tripogon 293

ovate, 5–11-flowered; lemmas 3–4 mm long, 3-veined, the veins ciliate below and excurrent as mucros. Savannas, ca. 50 m; Bolívar

(Río Orinoco). Guárico; Guyana, eastern Brazil. ◆Fig. 235.

94. TRIPOGON Roem. & Schult., Syst. Veg. 2: 34, 600. 1817. [Subfamily Chloridoideae, Tribe Cynodonteae] by Emmet J. Judziewicz Small, densely tufted perennials. Leaves basal; ligules membranous, ciliate; blades involute, filiform. Inflorescence long-exserted, a few-flowered spike; spikelets sessile, appressed to ascending, borne in 2 rows, these mostly not overlapping. Spikelets narrow, several-flowered, laterally compressed, disarticulating between the florets; glumes well developed, usually 1-veined, much shorter than the spikelet; florets with lemmas 3-veined, bidentate, mucronate to awned, the lateral teeth sometimes awned as well. Warm-temperate to tropical in both New World and Old World; 20–30 species, 1 in Venezuela. Tripogon spicatus (Nees) Ekman, Ark. Bot. 11(4): 36. 1912. —Bromus spicatus Nees in Mart., Fl. Bras. Enum. Pl. 2: 471. 1829. Tufted perennial 10–40 cm tall; inflorescence 4–7 cm long; spikelets 5–7 mm long, linear to elliptic, 5–7-flowered; lemmas 3–3.5 mm long, 3-veined, the midvein excurrent as a mucro, the lateral veins as small teeth. Savannas (often rocky or gravelly), 100–300 m; Bolívar (Altiplanicie de Nuria, La Vergareña, 46 km north of La Paragua), Amazonas (southeast of Puerto Ayacucho). Southwestern U.S.A., Mexico, Guatemala, Honduras, Nicaragua, Cuba, Haiti, Colombia, Ecuador, Peru, Brazil, Bolivia, Chile, Paraguay, Argentina, Uruguay. ◆Fig. 236.

Fig. 236. Tripogon spicatus

294

P OACEAE

95. TRIPSACUM L., Syst. Nat. ed. 10: 1261. 1759. [Subfamily Panicoideae, Tribe Andropogoneae] by Emmet J. Judziewicz Robust, cespitose perennials from short, stout rhizomes, the culms stout, solid. Leaves with ligules short, membranous; blades usually broad, elongate, with a prominent white, the lower surface with raised midrib. Inflorescences in both terminal and axillary clusters, exserted or not, each of 1–many fascicled, erect to curving spike-like racemes; racemes with basal portion bearing pistillate spikelets and the longer apical portion staminate spikelets. Pistillate (basal) portion of rachis with cylindrical, straw-colored, indurate, corky, greatly thickened internodes, at maturity readily disarticulating, each joint deeply excavated into a cupule the margins of which overlap the hard lower glume of a solitary, permanently enclosed, sessile spikelet (the pedicellate spikelet suppressed); spikelets partially sunken in the rachis internode, the lower glume ovate-triangular, indurate, concealing the membranous to hyaline upper glume and florets, the lower floret sterile, the upper floret pistillate. Staminate (apical) portion of racemes with rachis internodes slender, triquetrous, ± disarticulating, the spikelets paired and similar, one member of the pair sessile or subsessile, the other often pedicellate; lower glume as long as the spikelet or occasionally distinctly shorter, firm, narrowly elliptic to lanceolate, obtuse to asymmetrically acute, flat, and finely many-veined on the back, the marginal veins prominent, scabrous to ciliate-scabrid, the margins inflexed over the margins of the upper glume and enclosed florets; upper glume as long as the spikelet, delicately membranous, many-veined; florets in pairs, similar, both staminate, hyaline, the lemmas several-veined, the paleas well developed. Eastern U.S.A., Mexico, Central America, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay; ca. 13 species, 2 in Venezuela, 1 of these in the flora area. Tripsacum australe Cutler & E.S. Anderson, Ann. Missouri Bot. Gard. 28: 259. 1941. Robust, cespitose perennial, the culms 2– 3.5 m × 1–1.5 cm; leaves with lower sheaths woolly (at least when young), the blades 30– 140 × 2–4 cm, tapering to a pseudopetiolate base; racemes 1–4, each 15–33 cm long, the

basal 1/3 pistillate, the upper 2/3 staminate; pistillate spikelets 5.5–7 mm long; staminate spikelets 6.5–8.5 mm long. Savannas, ca. 100 m; Amazonas (Culebra). Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina. ◆Fig. 237.

96. ZEUGITES P. Browne, Civ. Nat. Hist. Jamaica 341. 1756. [Subfamily Arundinoideae, Tribe Centotheceae] Despretzia Kunth, Révis. Gramin. 2: [485]. 1831. Krombholzia Rupr. ex E. Fourn., Bull. Soc. Roy. Bot. Belgique 15: 464. 1876. Senites Adans., Fam. Pl. 2: 39, 604. 1763. by Gerrit Davidse Perennials, slender and creeping to coarse and reed-like. Leaves cauline; sheaths rounded on the back; ligule membranous; pseudopetioles usually well developed; blades lanceolate to ovate, flat, usually thin, transversely veined. Inflores-

Tripsacum 295

Fig. 237. Tripsacum australe

296

P OACEAE

Fig. 238. Zeugites sp. A

P O D O C A R PA C E A E 297

cence a terminal, open panicle; rachis and main panicle branches often viscid. Spikelets bisexual, pedicellate, 3–15-flowered, solitary, laterally compressed; glumes subequal in length, shorter than the spikelets, broad, obtuse or truncate, often irregularly toothed or lobed, conspicuously cross-veined, the upper glume generally narrower than the lower glume; florets dimorphic; lowest floret pistillate; upper 2–14 florets staminate; rachilla joint between the pistillate and staminate florets usually elongate; disarticulation below the glumes and the base of the staminate florets, the staminate florets falling as 1 unit; lemma of fertile floret broad, usually obtuse; fertile palea a little shorter to a little longer than the fertile lemma; styles 1 or 2; stigmas plumose; lodicules 2, truncate, vasculated; hilum punctate, embryo 1/4–1/3 as long as the caryopsis; staminate florets narrower than the pistillate floret, acute to subobtuse; palea almost as long as the lemma; lodicules 2, truncate, vasculated; stamens 3. Fruit a caryopsis; hilum punctate; embryo 1/4–1/3 as long as the caryopsis. Neotropics; 10 species, 2 in Venezuela, 1 of these in the flora area. Zeugites sp. A Reed-like perennial 1–2 m tall. Evergreen lowland forests, 100–200 m; Bolívar (lower Río Suapure), Amazonas (vicinity of Puerto Ayacucho). Endemic. ◆Fig. 238. Zeugites sp. A resembles Z. panamensis in having well-developed pseudopetioles to 40 mm long and nearly glabrous pulvini in leaves from the middle nodes of the culms. However in Z. sp. A, a pulvinus is developed only at the base of the pseudopetiole, not at the apex and base as in Z. panamensis and the closely related Z. pittieri. The glumes are

approximately 1/2 as long as the pistillate floret in Z. sp. A, but about 3/4 as long in the type of Z. panamensis. It is intriguing that this is apparently the only lowland species in the genus, which otherwise only occurs at middle elevations. The reed-like species were previously only known from Mesoamerica and Mexico. The nearest locality for any of these species is Zeugites panamensis in central Panama. The specimen from Bolívar is sterile, but matches quite well the fertile specimens from Amazonas.

PODOCARPACEAE by David J. de Laubenfels Trees or shrubs, rarely prostrate. Roots with nodules (of unknown function, not nitrogen fixing). Leaves evergreen, spirally placed or opposite and decussate, simple, petiolate or sessile, decurrent; margins entire; a single midvein or many parallel veins branching dichotomously near the base. Foliage buds weakly to strongly developed, consisting of a cluster of scales. Inflorescences terminal and/or axillary, spicate, paniculate, or solitary cones or modified cones, unisexual (plants monoecious or dioecious). Buds for pollen cones similar to foliage buds; pollen cones cylindrical, with numerous microsporophylls, each with 2 inverted pollen sacs below a small erect apex. Seed cones usually on current growth, subtended by a scaly or naked peduncle or nearly sessile, a compound structure with ovule-bearing structures in the axil of a fertile bract, 1–few basal sterile bracts and 1–many fertile bracts; the ovule structure a single ovule ± surrounded by a scale in the form of a filmy

298

P ODOCARPACEAE

epimatium (corresponding to the “seed scale” of Pinaceae, it subtends the ovule and cups it like an aril) or a leathery or membranous covering, or the ovule naked; the ovule inverted or erect, the associated structures (scales or bracts or both) usually becoming enlarged and fleshy at maturity. Seeds with a hard woody outer layer, naked or covered, usually with weak lateral keels and acute at the micropylar end, with abundant endosperm. Cotyledons 4 but fused to form 2 double structures with a double apex. Pantropics and subtropics to Mexico, Cuba, China, and Japan, and throughout the southern hemisphere temperate forests; 13 genera and ca. 175 species, 1 genus and 9 species in the flora area. 1. PODOCARPUS L’Her. ex Pers., Syn. Pl. 2: 580. 1807, nom. cons., non Labill. 1806. Shrubs to large trees or occasionally prostrate. Leaves spirally arranged, evergreen, simple, sessile to petiolate; blades linear to ovate, entire, with a single median vascular bundle. Growth bud well developed, several tough, erect or overlapping scales surrounding several overlapping membranous scales which emerge with growth and spread as the leafy shoot elongates. Buds for pollen cones axillary on previous year’s growth, rarely terminal. Pollen cones solitary or grouped in a cluster, a spike, or occasionally a panicle. Seed-bearing structure a highly modified cone subtended by a naked peduncle and 2–several enlarged bracts, 1–several of which are fertile, the bracts subtended by 2 small lanceolate foliola (subgenus Foliolatus Laubenf.) or without foliola (subgenus Podocarpus). Seeds inverted and completely covered by a leathery scale; bracts becoming greatly enlarged and fleshy when mature or rarely remaining leathery. Distribution the same as the family; ca. 100 species, 13 in Venezuela, 9 of these in the flora area (all in subgenus Podocarpus). Key to the Species of Podocarpus 1. 1. 2(1). 2. 3(2). 3. 4(3). 4. 5(3). 5. 6(5). 6. 7(6). 7. 8(6). 8.

Upper surface of leaves with a ridge over the midvein ............... P. roraimae Upper surface of leaves with a groove over the midvein ......................... 2 Terminal buds globular, with overlapping scales ....................... P. tepuiensis Terminal buds with erect or spreading scales .......................................... 3 Mature leaves ≤ 7 mm wide ....................................................................... 4 Mature leaves ≥ 7 mm wide ....................................................................... 5 Mature leaves ± ovate, > 5 mm wide; bud scales mostly 2–3 mm long, but 1 or 2 sometimes over 6 mm long and foliose ........................ P. buchholzii Mature leaves long and narrow, 4–5 mm wide; bud scales to 9 mm long but not foliose ............................................................................ P. aracensis Bud scales lanceolate, 6–25 mm long ..................................... P. steyermarkii Bud scales triangular or cuspidate, < 6 mm long ..................................... 6 Leaf apices acuminate; fruits with prominent crest ................................ 7 Leaf apices acute or rounded; fruits with weak crest or none ................. 8 Leaves ca. 8 mm wide ............................................................... P. acuminatus Leaves ≥ 18 mm wide ................................................................ P. magnifolius Leaves 8–10 mm wide; juvenile leaves linear; fruits ovoid .... P. brasiliensis Leaves 10–15 mm wide; juvenile leaves lanceolate; fruits globular ....................................................................................................... P. celatus

Podocarpus 299

Podocarpus acuminatus Laubenf., Novon 2: 329. 1992. Small sun-loving tree 4–5 m tall; leaves small, acuminate; bud scales noticeably cuspidate; fruits about 7 × 4 mm with a sharp apical crest; receptacle with extra bracts beside or between the larger 2, the bracts with spreading tips. Upper montane forests on tepuis summits and slopes, often among rocks, 1900–2400 m; Bolívar (Amurí-tepui), Amazonas (Sierra de la Neblina). Brazil (Amazonas: Serra da Neblina). Podocarpus aracensis Laubenf. & Silba, Phytologia 65: 330. 1988. Bushy tree to 6 m tall; leaves narrow, revolute with acute but slightly rounded tips and elongated erect bud scales; pollen cone with elongated scaly peduncle. Tepui scrub forests, in rock crevices, 1200–2500 m; Bolívar (Macizo del Chimantá [Murey-tepui]), Amazonas (Cerro Yaví). Brazil (Amazonas: Serra Aracá). Podocarpus brasiliensis Laubenf. in Luces & Steyerm., Fl. Venez. 11(2): 32, fig. 10. 1982. Small tree to 15 m tall; leaves linear and somewhat rounded at the apex; bud scales short, triangular, erect. Montane forests, often along streams, 800–1700 m; Bolívar (Auyán-tepui), Amazonas (Cerro Yapacana). Widespread on the interior plateaus of Brazil. Podocarpus buchholzii Laubenf. in Luces & Steyerm., Fl. Venez. 11(2): 31, fig. 9. 1982. Shrub 1–2 m or small tree to 10 m tall; leaves small, erect, triangular; bud scales often accompanied by 1 or 2 elongate foliose scales. Tepui scrub, boggy areas in Bonnetia forests, in crevices, among rocks at the base of cliffs, 1900–2500 m; Bolívar (Macizo del Chimantá), Amazonas (Sierra de la Neblina). Anzoátegui (Cerro Peonía); Brazil (Amazonas: Serra da Neblina, Rio Negro). Podocarpus celatus Laubenf. in Luces & Steyerm., Fl. Venez. 11(2): 35 fig. 12. 1982. Tree 6–25 m tall; leaves similar to P. brasiliensis when mature but markedly lanceolate when juvenile; bud scales short and sometimes almost overlapping. Lower montane to montane forests, 400–1400 m;

Bolívar (Cerro Marutaní, near Icabarú), Amazonas (Cerro Marahuaka, Río Coro Coro west of Cerro Yutajé). Venezuelan Andes; Colombia, Peru, Brazil, Bolivia. Podocarpus magnifolius J. Buchholz & N.E. Gray, J. Arnold Arb. 29: 133. 1948. Tree 10–25 m tall; leaves rather broad and acuminate at each end, long and linear when juvenile, usually distinctly revolute. Forests on upper tepui slopes, 700–1900 m; Bolívar (Ptari-tepui), Amazonas (Sierra de la Neblina). Widespread in northern Venezuela; Panama, Colombia, Peru, western Brazil, Bolivia. Podocarpus roraimae Pilger, Notizbl. Königl. Bot. Gart. Berlin 5: 299. 1913. Small tree 1–5(–10) m tall; leaves 2–3 cm × 5–7 mm, glaucous, especially on lower surface, bud scales short and erect, sometimes accompanied by 1 or 2 reduced leaves. Tepui scrub on rock outcrops, forest margins, Bonnetia forests, 1800–2700 m; Bolívar (Macizo del Chimantá, Roraima-tepui), Amazonas (Cerro Marahuaka, Sierra de la Neblina). Endemic. ◆Fig. 240. Podocarpus roraimae grows near the borders of both Guyana and Brazil and may eventually be found in those countries. Podocarpus steyermarkii J. Buchholz & N.E. Gray, J. Arnold Arb. 29: 133. 1948. Small tree 5–12 m tall; leaves linear and narrowly acute to slightly acuminate, in juvenile specimens distinctly lanceolate; foliage bud scales erect, narrowly lanceolate, and slightly broadened at the base, the whole bud longer than wide. Tepui summit and slope forests, 1700–2300 m; Bolívar (Carraotepui near Ptari-tepui, Cerro Jaua, Uaipántepui), Amazonas (Cerro Yaví, Sierra de la Neblina). Anzoátegui (Cerro Peonía). ◆Fig. 239. Podocarpus steyermarkii grows along the Brazilian border and most likely occurs in Brazil in Amazonas state. Podocarpus tepuiensis J. Buchholz & N.E. Gray, J. Arnold Arb. 29: 134. 1948. Shrubby tree 1–6(–15) m tall; bud scales overlapping, forming a small round ball, leaves 15–35(–60) × 3–5(–8) mm, juvenile form long and narrow. Tepui slope and summit forests, especially along streambanks,

300

P ODOCARPACEAE

also known from one lowland site on white sand in a seasonally flooded (black-water) forest close to a savanna, (100–)700–2000 m; widespread in mountains of Bolívar and Amazonas (the lowland site in Amazonas is from Budare in the upper Río Temi basin upstream from Yavita). Ecuador, Peru. ◆Fig. 241.

Fig. 239. Podocarpus steyermarkii

Fig. 240. Podocarpus roraimae

Fig. 241. Podocarpus tepuiensis

P O D O S T E M A C E A E 301

PODOSTEMACEAE by Paul E. Berry Aquatic herbs attached to rocky or other substrates (haptophytes or rheophytes) in rapidly flowing streams and rivers, with special root hairs or rhizoids that adhere strongly to the substrate, flowering and fruiting when emergent above receding waters. Stems present, sometimes obscure, branched or unbranched, usually in pairs along dorsiventrally flattened roots. Leaves di-, tri-, or pleiostichous, in the apparently stemless species often united at the base, entire or much-divided, sometimes with tufts of filaments on the upper surface or covered with stiff emergences; sheaths 0, 1, or 2 on either side of the base, sometimes connate into a 1- or 2-lobed, intrapetiolar ligule. Inflorescences of axillary and terminal, solitary or fascicled flowers inserted between the leaf bases or at the end of short shoots, or extra-axillary, or on 2-sided spike-like monochasia, the flowers alternating with leaf-like bracts. Flowers bisexual, actinomorphic to zygomorphic, enveloped by 2–5 thin basal leaves, or enveloped by a thin sac-like spathella, rarely many flowers in a single spathella. Tepals 2–many, petaloid or reduced to small or narrow scales, free or somewhat connate. Stamens 1–many, sometimes 2 or 3 on a common stalk (andropodium) with 1 tepal in or just below the fork, with or without 1 tepal on one or either side of the base of the andropodium; anthers sagittate, longitudinally dehiscent by 2 slits. Ovary superior, 2- or 3-locular (1-locular in some Asian species), sometimes on a short gynophore; carpels equal to very unequal; placentas axillary, with 2–many anatropous ovules; style present or absent, stigmas 1–3, variable in shape and size, free or basally connate. Fruit a 2- or 3-locular, generally ribbed, septifragal capsule, the smallest valve usually caducous in fruits with unequal valves. Seeds 2–many, the exotesta thick-walled and mucilaginous upon imbibition; embryo straight, endosperm lacking. Eastern Canada and U.S.A., Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia, Paraguay, Argentina, Uruguay, tropical Africa, Madagascar, Mascarene Islands, Asia, Australia; 48 genera and ca. 270 species (R. Rutishauser, Aquat. Bot. 57: 30. 1997), 8 genera and 26 species in the flora area. Vegetatively, the family resembles algae, and its species are a characteristic element of the fast-flowing streams and rivers of the region. Plants are vegetative during the rainy season and notoriously polymorphic in leaf form, but they develop flowers and fruits when the water level drops in the dry season. The family is embryologically unusual in lacking double fertilization and endosperm. The phylogenetic position of the family has long been an enigma, but molecular studies now place Podostemaceae consistently sister to Clusiaceae or Hypericaceae in the Malpighiales. The treatment below is based on the three-part family monograph by P. van Royen (The Podostemaceae of the New World. Part I. Meded. Bot. Mus. Herb. Rijks Univ. Utrecht 107: 1–151. 1951; The Podostemaceae of the New World. Part II. Acta Bot. Neerl. 2: 1–20. 1953; The Podostemaceae of the New World. Part III. Acta Bot. Neerl. 3: 215–263. 1954). It must be emphasized, however, that the family in the Neotropics is desperately in need of a modern systematic revision (see C. T. Philbrick & A. Novelo R. New World Podostemaceae: ecological and evolutionary enigmas. Brittonia 47: 210–222. 1995). Van Royen apparently lacked the necessary

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field experience to understand the broad variation in vegetative and reproductive characters that occurs in many taxa. For the flora area, this problem is particularly acute with many of the species described in Apinagia, Marathrum, and Rhyncholacis. Furthermore, these genera can be very difficult to distinguish morphologically (especially Apinagia and Marathrum), and they will probably prove to be polyphyletic or paraphyletic with further study (T. Philbrick, pers. comm.). The family presents excellent opportunities for researchers who will be able to combine field observations with detailed molecular and morphological studies. Future studies will likely result in significant changes in species and genus concepts in the family. Key to the Genera of Podostemaceae 1.

1.

2(1).

2.

3(2).

3.

4(3). 4.

5(4). 5. 6(4). 6. 7(6).

Leaves tristichous, 1–3 mm long, oval, sessile, and glabrous; plants mosslike, 3–10 cm long; perianth calyx-like, 3-lobed; anther 1; ovary 3-locular, with 3 divergent stigmas; capsule with 3 valves (subfamily Tristichoideae) ........................................................................... 7. Tristicha Leaves distichous or different from above; plants various in habit; perianth scale-like or with 5 segments; anthers 1–many; ovary 2-locular, with either 2 stigmas or a single style; capsule with 2 valves ............ 2 Flowers solitary, not enclosed by a spathella, the perianth with 5 lanceoblong tepals; stamens 5–25; ovary with a single style topped by a globular-papillose stigma (subfamily Weddellinoideae) ...... 8. Weddellina Flowers solitary, fasciculate, or on a spike-like inflorescence, enclosed by a membranous spathella, the perianth filiform or scale-like; stamens 1–many; ovary with 2 separate, nonpapillose stigmas (subfamily Podostemoideae) .................................................................................... 3 Inflorescences large, erect, sword-like, flattened spikes to 10–65 cm tall, with 40–90 pinkish flowers; leaves large, with lobed margins, 8–200 × 2–30 cm, with coarsely rigid outgrowths on the upper surface .................................................................................................... 4. Mourera Flowers solitary or fasciculate, axillary or terminal, never in a flattened spike-like inflorescence; leaves smaller or more divided than above, without coarse outgrowths on the upper surface ................................. 4 Plants very small, the vegetative leaves 0.5–2 cm long; flowers solitary, pedicels 1–14 mm long; stamens 1 or 2 (rarely 3) per flower .............. 5 Plants generally (much) larger than above, with longer vegetative leaves; flowers solitary or fasciculate, pedicels 5–150 mm tall; stamens generally > 3 per flower (3 or fewer in a few Apinagia species) ................... 6 Stamens 1 per flower, anther basifixed; pedicels 4–14 mm long .... 5. Oserya Stamens 2 or less often 3 per flower, anthers dorsifixed; pedicels 1–4 mm long ....................................................................................... 2. Jenmaniella Stigmas rigid and beaked, marcescent; midrib of carpels and valves winged ................................................................................ 6. Rhyncholacis Stigmas cylindric to linear, neither stiff or beaked, falling off when the fruit ripens; midrib of carpels and valves ribbed but not winged ....... 7 Plants stemless (without internodes); stigmas subulate with widened and emarginate top; each valve of capsules 3- or 4-ribbed ........ 3. Marathrum

Apinagia 303

7.

Plants stemless or stemmed (with internodes); stigmas not as above; each valve of capsule with (1)3–7 ribs .............................................. 1. Apinagia

1. APINAGIA Tul., Ann. Sci. Nat. Bot. sér. 3, 11: 90. 1849, emend. P. Royen, Meded. Bot. Mus. Herb. Rijks Univ. Utrecht 107: 25. 1951. Small to large herbs, either thalloid or with a single unbranched stem. Leaves distichous, connate at base in thalloid species, pinnately or palmately veined, or veinless, often with tufts of hairs on the upper surface. Flowers solitary or fasciculate, the inflorescences extra-axillary, sometimes much-branched with terminal and axillary flowers, or the flowers in compound but strongly contracted inflorescences arising from the base of an unbranched stem or a leaf. Tepals 2–many; if stamens 2 or 3 then with 1–3 tepals alternate with the stamens at the base of the ovary. Stamens 1–many, in 1 or 2 complete whorls, or in an incomplete whorl; anthers dehiscing introrsely or extrorsely. Ovary with 2 usually equal (rarely unequal) carpels, with 2–14 dark lines; stigmas cylindric to linear, free or basally connate. Capsule variously ribbed. Colombia, Venezuela, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia, Paraguay, Argentina, Uruguay; ca. 50 species, 10 in Venezuela, all in the flora area. Apinagia can be very difficult to distinguish from taxa described under Jenmaniella, Marathrum, or Rhyncholacis. As discussed above, these genera are likely polyphyletic, and there has probably been a severe oversplitting of species. Future field studies should carefully examine the variability of vegetative characters and stamen number, as well as other traits used below in the key. Key to the Species of Apinagia 1. 1. 2(1).

2.

3(2). 3. 4(3). 4. 5(4). 5. 6(5). 6. 7(4).

Leaves pinnate, the pinnae formed by bundles of linear segments to 15 cm long .............................................................................. A. guyanensis Leaves not pinnate, or if pinnate then the rachis always < 3 mm wide and pinnae different from above .................................................................. 2 Upper surface of leaves with tufts of filaments in 2 lines parallel to midrib; leaves lanceolate, usually with only apical part strongly dissected ................................................................................................ A. staheliana Upper surface of leaves with tufts of filaments scattered; leaves lanceolate or not, usually dissected along entire length, rarely at tip only ................................................................................................................ 3 Herbs with main vegetative stem to 100 cm long and a few short side branches ........................................................................ A. multibranchiata Smaller, much-branched herbs to 30 cm long, sometimes thalloid ......... 4 Stamens 10 or more, in a complete whorl ................................................. 5 Stamens 1–9, in an incomplete whorl ....................................................... 7 Stamens < 12; tepals 4–10; leaves elliptic or obliquely rectangular to rhombiform, sometimes cuneiform, to 10 × 4 cm ............... A. richardiana Stamens and tepals 10–30; leaves lanceolate, 3–26 × 0.5–5 cm .............. 6 Capsule with 2 long ribs and a shorter one on each side ........... A. longifolia Capsule with 6 or 8 long ribs ............................................................. A. kochii Stamens 2, on an andropodium; leaves repeatedly forked, ca. 10 cm long,

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the tips many-laciniate .......................................................... A. ruppioides Stamens 1–9, not on an andropodium; leaves not as above ..................... 8 Leaves elliptic or obliquely rectangular to rhombiform, tips dissected, upper surface with bundles of filaments; capsule with 2 long and 4 short ribs ........................................................................... A. richardiana 8. Leaves pinnatisect to bipinnatisect, all tips either entire or dissected, upper surface with or without bundles of filaments; capsule with similarly long ribs ......................................................................................... 9 9(8). Tips of leaves and lobes entire; upper surface of leaves with bundles of filaments ................................................................................ A. corymbosa 9. Tips of leaves and lobes finely dissected; upper surface of leaves without bundles of filaments ............................................................................ 10 10(9). Leaves pinnatisect to bipinnatisect ................................................... A. exilis 10. Leaves pinnate, secondary pinnae composed of bundles of subfiliform segments ................................................................................ A. brevicaulis 7. 8(7).

Apinagia brevicaulis Mildbr., Beitr. Kenntn. Podost. 42. 1904. Small, branched herb 1–2 cm long; leaves pinnate, 1–2 cm long; flowers lilac, tepals 4– 6, stamens 3–5 on one side of the flower. Waterfalls, 50–200 m; Bolívar (falls at Isla Cuchivero in Río Cuchivero). Endemic. ◆Fig. 246.

Similar to Apinagia corymbosa except the tip of the leaf and lobes dissected into many filiform threads and lacking the tufts of filaments on the upper surface. Rapids and waterfalls, 50–400 m; Bolívar (Río Caroní above San Félix and falls farther upstream). Guyana, Suriname. ◆Fig. 242.

Apinagia corymbosa (Tul.) Engl. in Engl. & Prantl, Nat. Pflanzenfam. ed. 2, 18a: 38. 1930 [1928]. —Neolacis corymbosa (Tul.) Wedd. in A. DC., Prodr. 17: 60. 1873. —Ligea richardiana var. corymbosa Tul., Ann. Sci. Nat. Bot. sér. 3, 11: 96. 1849. Colombia, Venezuela, Guyana, Suriname, French Guiana, northern Brazil; 2 varieties, 1 in Venezuela. The second variety, var. capillarifolia (Engl.) P. Royen, is found in northern Brazil.

Apinagia guyanensis (Pulle) P. Royen, Meded. Bot. Mus. Herb. Rijks Univ. Utrecht 107: 37. 1951. —Oenone guyanensis Pulle, Enum. Vasc. Pl. Surinam 193, t. 7. 1906. Medium to large stemless herb; base branched, fleshy, 5–10 cm high, 2–4 cm wide; leaves pinnate, to 70 cm long or longer, pinnae repeatedly forked, 3–16 cm long, ultimate divisions lance-linear; flowers white to pink, with 8–15 tepals, 8–23 stamens, pedicels 2–12 cm long; each valve of capsule with 3 ribs. Rapids and waterfalls, 100–300 m; Bolívar (Río Toro, Serranía de Imataca). Suriname, French Guiana, Brazil.

A. corymbosa var. corymbosa Small, strongly branched herb 4–18 cm tall; leaves 1–4 cm long, pinnatisect, basal ones sometimes pinnatilobed with a cuneate base, palmativeined; tepals 3–6, stamens 2–5 on one side of the flower. Waterfalls, 50–300 m; Bolívar (Río Caroní at Salto Pica-Pica near Boquita). Distribution as in species. Apinagia exilis (Tul.) P. Royen, Meded. Bot. Mus. Herb. Rijks Univ. Utrecht 107: 53, pl. 3, figs. 8–13. 1951. —Ligea richardiana var. exilis Tul., Ann. Sci. Nat. Bot. sér. 3, 11: 96. 1849. —Neolacis corymbosa var. exilis Tul. in A. DC., Prodr. 17: 60. 1873.

Apinagia kochii (Engl.) P. Royen, Meded. Bot. Mus. Herb. Rijks Univ. Utrecht 107: 38, pl. 3, figs. 1–4. 1951. —Oenone kochii Engl., Bot. Jahrb. Syst. 61(Beibl. 138): 1. 1927. —Carurú. Small to medium-sized herb; stem branched, internodes winged; leaves entire to pinnatilobed, 10–25 × 2–2.5 cm; flowers white, tepals 15–20, stamens 25–30. Rapids and waterfalls, 700–1000 m; Bolívar (Río Uairén in Río Kukenán basin). French Guiana, Brazil (Roraima).

Apinagia 305

Apinagia longifolia (Tul.) P. Royen, Meded. Bot. Mus. Herb. Rijks Univ. Utrecht 107: 36. 1951. —Oenone longifolia Tul., Ann. Sci. Nat. Bot. sér. 3, 11: 96. 1849. Medium-sized herb with distinct stem 5– 30 cm long, unbranched or slightly branched; leaves pinnatilobed to pinnatisect, 2–20 × 0.5–5 cm, with many tufts of 3–6 mm long filaments on upper surface; tepals 10–17, stamens 10–30; pedicel 4–9 cm long in fruit. Rapids and waterfalls, 30–600 m; Delta Amacuro (upper Río Amacuro, Serranía de Imataca), Bolívar (between Caicara and Puerto Ayacucho, Parque la Llovizna on Río Caroní, Río Paragua at Raudales de Maihia, Río Caura at Raudal la Mura and Salto Pará, Río Icabarú at Raudales de Yumariba), Amazonas (Río Orinoco at El Motín, mouth of Río Orinoquillo). Guyana, Suriname, French Guiana, northern Brazil. ◆Fig. 245. Apinagia longifolia is allied to A. multibranchiata and A. staheliana. Apinagia multibranchiata (Matthiesen) P. Royen, Meded. Bot. Mus. Herb. Rijks Univ. Utrecht 107: 39. 1951. —Oenone multibranchiata Matthiesen, Biblioth. Bot. 15(68): 48, t. 1, 5, 6, figs. 81–90C. 1908. Medium-sized annual herb; shoots strongly dimorphic, the submerged vegetative shoots usually unbranched and to > 1 m long, the emergent fertile shoots branched with short flowering shoots; vegetative leaves distichous, lanceolate to ovate, 5–15 (–24) × 0.5–3 cm, the tips rounded to toothed or sometimes fimbriate, with many tufts of filaments on the margins or upper surface; flowers solitary in the axil of one of a pair of leaves, covered in bud by a tubular sheath to 1 cm long; tepals ligulate, 6–16, ca. 0.5 mm long; stamens 6–23 in 1 or 2 whorls; stigmas subulate, filiform, 1–1.5 mm long; pedicels and filaments white to pink, pedicels 0.5–2 cm long at anthesis; flowers bee-pollinated; capsules with 3 short ribs on each valve, the pedicels elongating to 5–8 cm long; seeds ca. 600 per fruit [information from R. Rutihauser and M. Grubert. Bot. J. Linn. Soc. 132: 299–323. 2000]. Rapids and waterfalls at 30–900 m; Bolívar (Río Caroní at Macagua and at Salto Revaloso, Río Caura, Río Icabarú near Uaiparú, Salto Hacha at Canaima), Amazonas (Río Yawarui tributary of upper Río Matacuni). Carabobo.

Apinagia multibranchiata is similar to A. staheliana, and the two may prove to be conspecific. Apinagia richardiana (Tul.) P. Royen, Meded. Bot. Mus. Herb. Rijks Univ. Utrecht 107: 44. 1951. —Ligea richardiana Tul., Ann. Sci. Nat. Bot. sér. 3, 11: 96. 1849. —Uirín (Arekuna). Small to medium-sized herb; stem strongly branched, 1–30 cm long; leaves to 10 × 4 cm, elliptical to asymmetrically rectangular or rhombiform; flowers white to pink, tepals 4–10, stamens 3–10; each valve of capsule with 3 ribs (the central one more prominent). Rapids, waterfalls, 50–800 m; Delta Amacuro (Río Amacuro upstream of San Victor), Bolívar (Río Caura at Salto Mura, Río Icabarú at Raudales de Yumariba, below mouth of Río Nichare, Río Paragua at Raudales de Maihia, Salto Pará), Amazonas (Salto Yureba on Río Yureba). Widely scattered in northern Venezuela; Guyana, Suriname, French Guiana, Brazil. ◆Fig. 244. Apinagia richardiana is a common and variable species. The label of Bernardi 6678 (NY: Río Icabarú) states that the exposed leaves of this species were a favorite food of leaf-cutter ants (Atta sexdens). Apinagia ruppioides (Kunth) Tul., Ann. Sci. Nat. Bot. sér. 3, 11: 98. 1849. —Podostemum ruppioides Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 1: 246. 1815 [1816]. Medium-sized branched herb to ca. 15 cm tall; leaves repeatedly forked, ca. 10 cm long, ultimate divisions filiform; flowers poorly known, apparently with one forked filament and 2 stamens. Rapids at 30–300 m; Bolívar (Río Caura), Amazonas (Raudales de Atures, Raudales de Santa Bárbara del Orinoco). Colombia. ◆Fig. 243. Apinagia ruppioides has been collected growing on dead branches stuck in rapids. Apinagia staheliana (Went) P. Royen, Meded. Bot. Mus. Herb. Rijks Univ. Utrecht 107: 40. 1951. —Oenone staheliana Went, Verh. Kon. Ned. Akad. Wetensch., Afd. Natuurk., Tweede Sect. 2, 25: 19, t. 3, 4, figs. 20–25. 1926. Medium-sized to large herb; stem branched or not, to 2 m long; leaves lanceolate, 10–30 cm long, blade 1–15 cm long,

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Fig. 242. Apinagia exilis

Fig. 243. Apinagia ruppioides

Fig. 244. Apinagia richardiana

Fig. 245. Apinagia longifolia

Fig. 246. Apinagia brevicaulis

Marathrum 307

the dissected apical part 5–15 cm long; flowers red to white, in extra-axillary inflorescences to 25 cm long, or along a branched stem, united only at base with the foliar part, rarely these inflorescences strongly

branched; tepals 10–15, stamens 8–27; each valve of capsule with 3 ribs. Rapids, 30–400 m; Bolívar (Río Caroní, Río Caura), Amazonas (upper Río Orinoco at Raudal de los Guaharibos). Guyana, Suriname.

2. JENMANIELLA Engl., Bot. Jahrb. Syst. 61(Beibl. 138): 7. 1927. Tiny stemless herbs, rarely with a short stem. Leaves distichous, mostly a few times forked or pinnate with forked pinnae. Flowers few, solitary. Tepals 2–7, in a complete or incomplete whorl. Stamens 1–7, in a complete or incomplete whorl, free or sometimes on an andropodium with 2 stamens, sometimes all differing in size and in one flower. Ovary ellipsoid, terete, or subterete, with 2 equal carpels, 6ribbed, but the ribs on the sutures sometimes marked as twin ribs, on a short gynophore; styles mostly subulate, free. Capsules with equal or unequal valves, each valve 3- or 5-ribbed. Venezuela, Guyana, northern Brazil; 7 species, 1 in Venezuela. Jenmaniella ceratophylla Engl., Bot. Jahrb. Syst. 61 (Beibl. 138): 7, t. 8, figs. A–F. 1927. Venezuela, Guyana; 3 varieties, 1 in Venezuela. The other two varieties are known only from Guyana. J. ceratophylla var. parva P. Royen, Meded. Bot. Mus. Herb. Rijks Univ.

Utrecht 107: 125, 137, pl. 16, figs. 14, 15. 1951. Tiny herb with thalloid shoots; leaves 0.5– 1.5 cm long; tepals 2 or 3; stamens 2(3); pedicels 1–3 mm long. Rapids and waterfalls at 50–2000 m; Bolívar (Río Caroní above San Félix and at mouth of Río Curapacay, Río Erebato, Río Marajano on summit of Cerro Jaua). Endemic. ◆Fig. 247.

Fig. 247. Jenmaniella ceratophylla var. parva

3. MARATHRUM Bonpl. in Humb. & Bonpl., Pl. Aequinoct. 1: 39, t. 11. 1808 [1806]. Annual or perennial, small to medium-sized herbs, base often thalloid, usually branched. Roots elongate, prostrate and flattened or short and fan-shaped. Leaves mostly distichous, either repeatedly forked or pinnate, sometimes subentire with a few lobes along the margin. Flowers 1–many, solitary or in fascicles. Tepals 3–25, in a complete or incomplete whorl. Stamens 2–25, in a complete or incomplete whorl. Ovary 2-locular, carpels equal or unequal, 6–8-ribbed; stigmas 2, basally connate, often emarginate at tip. Capsule similar to ovary, with 2 equal valves, persistent, each valve 3- or 4-ribbed, suture margins thickened. Seeds numerous.

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Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Brazil; ca. 25 species, 4 in Venezuela, all in the flora area. Marathrum is part of a taxonomically difficult complex with Apinagia and Rhyncholacis. Species delimitations are very problematical based on current information and the lack of a better understanding of how variable species may be vegetatively and reproductively. This is shown by the overlap in vegetative and reproductive characters in the key below. Van Royen (P. Royen, Meded. Bot. Mus. Herb. Rijks Univ. Utrecht 107: 90–91. 1951) cited Tate 1243 (NY), from the lower slopes of Auyán-tepui at 500 m, as a possible new species of Marathrum. It has numerous, fascicled leaves 1.5–20 cm long that are repeatedly pinnate and has distinctly pointed stipules. Key to the Species of Marathrum 1.

1.

2(1). 2. 3(2). 3.

Leaves either entire or pinnately lobed to parted, cuneate to spathulate in outline, rhombiform in younger leaves, lobes few and entire, or each lobe with 2–5 narrow lobes, the apex with many lanceolate segments to 5 mm long; tepals and stamens 5–8 each ...................................M. utile Leaves repeatedly pinnate or forked, sometimes pinnate with repeatedly forked segments, rarely cuneate with forked lobes at tip; tepals 3– 12; stamens 2–40 ................................................................................... 2 Tepals 8–12; stamens 4–40, arranged around the flower ..... M. squamosum Tepals 3–5; stamens 2–4, grouped on one side of the flower ................... 3 Leaves 3–15 cm long, repeatedly pinnate; flowers fascicled; tepals 3– 5; stamens 3 or 4 ................................................................. M. capillaceum Leaves 1–5 cm long, repeatedly forked, sometimes cuneate with forked lobes at tip; flowers solitary; tepals 3 or 4; stamens 2 .... M. aeruginosum

Marathrum aeruginosum P. Royen, Meded. Bot. Mus. Herb. Rijks Univ. Utrecht 107: 84, 132, pl. 8, figs. 1, 2. 1951. Small herb with opposite shoots along branched, compressed, irregular base 2–10 mm diameter; leaves repeatedly forked or cuneate with forked lobes apically, 1–5 cm long; flowers solitary, tepals 3 or 4, stamens 2. Rapids, 100–200 m; Amazonas (Raudales de Santa Bárbara del Orinoco). Endemic. ◆Fig. 248. Marathrum capillaceum (Pulle) P. Royen, Meded. Bot. Mus. Herb. Rijks Univ. Utrecht 107: 86. 1951. —Lophogyne capillacea Pulle, Enum. Vasc. Pl. Surinam 194, t. 8. 1906. Small to medium-sized herb with fleshy, thalloid base ca. 3.5 diameter and 1.5 cm thick; leaves 3–15 cm long, repeatedly pinnate, ultimate divisions triangular to lanceolate; flowers fascicled, tepals 3–5, sta-

mens 3 or 4. Rapids and waterfalls, 50–300 m; Bolívar (Río Caura, Río Erebato). Suriname, French Guiana, Brazil. Marathrum squamosum Wedd. in A. DC., Prodr. 17: 54. 1873. Small to medium-sized herb with base to 1 cm wide and 0.5 cm tall; leaves 2.5–40 cm long, repeatedly pinnate or pinnate and pinnae repeatedly forked, ultimate divisions filiform and 2–18 mm long; flowers solitary or fascicled, pedicel to 8 cm long, tepals 8–12, stamens 4–25. Colombia, southern Venezuela, French Guiana, northern Brazil; 2 varieties, 1 in Venezuela. The second variety described is var. phellandrifolium (Engl.) P. Royen, which is known from northern Brazil. M. squamosum var. squamosum Marathrum squamosum var. spruceanum Wedd. in A. DC., Prodr. 17: 54. 1873.

Mourera 309

Fig. 248. Marathrum aeruginosum

Rapids, 100–200 m; Amazonas (San Carlos de Río Negro). Northern Brazil (upper Rio Negro, upper Rio Vaupés). Marathrum utile Tul., Ann. Sci. Nat. Bot. sér. 3, 11: 95. 1849. Small to medium-sized herb with shoots opposite along the roots; leaves 3–35 × 1–5.5 cm, entire to pinnately divided, cuneate to

Fig. 249. Marathrum utile

spathulate, rhombiform when young, fewlobed, apically with many lanceolate segments to 5 mm long; flowers solitary or fascicled, pedicel widened at the top, 1–4 cm long, tepals and stamens 5–8 each; fruit 8ribbed. Rapids, 100–200 m; Amazonas (Raudales de Santa Bárbara del Orinoco). Barinas, Mérida, Trujillo; Costa Rica, Colombia. ◆Fig. 249.

4. MOURERA Aubl., Hist. Pl. Guiane 582, t. 233. 1775. Small to very large, conspicuous herbs, stemless or with a short stem formed by the fusion of leaf bases. Leaves distichous, small to very large, elliptic in outline and then with a strongly fimbriate margin, or cuneate in outline and pinnately lobed or repeatedly forked, sometimes very coarse with many rigid outgrowths on upper surface. Flowers in 2-sided spike-like inflorescences, few- to many-flowered, rarely very short or reduced to 1 flower. Flowers alternating with decurrent bracts, the latter on both sides with a distinct wing. Tepals 5–20, free, in a complete whorl. Stamens 5–35 in a complete or incomplete whorl, sometimes in 2 complete whorls. Ovary 6–14-ribbed. Capsule with 2 equal valves, each 3- or 5-ribbed. Colombia, Venezuela, Guyana, Brazil, Argentina; 6 species, 1 in Venezuela. Mourera fluviatilis Aubl., Hist. Pl. Guiane 582, t. 233. 1775. Large herb; leaves polymorphic, entire to pinnatisect, 8–200 × 2–30 cm with rigid outgrowths on upper surface; inflorescences erect, sword-like, 5–65 cm tall, 2–30 cm wide, with 40–90 pink to violet flowers; tepals 16–

20, scale-like to lanceolate, < 0.5 mm wide; stamens 20–35, 6–12 mm long, in 1 or 2 whorls; each valve of capsule with 3 or 5 ribs; seeds ca. 2000 per fruit. Rapids and waterfalls, 30–400 m; Bolívar (Raudal Budare on Río Suapure, Río Caroní above San Félix and at Salto Revaloso, upper Río Caura, Río

310

P ODOSTEMACEAE

Toro), Amazonas (Río Yureba at Salto Yureba). Guyana, Suriname, French Guiana, Brazil, Bolivia. ◆Fig. 250. Mourera fluviatilis is the largest-leaved species in the Podostemaceae, and it is distinctive in its large, erect, flattened inflorescence.

Fig. 250. Mourera fluviatilis

5. OSERYA Tul. & Wedd., Ann. Sci. Nat. Bot. sér. 3, 11: 105. 1849. Small, annual or perennial herbs. Roots elongate, prostrate, flattened. Leaves distichous, blade repeatedly forked, sometimes with a distinct sheath. Flowers axillary, borne singly, zygomorphic, pedicellate, erect, covered by a tubular spathella. Tepals 2 or 3, scale-like. Stamen 1, deciduous; anther basifixed, dehiscing extrorsely to laterally. Ovary 2-locular, with 2 unequal carpels. Stigmas 2, free or basally connate, conical; ovules numerous. Capsules with 2 unequal valves, one of the valves sometimes caducous, 6–14-ribbed, suture margins thickened. Seeds numerous. Mexico, Venezuela, Guyana, Suriname, French Guiana, northern Brazil; 7 species, 1 in Venezuela. Oserya was first reported from Venezuela by J. Velásquez (Plantas Acuáticas Vasculares de Venezuela, p. 208. 1994), and the presence of the genus in the flora area is based on this report (no specimens were seen).

Rhyncholacis 311

Fig. 251. Oserya perpusilla

Oserya perpusilla (Went) P. Royen, Meded. Bot. Mus. Herb. Rijks Univ. Utrecht 119: 220, pl. 1, figs. 15–24. 1954. —Apinagia perpusilla Went in Pulle, Verh. Kon. Ned. Akad. Wetensch., Afd. Natuurk., Tweede Sect. 2, 16: 43, t. 1, 2, fig. 9, t. 12. 1910.

Tiny plant, ca. 3 cm long; leaves cuneate, distally a few times forked, 1–2 cm long; petioles 5–10 mm wide; flowers white; pedicels 4–14 mm long; stamen 1–2 mm long; ovary ellipsoid, 12-ribbed; stigmas free. Rapids, ca. 100 m; Bolívar (Río Cuyuní). Guyana, Suriname, French Guiana. ◆Fig. 251.

6. RHYNCHOLACIS Tul., Ann. Sci. Nat. Bot. sér. 3, 11: 95. 1849. Small to large herbs. Leaves distichous, repeatedly forked or pinnate with the lobes sometimes repeatedly forked, or simple and then always cuneate in outline and palmately lobed to parted with the lobes strongly dissected at the tip. Flowers solitary or fascicled. Tepals 3–20, in a complete or incomplete whorl, or all at one side of the flower. Stamens 4–25, in 1 or 2 complete whorls, or all at one side of the flower. Ovary with 2 equal carpels, midrib of each carpel winged (from the extension of the stigma bases). Stigmas 2, often rigid or beaked (rostrate), subulate or clavate, 3-sided or terete at base, with the midvein passing into the midrib of the fruit making the valves winged; capsule usually 6(8)-ribbed. Colombia, Venezuela, Suriname, French Guiana, Brazil; ca. 25 species, 7 in Venezuela, all in the flora area. Rhyncholacis can be recognized in flower and fruit by the usually stiff, beaked stigmas with midribs that descend into the winged midrib of the capsule. Although some species of Rhyncholacis lack the rostrate style, they have the characteristic winged midrib of the capsule. Key to the Species of Rhyncholacis 1. 1.

Stamens 2–6; leaves 4–9 cm long ............................................... R. oligandra Stamens 6–30; leaves 3–80 cm long .......................................................... 2

312

2(1). 2. 3(2). 3. 4(2). 4. 5(4). 5. 6(5).

6.

P ODOSTEMACEAE

Leaves repeatedly forked, sometimes repeatedly pinnate when young ................................................................................................................ 3 Leaves repeatedly pinnate both when young and mature ....................... 4 Anthers with 2 acute tips, the connective elongate beyond the anther cells ............................................................................................ R. coronata Anthers obtuse, the connective not elongate beyond the anther cells ........................................................................................ R. hydrocichorium Ultimate segments of leaves flattened, not filiform .................. R. applanata Ultimate segments of leaves filiform ........................................................ 5 Primary pinnae of leaves decurrent with a wing along the rachis of the leaves ....................................................................................... R. divaricata Primary pinnae of leaves not decurrent along the rachis of the leaves ................................................................................................................ 6 Midrib of ovary winged, but the 4 other ribs barely visible; leaf rachis subfiliform, abruptly widened at base; primary pinnae to 12 cm long; pedicel of fruit 5–6 cm long .................................................. R. flagellifolia Midrib of ovary winged, with the 4 other ribs clearly visible; leaf rachis terete, but subfiliform toward tip, gradually widening toward base; primary pinnae to 25 cm long; pedicel of fruit 8–15 cm long ................................................................................................ R. penicillata

Rhyncholacis applanata K.I. Goebel, Pflanzenbiolog. Schilder. 2: 377, t. 29, figs. 1, 2. 1893. Venezuela, Guyana; 2 varieties, 1 in Venezuela. The other variety described for this species is var. laxipinnata P. Royen, supposedly endemic to Guyana. R. applanata var. applanata Medium-sized herb with branched or unbranched base of the stem 1–5 cm high; leaves repeatedly pinnate, 0.3–1.7 cm wide, primary pinnae 1.5–15 cm long, ultimate divisions lanceolate to linear, < 1 mm long; flowers fascicled, pedicels 4–8 cm long, tepals and stamens each 8–20; styles marcescent, ca. 1.5 mm long. Rapids and waterfalls, 400– 1200 m; Bolívar (headwaters of Cerro Venamo, Río Aparamán at Salto Yuray-merú, Río Kamoyrán in Gran Sabana, northeastern slopes of Sierra de Maigualida), Amazonas (Río Coro Coro near Yutajé). Guyana. Rhyncholacis coronata P. Royen, Meded. Bot. Mus. Herb. Rijks Univ. Utrecht 107: 97, 133, pl. 9, figs. 8, 9. 1951. Medium-sized herb with irregular base of stem ca. 3 cm high and 1–2 cm wide; leaves repeatedly pinnate or forked when young, 10–20 cm long; flowers fascicled, pedicels 1–3 cm long; tepals 8–10, stamens 8–18. Water-

falls, 400–500 m; Bolívar (Río Icabarú at Salto Yumariba). Guyana. ◆Fig. 252. Rhyncholacis divaricata Matthiesen, Biblioth. Bot. 15(68): 19, 49, t. 8, figs. 56–60. 1908. Small herb, the base of the stem irregular, unbranched, ca. 1 cm long; leaves repeatedly pinnate, 10–15 cm long, ultimate divisions filiform and ca. 2.5 mm long; flowers solitary or in fascicles of 2–4, pedicels to 2.5 cm long; tepals and stamens 6–9 each. Rapids and waterfalls, 50–600 m; Bolívar (Río Caroní above San Félix and at Salto Curapacay, Río Tírica), Amazonas (Río Padamo). Endemic. Rhyncholacis flagellifolia P. Royen, Meded. Bot. Mus. Herb. Rijks Univ. Utrecht 107: 108, 136, pl. 9, figs. 12–16. 1951. Small to medium-sized herb; base of stem cuneiform, compressed, branched, 0.5–5 cm wide; leaves repeatedly pinnate, 5–35 cm long, ultimate divisions filiform, 0.5–3 mm long; flowers white to pink, pedicels 1–3 cm long; tepals and stamens 6–9 each. Rapids, 50–1000 m; Bolívar (Río Caroní, Río Uairén in Río Kukenán basin). Northern Brazil. ◆Fig. 253. Rhyncholacis hydrocichorium Tul., Ann. Sci. Nat. Bot. sér. 3, 11: 95. 1849.

Rhyncholacis 313

Fig. 252. Rhyncholacis coronata

Fig. 254. Rhyncholacis penicillata

Fig. 253. Rhyncholacis flagellifolia

Fig. 255. Rhyncholacis hydrocichorium

314

P ODOSTEMACEAE

Small herb; leaves cuneate to elliptic, 2.5– 20 cm long, repeatedly forked, pinnae to 25 mm wide, ultimate divisions filiform; flowers fascicled, pedicels with 2 narrow wings, 2–11 cm long; tepals and stamens each 7–14, pinkish mauve; pedicels to 6–16 cm long in fruit. Rapids and waterfalls, 50–400 m; Bolívar (Río Aza west-southwest of Chiguao, Río Cuyuní, Salto Guaiquinima on Río Paragua, near San Martín de Turumbán). Guyana. ◆Fig. 255. Rhyncholacis oligandra Wedd. in DC., Prod. 17: 57. 1873. Venezuela, Guyana, Brazil(?); 2 varieties, 1 in Venezuela. The second variety described for this species is var. tenella P. Royen, which occurs in Guyana. R. oligandra var. oligandra Small, delicate herb; leaves repeatedly

pinnate or forked, 1–6 cm long; flowers solitary or fascicled, pedicel 0.5–2.5 cm long; tepals and stamens each 5 or 6. Rapids and exposed rocks, 100–200 m; Amazonas (Río Casiquiare region including Río Pasiba and Río Pasimoni, possibly San Carlos de Río Negro). Northern Brazil(?). Rhyncholacis penicillata Matthiesen, Biblioth. Bot. 15(68): 16, 48. 1908. Medium-sized herb; base of stems thick and robust, leaves repeatedly pinnate, 15–50 cm long, ultimate divisions filiform; flowers in fascicles of ca. 20, pedicels 3.5–10 cm long; tepals and stamens 7–10 each. Rapids and waterfalls, 50–1200 m; Bolívar (Río Aro, Río Caroní above San Félix, Río Parupa, Río Tírica above Techiné-merú, Salto Hacha near Canaima, Salto Uku-merú below Kamarata), Amazonas (near Isla Carestia in Río Orinoco). Endemic. ◆Fig. 254.

7. TRISTICHA Thouars, Gen. Nov. Madagasc. 2. 1806. Dufourea Bory ex Willd., Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck. Gesammten Naturk. 3: 100. 1809, non Ach. 1809. Heavily branched, moss-like herbs growing in large mats. Leaves tristichous. Flowers solitary at the end of ordinary shoots or on brachyblasts, at first enclosed between 2 or 3 distinctly larger leaves. Perianth well developed, 3-merous, petaloid lobes free or connate at base. Stamen 1 (rarely 2). Ovary 3-carpellate, 3-locular; stigmas 3, oblong-elliptic, slightly connate at the base. Capsule 3-valved, each valve 3-ribbed. Seeds numerous. Mexico, Central America, Cuba, Colombia, Venezuela, Guyana, French Guiana, Brazil, Paraguay, Argentina, Uruguay, Africa, Madagascar, Indian Ocean islands; 1 species.

Fig. 256. Tristicha trifaria

Weddellina 315

Tristicha trifaria is the most widespread species in the family and can grow in full sun to deep shade. Kato et al. (M. Kato, Y. Kita, and S. Koi. Australian Systematic Botany 16: 177–183. 2003) used molecular phylogenies to determine that Tristicha is monotypic and that the previously recognized T. australis C. Cusset & G. Cusset from northern and western Australia belongs instead in the genus Malaccotristicha. Tristicha trifaria (Bory ex Willd.) Spreng., Syst. Veg. 1: 6, 22, 95. 1825 [1824]. —Dufourea trifaria Bory ex Willd., Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck. Gesammten Naturk. 5: 62. 1811. Dense, small, mat-forming herb on rocks

in rapids, 50–800 m; Delta Amacuro (Río Amacuro past Río Matanaima near Salto de Quebradero, Serranía de Imataca), Bolívar (Río Cuyuni, Río Icabarú, Río Kanarakuni, Salto Pará on Río Caura). Barinas, Carababo, Mérida; rest of distribution as in genus. ◆Fig. 256.

8. WEDDELLINA Tul., Ann. Sci. Nat. Bot. sér. 3, 11: 113. 1849. Small herbs in dense mats, of branched sterile shoots and unbranched fertile shoots each with 1 terminal flower (these can be clumped into dense fascicle-like clusters). Sterile shoots repeatedly forked, densely to subdensely covered with 2–6dentate scales and small alternate leaves; the leaves repeatedly forked and covered with scales, ultimate divisions with tufts of minute filaments in the axil of 1–6-dentate scales; stem of fertile shoot with a few scale-like leaves. Flowers actinomorphic,

Fig. 257. Weddellina squamulosa

316

P ODOSTEMACEAE

enveloped by scale-like leaves when young. Tepals 5(6), free or slightly connate at base. Stamens 5–25, in a single complete whorl. Ovary 2-carpellate, 6-ribbed; style elongate, with a globose, papillose stigma; ovules numerous in each locule. Capsule with 2 equal valves, each with 4–6 undulate ribs. Colombia, Venezuela, Guyana, Suriname, French Guiana, Brazil; 1 species. Weddellina is unique in the family in its prominent style and globular-papillose stigma and gynoecium with an apical septum but an incomplete septum in the center (I. Jäger-Zürn. Aquatic Botany 57: 151–182. 1997; see Fig. 257). Weddellina squamulosa Tul., Ann. Sci. Nat. Bot. sér. 3, 11: 113. 1849. Rapids and waterfalls, 50–500 m; Bolívar (Río Caroni at Macagua, Salto Yumariba on

Río Icabarú), Amazonas (near Isla Carestia in Río Orinoco, Raudales de Atures, Río Cataniapo, upper Río Orinoco). Distribution as in genus. ◆Fig. 257.

POLYGALACEAE by Gerardo A. Aymard C., Paul E. Berry, and Bente Eriksen Herbs, subshrubs, erect or scandent shrubs, lianas, or rarely small trees or saprophytes. Leaves simple, alternate, verticillate, or rarely opposite, sometimes reduced to scales, sometimes with 2 flat circular glands at the apex of petioles or basal portion of adaxial leaf blade; stipules absent. Inflorescences racemose or paniculate, axillary or terminal, sometimes a spike or a single flower. Flowers bisexual, zygomorphic or sometimes actinomorphic, 2-bracteolate; sepals 5, lobes ± free from base, unequal in zygomorphic flowers, the 2 inner ones large and often petaloid (“wings”), or tubular (Diclidanthera, Moutabea); petals 3 or sometimes 5, declinate, if 3, the lower petal usually keel-shaped, often appendaged, the upper 2 ligulate or ovate, usually adnate to the lower petal and connate to the staminal sheath, if 5, connate above the middle into a tube and adnate to the sepals, forming a hypanthium. Stamens 4–10; filaments fused with the corolla or subdiadelphous and then adnate to the lateral petal lobes; anthers 2- or 4-locular, dorsifixed or basifixed, erect or pendent, dehiscent by an apical pore or by longitudinal slits; disk present, intrastaminal, annular or developed into an adaxial gland. Ovary superior, 1–8locular; style simple, straight, curved, or geniculate; stigma capitate or bilobed; ovules usually 1 per locule, sometimes several, pendulous or axile. Fruit a capsule, drupe, berry, or samara. Seeds 1 per locule, rarely 2 (Bredemeyera, Polygala), pilose, sometimes glabrous, usually arillate; endosperm straight, abundant or scarce, usually oily. Cosmopolitan; ca. 17 genera and 1000 species, 7 genera and ca. 80 species in Venezuela, 7 genera and 63 species in the flora area.

Barnhartia 317

Key to the Genera of Polygalaceae 1. 1. 2(1). 2. 3(2). 3. 4(1). 4. 5(4). 5. 6(4). 6. 7(6). 7.

Herbs, subshrubs, erect shrubs, or small trees ........................................ 2 Scandent shrubs or lianas ......................................................................... 4 Small trees; corolla tubular, 5-merous; style straight; fruit a berry .................................................................................................. 5. Moutabea Herbs, subshrubs, or shrubs; corolla papilionaceous, 3-merous; style curved or geniculate; fruit a drupe, capsule, or samara ...................... 3 Shrubs; style geniculate; fruit a drupe or samara; seed 1 per fruit ................................................................................................... 4. Monnina Herbs or subshrubs; style curved; fruit a capsule; seeds 2 per fruit .................................................................................................... 6. Polygala Corolla papilionaceous, 3-merous, zygomorphic; style curved or geniculate; fruit a capsule or samara .............................................................. 5 Corolla with a short or large tube, 5-merous, actinomorphic or subzygomorphic; style straight; fruit a berry ............................................. 6 Filaments lacking nectaries; style curved; fruit a 1-seeded samara; seeds glabrous .................................................................................. 7. Securidaca Filaments with nectaries; style geniculate; fruit a 2-seeded capsule; seeds with long trichomes ............................................................ 2. Bredemeyera Filaments forming a tube; receptacle with an annular nectary .................................................................................................. 5. Moutabea Filaments free, adnate to the corolla tube or the petals; receptacle lacking nectaries ................................................................................................. 7 Corolla slightly zygomorphic, calyx with sepals ± free from the base; stamens 7 or 8; locules 2 or 3 ................................................ 1. Barnhartia Corolla actinomorphic, calyx with sepals connate and tubular in lower half; stamens 8 or 10; locules 4 or 5 ................................. 3. Diclidanthera

1. BARNHARTIA Gleason, Bull. Torrey Bot. Club 53: 297. 1926. Lianas or scandent shrubs. Leaves alternate, petiolate; blades subcoriaceous, narrowly elliptic-oblong; margins entire. Inflorescence of 1–3 simple or branched spikes from the upper leaf axils, 2–5 cm long. Flowers slightly zygomorphic, disk absent. Sepals 5, imbricate, inserted on the summit of the hypanthium, distinct, pubescent; petals 5, valvate in bud, inserted on the summit of the hypanthium, 4 somewhat connate at the base into 2 pairs, the fifth free and distinct, densely hirsute within, clawed, forming a short tube. Stamens 7 or 8, epipetalous, 5 opposite the petals and 2 or 3 alternate with them (each pair of petals bearing 2 opposite stamens and 1 alternate one, the odd petal bearing 1 opposite and sometimes 1 lateral alternate stamen); filaments inserted near the summit of the claw, short, flat; anthers flattened-ellipsoid, 2-locular, opening by a tangential cleft, the introrse valves shriveled and pendent. Ovary with a broad base, slightly flattened, 2- or 3-locular; ovules 1 per locule, pendulous; style straight, pilose; stigma capitate, 2-lobed. Fruit a berry. Southern Venezuela, Guyana, Suriname, French Guiana, Amazonian Brazil; 1 species.

318

P OLYGALACEAE

Barnhartia floribunda Gleason, Bull. Torrey Bot. Club 53: 297. 1926. —Kurawa-dek (Pemón). Canopy liana; petals white; fruits globose, glabrous, yellow-orange when ripe. Lower montane forests, 300–1000 m; Bolívar (base of Cerro Camarón, Cerro Ichún, El Polaco mining camp, Río Erebato, Serranía Marutaní). Guyana, Suriname, French Guiana, Brazil (Amazonas). ◆Fig. 258.

Fig. 258. Barnhartia floribunda

2. BREDEMEYERA Willd., Ges. Naturf. Freunde Berlin Neue Schriften 3: 412, t. 6. 1801. Catocoma Benth., J. Bot. (Hooker) 4: 101. 1842. Scandent shrubs or lianas. Leaves alternate, eglandular, petiolate or subsessile. Inflorescences paniculate. Flowers zygomorphic; sepals 5, ± free from the base, the 2 inner ones larger and petaloid (wings); corolla papilionaceous, petals 5, white, yellow, or greenish, united at the base with the staminal tube, the lower petal boat-shaped (keel), lacking a crest. Stamens 8, forming a sheath adnate to the upper petal; anthers basifixed, opening by an apical pore. Style short, geniculate; stigma bilobed, carpels 2; 1 ovule per locule. Fruit a 2-valved capsule, elongate, cuneiform, loculicidally dehiscent. Seeds oblong, covered with long trichomes; aril short or absent. Neotropics (Mexico to Argentina); 20 species, 7 in Venezuela, 6 of these in the flora area. Key to the Species of Bredemeyera 1.

1.

Leaf venation camptodromous (the secondary veins curve toward the margin without forming loops), the blades sparsely appressed-pubescent to puberulent on the lower surface; inflorescence and pedicels gray-tomentose, glabrescent when mature; pedicels 2–3 mm long; flowers 6–8 mm long; fruit 25–30 mm long .......................... B. floribunda Leaf venation camptodromous or brochidodromous (secondary veins curve markedly before reaching the margin and unite with the vein above forming a loop), the blades glabrescent to pubescent on the lower

Bredemeyera 319

2(1). 2. 3(2). 3. 4(2). 4.

5(4). 5.

surface; inflorescence and pedicels glabrous to densely tomentose; pedicels and flowers smaller than above; fruit < 25 mm long ............. 2 Leaves coriaceous, strongly reticulate on both surfaces .......................... 3 Leaves chartaceous or subcoriaceous, prominulous-reticulate on both surfaces .................................................................................................. 4 Leaves obovate, apex emarginate; style pilose at base; fruit 9–10 mm long, subgloboid or obovoid ..................................................... B. altissima Leaves elliptic to ovate-elliptic, apex acuminate; style glabrous at base; fruit 12–20 mm long, spathulate in outline ................................ B. lucida Upper surface of leaves dull, the apex emarginate; inflorescences and pedicels densely yellow-pubescent ..................................................B. sp. A Upper surface of leaves shiny, the apex acute to acuminate; inflorescences and pedicels puberulent with patent trichomes or tomentose (not yellow) ............................................................................................. 5 Leaves ovate; petioles densely strigulose; flowers 2–3 mm long; sepals sparsely puberulent within .................................................... B. myrtifolia Leaves lanceolate-elliptic to elliptic; petioles tomentose; flowers 1–2 mm long; sepals glabrous within except at the base .................... B. densiflora

Bredemeyera altissima (Poepp. & Endl.) A.W. Benn. in Mart., Fl. Bras. 13(3): 50. 1874. —Catocoma altissima Poepp. & Endl., Nov. Gen. Sp. Pl. 3: 65. 1845. Liana; flowers greenish white. Semideciduous, white-sand shrublands, lowland evergreen forests, 50–200 m; Bolívar (Caicara to Ciudad Bolívar road, Represa de Guri, Río Parguaza), Amazonas (Caño San Miguel, Capibara, Puerto Ayacucho to El Burro road, Río Baría, San Carlos de Río Negro). Apure, Táchira; Colombia, Guyana, Peru, Brazil, Bolivia.

Benth., J. Bot. (Hooker) 4: 102. 1842. —Chinák (Pemón). Liana; flowers white with yellow. Semideciduous, evergreen lowland and montane forests, 50–1100 m; Bolívar (Cerro Guaiquinima, Ciudad Bolívar, Represa de Guri, Río Kukenán, Río Paragua), Amazonas (San Carlos de Río Negro). Anzoátegui, Aragua, Distrito Federal, Falcón, Guárico, Lara, Miranda, Monagas, Nueva Esparta, Portuguesa, Sucre, Táchira, Trujillo, Zulia, Yaracuy; Colombia, Guyana, Peru, Brazil, Bolivia, Paraguay. ◆Fig. 259.

Bredemeyera densiflora A.W. Benn. in Mart., Fl. Bras. 13(3): 52. 1874. Colombia, Venezuela, Guyana, Suriname, Peru, Brazil, Bolivia; 2 varieties, 1 in Venezuela.

Bredemeyera lucida (Benth.) Klotzsch ex Hassk., Ann. Mus. Bot. LugdunoBatavum 2: 189. 1852. —Catocoma lucida Benth., J. Bot.(Hooker) 4: 103. 1841. —Bejuco colorado, Bejuco de bocachico, Bejuco puguacari. Liana; flowers greenish white. Semideciduous to evergreen lowland and montane forests, granitic outcrops, 100–600 m; Delta Amacuro (Los Castillos), widespread in Bolívar and Amazonas. Apure, Cojedes, Falcón, Guárico, Mérida, Miranda, Sucre, Táchira, Trujillo, Zulia; Mexico, Guatemala, Belize, Honduras, Panama, Colombia, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia.

B. densiflora var. glabra A.W. Benn. in Mart., Fl. Bras. 13(3): 52. 1874. Liana; flowers greenish white. Evergreen lowland to lower montane forests, 100–800 m; Bolívar (Cerro Guaiquinima), Amazonas (between Caño Cotúa and Cerro Yapacana, San Fernando de Atabapo, Santa Bárbara del Orinoco). Táchira; Colombia, Guyana, Suriname, Peru, Brazil, Bolivia. Bredemeyera floribunda Willd., Ges. Naturf. Freunde Berlin Neue Schriften 3: 12. 1801. —Catocoma floribunda

Bredemeyera myrtifolia A.W. Benn. in Mart., Fl. Bras. 13(3): 50. 1874. —Bejuco

320

P OLYGALACEAE

carena, Bejuco de curare, Miel de pajarito. Bredemeyera parviflora A.W. Benn. in Mart., Fl. Bras. 13(3): 51. 1874. Liana; flowers greenish white. Evergreen lowland forests, 100–200 m; Amazonas (upper Río Autana, Río Cuao, San Carlos de Río Negro, San Fernando de Atabapo). Táchira, Zulia; Colombia, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 260.

Bredemeyera sp. A Liana; flowers pale green; fruit green. Evergreen lowland to lower montane forests, 200–400 m; Bolívar (Río Asa, Sierra Auraima west of Río Paragua). This taxon is based on Blanco 689 (NY, VEN) and Steyermark 90830 (NY, VEN).

Fig. 259. Bredemeyera floribunda

Fig. 260. Bredemeyera myrtifolia

Diclidanthera 321

3. DICLIDANTHERA Mart., Nov. Gen. Sp. Pl. 2: 139. 1826 [1827]. Small scandent trees or lianas. Leaves alternate, petiolate, with pellucid dots, sometimes with a cupulate gland at base of leaf blade or at the nodes; stipules absent or early deciduous. Inflorescences axillary or terminal racemes or panicles. Calyx and corolla united at the base; calyx 5-lobed, tubular, tube ± as long as lobes, the narrow divisions imbricate, lobes caducous in fruit; corolla elongate-cylindric, actinomorphic, 5-lobed, connate basally, white to greenish, lacking nectaries. Stamens 8 or 10, adnate to corolla tube, anthers sessile or nearly so, basifixed, glabrous. Ovary superior, globose, 4- or 5-locular; ovules 1 per locule, anatropous, pendulous; style straight; stigma capitate. Fruit a globose berry. Seeds pubescent. Colombia, Venezuela, Peru, Brazil, Bolivia; ca. 9 species, 5 in Venezuela, 4 of these in the flora area. 5 mm

1 cm 1 cm

5 cm Fig. 261. Diclidanthera wurdackiana

322

P OLYGALACEAE

Key to the Species of Diclidanthera 1. 1. 2(1). 2. 3(2). 3.

Leaves appressed- or erect-pubescent to pilose on the lower surface ............................................................................................ D. wurdackiana Leaves glabrous to glabrescent on lower surface ..................................... 2 Leaves coriaceous, papillose on lower surface; inflorescences branched; fruit 5–7 mm diameter ................................................................... D. sp. A Leaves chartaceous to subcoriaceous, smooth on lower surface; inflorescences unbranched; fruit 10–20 mm diameter ..................................... 3 Calyx lobes oblong, 6–7.5 × 2.5–3 mm, stamens 8 ...................... D. octandra Calyx lobes obovate, 9–12 × 1–1.5 mm, stamens 10 .............. D. bolivarensis Diclidanthera bolivarensis Pittier, Bol. Soc. Venez. Ci. Nat. 7: 313. 1942. Diclidanthera orinocensis Pittier, Bol. Soc. Venez. Ci. Nat. 7: 313. 1942. Liana; flowers white to greenish. Semideciduous to evergreen lowland forests, 100–200 m; Bolívar (Caicara to El Burro road, Ciudad Bolívar, El Temblador on middle Río Caura, Río Suapure), Amazonas (Puerto Ayacucho, Río Cataniapo). Anzoátegui, Apure; Colombia, Peru, Brazil. ◆Fig. 262. Diclidanthera octandra Gleason, Phytologia 1: 110. 1934. Liana; flowers white to greenish. Evergreen lowland forests, 100–200 m; Amazonas (Isla Ratón). Peru, Brazil, Bolivia. Diclidanthera wurdackiana Aymard & P.E. Berry, Biollania (Edición Especiál) 6: 191. 1997.

Fig. 262. Diclidanthera bolivarensis

Monnina 323

Liana; flowers whitish yellow; fruit green. Evergreen lowland to lower montane forests, 100–600 m; Bolívar (Los Pijiguaos), Amazonas (Río Putaco-Río Ocamo, 15 km southeast of San Fernando de Atabapo, Trapichote). Endemic. ◆Fig. 261.

Diclidanthera sp. A Liana; fruit green. Evergreen lowland forests, 100–200 m; Amazonas (San Carlos de Río Negro). Endemic. This species is based on Christenson et al. 1419 (NY, US, VEN).

4. MONNINA Ruiz & Pav., Fl. Peruv. 1: 169. 1798. by Bente Eriksen Shrubs and treelets, erect or scrambling. Stems and branches terete, glabrous, solid or fistulous, glabrescent to densely hirtellous. Leaves alternate, short-petiolate to subsessile, with or without nodal glands; blades entire, rarely undulate, glabrescent to hirtellous; venation brochidodromous and ± conspicuous. Inflorescences terminal or axillary, simple or compound racemes. Flowers each subtended by a bract and 2 bracteoles, irregular, bisexual. Outer sepals 3, usually shorter than the mature flower, herbaceous to carnose, free or the lower 2 ± connate; inner sepals (wings) 2, large, lateral, petaloid, blue to purple; petals 3, the lower (keel) carinate, blue to purple with a yellow tip or yellow throughout, the upper 2 united in their lower part with the filament sheath (see below), but free from each other, forming a conduplicate petal-stamen sheath, the upper lobes blue to purple. Stamens 8, diadelphous; filaments fused for most or rarely all of their length into a filament sheath, to the lateral margins of which the upper petals are adnate; anthers 2-locular, opening by a short, curved slit. Ovary 1-locular, subtended by a gland extending toward the adaxial side; style compressed, geniculate to curved, apically unequally cleft, the posterior lobe with a sessile or ± stalked capitate stigma, the anterior lobe sterile, often forming a tooth. Fruit a drupe, blue or black at maturity. Seeds 1 per fruit; endosperm present. Southern U.S.A., Mexico, Central America, Colombia, Venezuela, Ecuador, Peru, Bolivia; ca. 150 species, ca. 15 in Venezuela, 1 of these in the flora area. Monnina cacumina N.E. Br., Trans. Linn. Soc. London, Bot. ser. 2, 6: 19. 1901. Monnina duidae S.F. Blake, Bull. Torrey Bot. Club 58: 381. 1931. —Monnina cacumina var. duidae (S.F. Blake) Steyerm., Acta Bot. Venez. 2: 232. 1967. Monnina uaipanensis Wurdack, Mem. New York Bot. Gard. 9: 479. 1957. Erect or scandent shrub 1–6 m tall; leaves coriaceous, deep green on upper surface, pale green on lower surface; bracts narrow, longer than the buds; outer lower sepals partly connate, acute; drupe wine red, becoming bluish black at maturity. Exposed or in forests, on rocks, tepui summits, 1300–2800 m; scattered in Bolívar and Amazonas. Guyana, Brazil (Amazonas: Serra da Neblina; Roraima: Serra do Sol). ◆Fig. 263. Steyermark suggested that Monnina duidae and M. uaipanensis were based on very subtle characters and were better con-

Fig. 263. Monnina cacumina

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sidered varieties of a single species. The view taken here is that the differences are just

part of the natural variation, so separate entities are not maintained.

5. MOUTABEA Aubl., Hist. Pl. Guiane 679, t. 274. 1775. Erect or scandent shrubs, lianas, or rarely small trees. Leaves alternate, petiolate; blades thick, dark to black when dry, coriaceous, sometimes subcoriaceous to membranous, the venation usually obscure, glabrous and with pellucid dots; margins entire; stipules absent. Inflorescences short, axillary, paniculate. Flowers slightly zygomorphic, white or yellow. Calyx adnate to the corolla into a tube; sepals 5, imbricate, ± equal, caducous in fruit; corolla a long tube, petal lobes 5, white or yellow, fragrant, imbricate, subequal except the lower one slightly boat-shaped, connate basally. Stamens 8, joined into 2 groups of 4 on the margin of an oblique sheath that is adnate to the corolla tube; anthers sessile, basifixed. Ovary superior, 4- or 5locular, surrounded by a nectary disk; ovules 1 per locule, pendulous; style filiform, straight, slightly dilated apically. Fruit a leathery berry or drupe, globose, with a fleshy pericarp, 2–5(rarely more)-seeded, edible. Seeds lacking an aril, without endosperm, pubescent or glabrous, shiny. Costa Rica, Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; ca. 10 species, 5 in Venezuela, all in the flora area. As the treatment below shows, Moutabea is badly in need of a taxonomic revision. Key to the Species of Moutabea 1. 1.

2(1).

2.

3(2). 3. 4(3).

4.

Leaf blades membranous, venation evident on the upper surface, the midrib flat on the lower surface; petioles ca. 1 mm wide ............. M. sp. A Leaf blades subcoriaceous to rigid-coriaceous, venation obscure on the upper surface, the midrib raised on the lower surface; petioles 1–5 mm wide ........................................................................................................ 2 Plants growing in scrubby savannas on sandy soils; leaves widely ovate to ovate or elliptic; inflorescences 1.5–2 cm long; flowers 5–7 mm long; calyx glabrous externally ............................................................... M. sp. B Plants growing in evergreen forests; leaves obovate, oblanceolate, ovate, elliptic-ovate, or oblong; inflorescences > 2.5 cm long; flowers 10– 22 mm long; calyx puberulent externally ............................................. 3 Flowers 18–24 mm long; sepals acute, not ciliate ............ M. aff. chodatiana Flowers 10–15 mm long; sepals oblong to narrowly ovate, ciliate .......... 4 Leaves 6–18 cm long, obovate or ovate, base cuneate to acute; inflorescence sparsely pubescent, pedicels 2–2.5 mm long, bracts adjacent to the calyx widely ovate .......................................................... M. guianensis Leaves 15–30 cm long, elliptic to oblong, sometimes oblanceolate, base attenuate; inflorescence glabrescent, flowers sessile, bracts adjacent to the calyx ovate-lanceolate ........................................................ M. aculeata

Moutabea aculeata (Ruiz & Pav.) Poepp. & Endl., Nov. Gen. Sp. Pl. 2: 62. 1838. —Acosta aculeata Ruiz & Pav., Syst. Veg. Fl. Peruv. 1: 5, t. 4, fig. A. 1798. —Ojo de venado.

Moutabea longifolia Poepp. & Endl., Nov. Gen. Sp. Pl. 2: 62, t. 186. 1838. Liana or small tree; flowers white; fruit orange, aril white, edible. Evergreen lowland forests, 100–400 m; widespread in Bolívar

Moutabea 325

and Amazonas. Mérida; Costa Rica, Panama, Colombia, Ecuador, Peru, Brazil, Bolivia. It is not clear if Moutabea longifolia should be considered a synonym of M. aculeata, but here we follow Wendt (T. Wendt. Lundellia 3: 6–12. 2000) and Eriksen et al. (Bente Eriksen, Bertil Ståhl, and Claes Persson. Polygalaceae. In: Flora of Ecuador No. 65. 2000).

Moutabea aff. chodatiana Huber, Bol. Mus. Paraense Hist. Nat. Ethnogr.3: 426. 1902. —Ojo de venado. Liana; flowers white. Evergreen lowland forests, 100–200 m; Bolívar (Río Villacoa), Amazonas (Río Casiquiare between Solano and mouth of Río Pasimoni, Río Siapa). Brazil (Pará, Roraima).

Fig. 264. Moutabea guianensis

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Moutabea guianensis Aubl., Hist. Pl. Guiane 680, t. 274. 1775. —Fruto de macaco, Ojo de venado, Ojo de zamuro. Liana or small tree; flowers white; fruit orange, aril white, edible. Evergreen lowland to montane forests and shrublands, 100–800 m; Bolívar (167 km south of Caicara del Orinoco, La Escalera, Río Botanamo, upper Río Caura), Amazonas (lower slopes of Cerro Aracamuni, Maroa, Río Baría, Río Casiquiare, Río Cataniapo, Río Cuao, Río Cunucunuma, Río Emoni, Río Matacuni, Río Ocamo, Río Siapa, Río Sipapo, Río Yatúa). Apure, Aragua, Zulia; Guyana, Suriname, French Guiana, Brazil. ◆Fig. 264. Moutabea sp. A. —Bejuco parásito. Liana; flowers white; fruit yellow. Evergreen lowland forests, 100–400 m; Delta Amacuro (Río Cuyubini basin), Bolívar (20–

25 km southwest of El Manteco, Río Erebato), Amazonas (Río Cunucunuma, Río Matacuni). Brazil (Amazonas). This taxon is based on Liesner & González 5969 (MO, NY, VEN), Boom & Marín 10478 (NY, PORT), Stergios & Velázco 14270 (MO, PORT, TFAV, VEN), Steyermark 87617 (VEN), and Peréz & Sosa 338 (MO, TFAV, VEN). Moutabea sp. B Liana; flowers white, fruit yellow. Scrubby savannas on sandy soils, 100–200 m; Amazonas (Caño Cotúa at base of Cerro Yapacana, Río Asisa). Endemic. This taxon is based on Huber 1570 (NY, US, VEN), Huber 1710 (US, VEN), Huber 1810 (VEN), Huber 2000 (US, VEN), Huber 5938 (US, VEN), and Huber & Kral 7967 (NY, US, VEN).

6. POLYGALA L., Sp. Pl. 701. 1753. Herbs, less often subshrubs, or rarely shrubs. Roots often with odor of methyl salicylate (oil of wintergreen). Leaves alternate, opposite, or verticillate, short-petiolate or sessile. Inflorescences terminal or lateral racemes, spikes, sometimes contracted into heads. Sepals 5, unequal, the outer 3 free or sometimes 2 basally connate, the inner 2 large, petaloid, wing-like; corolla papilionaceous, petals 3, white, pink, reddish, greenish, blue, or purple, basally connate, the lower one keel-shaped, lobed, or occasionally unappendaged, the upper 2 ligulate to ovate, connate to the keel or adnate to the staminal sheath or both. Stamens 8; filaments connate nearly to the apex into a sheath split above and adnate to the upper petals, anther dorsifixed or basifixed. Ovary 2-locular; ovules solitary; style curved; stigma dilated, entire to 2-lobed, often tufted; nectary on receptacle annular, or none. Fruit capsular, compressed, obovate to orbicular, generally loculicidally dehiscent. Seeds usually 2, often pilose, sometime glabrous, mostly arillate, with or lacking endosperm. Cosmopolitan; ca. 500 species, ca. 30 in Venezuela, 24 of these in the flora area. Key to the Species of Polygala 1. 1. 2(1). 2. 3(2).

3.

Leaves 8–60 mm wide ................................................................................ 2 Leaves 0.1–7 mm wide ............................................................................... 4 Pedicels 4–6 mm long, flowers 15–25 mm long ......................... P. spectabilis Pedicels 1–3 mm long, flowers 2–10 mm long .......................................... 3 Leaves ovate to broadly ovate, coriaceous, strongly reticulate on both surfaces, blades and inflorescences lacking incurved-appressed trichomes; flowers blue; sepals lacking glandular trichomes at the margins; fruit 6–10 mm long, hirsute ........................................................... P. caracasana Leaves elliptic or elliptic-lanceolate, chartaceous or subcoriaceous, reticulate on both surfaces, blades and inflorescences with incurved-ap-

Polygala 327

4(1). 4. 5(4). 5. 6(5). 6.

7(6).

7. 8(5). 8.

9(8). 9. 10(9). 10. 11(10).

11.

12(10). 12. 13(12). 13. 14(9). 14.

pressed trichomes; flowers purple or greenish lavender; sepals with glandular trichomes at the margins; fruit 2–5 mm long, glabrous ..................................................................................................... P. violacea Leaves reduced to scales, scales 0.5–1 mm long sometimes lacking altogether; inflorescences subconical, 2–5 mm long ......................... P. subtilis Leaves not scale-like, > 2 mm long; inflorescences elongate, capitate, cylindric, conic-cylindric, or conical, > 6 mm long ................................... 5 Leaves verticillate all along the stems ..................................................... 6 Leaves alternate or verticillate only at the base and alternate or opposite at the top of the stem ............................................................................. 8 Stems glandular-pilose; leaves linear-acicular, not punctate on lower surface; seeds with hooked trichomes ........................................ P. glochidiata Stems glabrous or if pilose then lacking glandular trichomes; leaves elliptic, lanceolate, or lanceolate-elliptic, punctate on lower surface; seeds without hooked trichomes ..................................................................... 7 Stems winged; leaves shortly pedicellate, lanceolate or lanceolate-elliptic; outer sepal with 2 large glands on the back; capsule glabrous; seed with aril ................................................................................ P. asperuloides Stems not winged; leaves sessile, elliptic; outer sepal lacking glands; capsule sparsely appressed-pubescent; seed lacking aril .............. P. galioides Plants slender, 2–5 cm tall; upper sepal with a biglandular thickening at the base .................................................................................. P. microspora Plants more robust, 10–70 cm tall; upper sepal lacking a biglandular thickening at the base, or if glands present (P. asperuliodes, P. paniculata) the plants always > 5 cm tall ............................................ 9 Inflorescences compressed, densely flowered, cylindric, cylindric-ovoid, globose, or capitate, 7–18 mm wide .................................................... 10 Inflorescences elongate, loosely flowered, conic-cylindric or conical, 1– 6 mm wide ............................................................................................ 14 Leaves ovate to obovate or elliptic-lanceolate ........................................ 11 Leaves linear-acicular to linear-lanceolate ............................................. 12 Leaves ovate to obovate, not punctate; flowers 1.5–3 mm long, pink; capsules orbicular to orbicular-ovoid, 1.2–1.5 mm long, lacking orange glands .........................................................................................P. timoutou Leaves elliptic-lanceolate, sparsely punctate; flowers 4–5 mm long, greenish white; capsules oblong, 2.5–3 mm long, with orange glands ............................................................................................... P. timoutoides Inflorescence cylindric-ovoid; capsules suborbicular, lacking glands; seeds ellipsoid, aril present ............................................................. P. hygrophila Inflorescence capitate or globose; capsules ellipsoid or ovoid-oblong, orange, gland-dotted; seeds conical, lacking aril ................................... 13 Pedicels 0.4–1.5 mm long; capsules ellipsoid; seeds with a short ring of reddish brown trichomes at the base ................................... P. adenophora Pedicels 1.5–4.5 mm long; capsules ovoid-oblong; seeds lacking a short ring of reddish brown trichomes at the base ........................ P. longicaulis Leaves with erect and incurved trichomes on both surfaces; sepals with glandular trichomes at the margins .......................................... P. violacea Leaves glabrous on both surfaces, or if pubescent, lacking incurved tri-

328

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chomes; sepals lacking glandular trichomes at the margins ............. 15 15(14). Seeds conical ............................................................................................ 16 15. Seeds oblong, reniform, ellipsoid, or ovate ............................................. 20 16(15). Seeds bearing 7 or 8 appendages on the hilum, subulate, 3–4 mm long ............................................................................................ P. sanariopoana 16. Seeds lacking appendages on the hilum ................................................. 17 17(16). Capsules lacking orange glands; seeds not comose, aril 2-lobed ............................................................................................. P. trichosperma 17. Capsules bearing a double line of orange glands; seeds comose, aril not lobed, or absent .................................................................................... 18 18(17). Bracts glandular-ciliate in the margins; ovary with a tuft of trichomes at the top .................................................................................... P. longicaulis 18. Bracts not glandular-ciliate in the margins; ovary with a tuft of trichomes at the top .............................................................................................. 19 19(18). Leaves linear, ca. 5 mm long, epunctate; inflorescences 15–20 mm long; bracts gland-dotted ..............................................................P. savannarum 19. Leaves obovate or oblanceolate, 5–8 mm long, punctate; inflorescences 5– 14 mm long; bracts not gland-dotted ..................................... P. sipapoana 20(15). Stems pilose to densely pubescent .......................................................... 21 20. Stems glabrous or sparsely puberulent .................................................. 25 21(20). Stems lacking glandular trichomes ........................................................ 22 21. Stems with glandular trichomes ............................................................. 23 22(21). Leaves punctate; pedicels papillose; capsule ovate-oblong, papillose, lacking glands; inflorescence rachis with hooked trichomes; seeds lacking appendages .............................................................................. P. brevialata 22. Leaves epunctate; pedicels not papillose; capsule elliptic or ovate-elliptic, not papillose; inflorescence rachis lacking hooked trichomes; seeds with 2 appendages at the top ..................................................... P. violacea 23(21). Leaves with glandular trichomes on the margins; rachis of the inflorescence densely glandular-papillose, pedicels 0.1–0.4 mm long .......... P. blakeana 23. Leaves glabrous on the margins; rachis of the inflorescence glabrous or sparsely glandular, pedicels 0.6–1.4 mm long .................................... 24 24(23). Inflorescences 0.6–1.5 cm long, sparsely hirsute-glandular; outer sepals lacking glands; seeds lacking aril, glabrous ........................... P. spruceana 24. Inflorescences 5–20 cm long, glabrous, outer sepals bearing 2 glands basally; seeds with aril, appressed-pubescent ...................... P. paniculata 25(20). Stems winged-angled; leaves elliptic or obovate-elliptic; outer sepals with 2 glands on the back ............................................................ P. asperuloides 25. Stems lacking wings; leaves linear; outer sepals lacking glands on the back ...................................................................................................... 26 26(25). Pedicels winged; ovary with a tuft of appressed trichomes at the top; capsules with a row of glands ...................................................... P. tenella 26. Pedicels not winged; ovary without appressed trichomes at the top; capsules lacking a row of glands .............................................................. 27 27(26). Pedicels 1–1.5 mm long; bracts lanceolate; seeds lacking glandular trichomes, aril present ................................................................... P. appressa 27. Pedicels < 0.8 mm long; bracts ovate; seeds with hooked glandular trichomes with one gland at the tip; aril lacking ............................. P. exigua

Polygala 329

Polygala adenophora DC., Prodr. 1: 327. 1824. —Echipipin (Panare), Pariwámuranó (Pemón). Polygala incarnata Aubl., Hist. Pl. Guiane 739. 1775, non L. 1753. Polygala adenophora var. gracilis Chodat, Mém. Soc. Phys. Genève 31(2): 179. 1874. Polygala adenophora var. robusta Chodat, Mém. Soc. Phys. Genève 31(2): 179. 1874. Herb; flowers white or pink. Savannas, 50–1600 m; Bolívar (Altiplanicie de Nuria, Caicara, Ciudad Piar, El Paují, Gran Sabana, Kavanayén, Maniapure, Maripa, Río Asa, upper Río Paragua, Río Suapure, Río Tírica, base of Roraima-tepui, Santa Elena de Uairén, Urimán), Amazonas (Canaripó, Caño Coro Coro, Caño Pimichín, Caño San Miguel, Cerro Camani, Cerro Parú, Isla Ratón, La Esmeralda, Morganito on Río Orinoco, Puerto Ayacucho, Río Asisa, Río Autana, San Antonio del Orinoco, Santa Barbára del Orinoco). Cojedes, Guárico; Mexico, Guatemala, Belize, Honduras, Nicaragua, Colombia, Trinidad, Guyana, Suriname, French Guiana, Brazil. Polygala appressa Benth., J. Bot. (Hooker) 4: 100. 1842. Polygala appressa var. gracillima A.W. Benn. in Mart., Fl. Bras. 13(3): 39. 1874. Polygala appressa var. kavanayena Steyerm., Fieldiana, Bot. 22: 298. 1952. Polygala appressa subsp. cumbrensis Wurdack, Mem. New York Bot. Gard. 23: 120. 1972. Herb; flowers white or pale pink. Savannas, 100–1600 m; Bolívar (Auyán-tepui, Caicara, Canaima, Cerro Guiaquinima, Chirikayén, Ciudad Piar, El Manteco, widespread in Gran Sabana), Amazonas (Canaripó, Caño Caname, Caño Cotúa, Caño San Miguel, Caño Yagua, Cerro Cariche, Cerro Parú, base of Cerro Yapacana, Guarinuma, Río Guayapo, lower Río Pasimoni). Guyana, Suriname, French Guiana, Brazil. ◆Fig. 270. Polygala asperuloides Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 403. 1821 [1823]. Herb; flowers white or pink. Savannas, river edges, 100–1000 m; Bolívar (El Abismo,

Entreríos, Raudales Maijía on Río Paragua, Represa Guri, middle Río Caura, Santa Elena de Uairén). Apure, Falcón, Lara, Portuguesa; Mexico, Central America, Colombia, Ecuador, Brazil, Bolivia, Argentina. ◆Fig. 268. Polygala blakeana Steyerm., Fieldiana, Bot. 28: 298. 1952. Herb; flowers white. Savannas, edges of Mauritia palm swamps, 100–1200 m; Bolívar (Cerro Altamira, Cerro Bolívar, Ciudad Piar, Hato La Vergareña, lower Río Caroní, Uaipán-tepui), Amazonas (Río Manapiare, Río Parucito). Guárico, Monagas; Guyana, Suriname, French Guiana, Brazil. ◆Fig. 274. Polygala brevialata Chodat, Mém. Soc. Phys. Genève 31(2): 234. 1893. Polygala subsecunda S.F. Blake, Bull. Torrey Bot. Club 51: 87. 1924. Polygala aristeguietae Wurdack, Bol. Soc. Venez. Ci. Nat. 22: 3. 1961. Herb; flowers white and purple. Savannas, granitic outcrops, 100–300 m; Bolívar (between Ciudad Bolívar and Río Caroní, northeast of Maripa, Puerto Ordaz), Amazonas (Puerto Ayacucho). Apure, Cojedes, Distrito Federal, Falcón, Guárico, Monagas; Colombia. Polygala caracasana Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 407. 1821 [1823]. Polygala columbica Chodat, Mém. Soc. Phys. Genève 31(2): 29. 1893. Polygala retifera S.F. Blake, Contr. U.S. Natl. Herb. 20: 527. 1924. Subshrub or perennial herb to 0.5 m tall; leaves membranous; flowers blue. Edges of evergreen lowland and submontane forests, 300–500 m; Bolívar (east of Miamo-Altiplanicie de Nuria). Aragua, Barinas, Carabobo, Distrito Federal, Falcón, Lara, Mérida, Miranda, Monagas, Nueva Esparta, Portuguesa, Sucre, Trujillo; Colombia, Trinidad, Ecuador. ◆Fig. 267. Eriksen et al. (Bente Eriksen, Bertil Ståhl, and Claes Persson. Polygalaceae. In: Flora of Ecuador No. 65. 2000) placed Polygala caracasana and members of Polygala sect. Hebecarpa Chodat in the genus Badiera, based on molecular evidence. We

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are awaiting the results of further studies before recognizing this genus. Polygala exigua A.W. Benn. in Mart., Fl. Bras. 13(3): 17. 1874. Panama, Colombia, Venezuela, Guyana, Brazil; 2 varieties, 1 in Venezuela. P. exigua var. fendleri (Chodat) Marques, Arch. Jard. Bot. Río de Janeiro 29: 69. 1988. —Polygala fendleri Chodat, Mém. Soc. Phys. Genève 31: 167. 1893. Herb; flowers white. Savannas, 300–1600 m; Bolívar (Ciudad Piar, El Pao, Gran Sabana, Roraima-tepui). Anzoátegui, Distrito Federal, Guárico, Portuguesa, Yaracuy; Panama, Colombia, Guyana, Brazil. ◆ Fig. 269. Polygala galioides Poir. in Lam., Encycl. 5: 503. 1804. Herb; flowers white. Savannas, 100–1000 m; Bolívar (35 km southwest of Caicara, La Paragua, Río Pao, Santa Elena de Uairén). Anzoátegui, Cojedes, Guárico, Portuguesa, Sucre; Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay. Polygala glochidiata Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 400. 1821 [1823]. U.S.A. (Arizona), Mexico, Central America, Cuba, Colombia, Venezuela, Guyana, Brazil, Bolivia, Paraguay, Argentina; 2 varieties, both in Venezuela, 1 of these in the flora area. P. glochidiata var. glochidiata. —Canilla de tintín, Pata de grillo. Herb; flowers white. Savannas, edges of Mauritia palm swamps, 50–1800 m; Bolívar (widespread). Anzoátegui, Aragua, Cojedes, Distrito Federal, Guárico, Monagas, Portuguesa; U.S.A. (Arizona), Mexico, Central America, Cuba, Colombia, Guyana, Brazil, Bolivia, Paraguay, Argentina. ◆Fig. 266. Polygala hygrophila Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 395. 1821 [1823]. —Waná-ká-puh (Pemón). Herb; flowers greenish white or purple. Savannas, edges of Mauritia palm swamps, 50–1600 m; Bolívar (Altiplanicie de Nuria,

Cerro Bolívar, widespread in Gran Sabana, Hato La Vergareña, Maripa, Urimán), Amazonas (La Esmeralda, Sierra Parima). Apure, Monagas, Táchira; Mexico, Central America, Hispaniola, Colombia, Guyana, Suriname, Brazil, Bolivia, Paraguay, Argentina. ◆Fig. 278. Polygala longicaulis Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 396. 1821 [1823]. —Ku’pe kwamen (Panare), Mentol, Siempre viva sabanera, Yaríkuto-orainó (Pemón). Herb; flowers white or purple. Savannas, 50–1600 m; Bolívar (widespread), Amazonas (Cerro Guanay, Puerto Ayacucho, Raudales de Atures, upper Río Ventuari, Samariapo, Santa Bárbara del Orinoco). Anzoátegui, Apure, Aragua, Carabobo, Cojedes, Distrito Federal, Guárico, Lara, Mérida, Monagas, Portuguesa, Sucre, Zulia; Mexico, Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Brazil, Bolivia, Paraguay. ◆Fig. 276. Polygala microspora S.F. Blake, Bull. Torrey Bot. Club 1: 397. 1929. Herb 2–5 cm tall; flowers white. Savannas, granitic outcrops, 50–1100 m; Bolívar (Río Kukenán, Roraima-tepui), Amazonas (Puerto Ayacucho-El Burro road, Samariapo). Guyana, Brazil, Bolivia. ◆Fig. 265. Polygala paniculata L., Sys. Nat. ed. 10, 2: 1154. 1759. —Parikuá-muranó (Pemón). Herb; flowers pink or white. Savannas, 100–2000 m; Bolívar (Cerro Akurima, Gran Sabana, Karaurín-tepui, La Escalera, Sororopán-tepui, Urimán). Widespread elsewhere in Venezuela; U.S.A. (Hawaii, Texas), Mexico, Central America, West Indies, Colombia, Guyana, Ecuador, Peru, Brazil, Bolivia, Argentina, Indonesia, Malaysia, Samoa, Fiji. ◆Fig. 273. Polygala sanariapoana Steyerm., Fieldiana, Bot. 28: 301. 1952. Herb; flowers white and purple. Savannas, granitic outcrops, 100–1600 m; Amazonas (Cerro Cuao, Puerto Ayacucho, Río Autana, Río Cabezón-Río Atabapo, Río Coromoto-Tobogán de la Selva, Río Sipapo, middle Río Ventuari, Samariapo). Colombia (Caquetá, Vaupés). ◆Fig. 272.

Polygala 331

Polygala savannarum Chodat, Bot. Jahrb. Syst. 52(Beibl. 115): 79. 1914. Herb; flowers white and purple. Savannas, 50–500 m; Bolívar (Hato La Vergareña, Río Maniapure, Serranía Carichana), Amazonas (Caño Picure, Galipero, Puerto Ayacucho, Raudales de Atures, Río Cataniapo, Río Coro Coro west of Cerro Yutajé, Samariapo, Santa Bárbara del Orinoco). Anzoátegui, Apure, Barinas, Guárico, Portuguesa; Colombia, Guyana, Brazil. ◆Fig. 277. Polygala sipapoana Wurdack, Mem. New York Bot. Gard. 8: 129. 1953. Herb; flowers white and purple. Highland sandstone outcrops, 1500–2000 m; Amazonas (Caño Cabeza de Manteco on Cerro Cuao, Cerro Sipapo). Endemic. Polygala spectabilis DC., Prodr. 1: 331. 1824. Subshrub or perennial herb to 1 m tall; leaves membranous; flowers pink. Savannas, 100–1100 m; Bolívar (Río Kukenán), Amazonas (Victorino-Río Guianía). Colombia, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia. ◆Fig. 280. Polygala spruceana A.W. Benn., J. Bot. 17: 203. 1879. Small herb; flowers white. Savannas, 50– 1400 m; Bolívar (Canaima, Carichana, lower Río Caroní, Uaipán-tepui, Urimán), Amazonas (Cerro Camani, Cerro Morrocoy, La Esmeralda, Puerto Ayacucho, Río Coro Coro west of Cerro Yutajé, San Juan de Manapiare, Santa Bárbara del Orinoco). Guárico; Guyana, Brazil (Pará). ◆Fig. 281. Polygala subtilis Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 393. 1821 [1823]. Herb; flowers white or greenish white. Savannas, edges of Mauritia palm swamps, 50– 1100 m; Bolívar (base of Auyán-tepui, Caicara, Canaima, Hato La Vergareña, Maripa, lower Río Caroní, Río Yuruaní, Santa Elena de Uairén, Uonquén, Urimán), Amazonas (Cerro Camani, Cerro Morrocoy, Culebra, La Esmeralda, Puerto Ayacucho, Río Parucito, Sabana del Oso-Río Ventuari, San Juan de Manapiare, Santa Bárbara del Orinoco, Samariapo). Guárico, Portuguesa; Colombia, Guyana, Suriname, Brazil. ◆Fig. 282.

Polygala tenella Willd., Sp. Pl. 3(2): 878. 1802. —Pari-co-guác-ariskú (Pemón). Polygala gracilis Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 401. 1821 [1823]. Polygala paludosa A. St.-Hil. & Moq., Mém. Mus. Natl. Hist. Nat. 17: 343. 1828. Polygala leptocaulis Torr. & A. Gray, N. Amer. Fl. 1: 30. 1838. Polygala alopecurus Chodat, Mém. Soc. Phys. Genève, 31: 227. 1893. Polygala funckii Chodat, Mém. Soc. Phys. Genève 31: 224. 1893. Herb; flowers white and purple. Savannas, edges of Mauritia palm swamps, 100– 400 m; Bolívar (Altiplanicie de Nuria, Ciudad Piar, El Manteco, Encrucijada de El Pao, Maniapure, lower Río Caroní), Amazonas (Caño Santo on Serranía de Guanay, La Esmeralda, Raudal de Maipures, Río Manapiare). Apure, Aragua, Cojedes, Distrito Federal, Guárico, Mérida, Portuguesa, Trujillo; U.S.A. (introduced in Florida, Louisiana), Mexico, Central America, Cuba, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina, Uruguay. ◆Fig. 279. Bernardi (L. Bernardi. Consideraciones taxonómicas y fitogeográficas acerca de 101 Polygalae Americanas. Cavanillesia Altera 1: 1–455. 2000) studied the types, original descriptions, and illustrations, and concluded that Polygala gracilis, P. leptocaulis, P. paludosa, and P. tenella lack significant morphological features to be treated as distinct species. Polygala timoutoides Chodat, Mém. Soc. Phys. Genève 30: 112. 1889. Costa Rica, Venezuela, Guyana, Brazil, Bolivia, Paraguay; 2 varieties, both in Venezuela, 1 of these in the flora area. P. timoutoides var. maguirei (Wurdack) Marques, Arch. Jard. Bot. Río de Janeiro 29: 39. 1988. —Polygala maguirei Wurdack, Mem. New York Bot. Gard. 23: 120. 1972. Herb; flowers and bracts greenish white. Savannas, edges of Mauritia palm swamps, 50–1200 m; Bolívar (Gran Sabana, Las Bonitas, Maripa), Amazonas (Río Asisa). Costa Rica, Guyana, Brazil, Bolivia, Paraguay. ◆Fig. 275.

332

P OLYGALACEAE

Polygala timoutou Aubl., Hist. Pl. Guiane 737, t. 295. 1775. Herb; bracts greenish white; flowers red or pink. Savannas, 100–1400 m; Bolívar (Auyán-tepui, Canaima, Chiguao, Gran Sabana, Hato La Vergareña, lower Río Caroní, Río Tirica 45 km east of Los Pijiguos, Urimán), Amazonas (Cerro Camani, Cerro Morrocoy-Río Manapiare, Culebra-Río Cunucunuma, La Esmeralda). Apure, Guárico, Portuguesa, Táchira; Costa Rica, Panama, Colombia, Trinidad, Guyana, Suriname, French Guiana, Brazil, Bolivia, Paraguay. Polygala trichosperma L., Syst. Nat. ed. 12, 3: 232. 1768. Polygala variabilis Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 397. 1821 [1823]. Herb; flower white to pale pink. Savannas, 50–1400 m; widespread in Bolívar and Amazonas. Anzoátegui, Apure, Guárico, Trujillo, Zulia; Mexico, Central America, Colombia, Guyana, Suriname, French Guiana, Brazil.

—Polygala monticola var. bryzoides (A. St.-Hil.) Steyerm., Fieldiana, Bot. 28: 300. 1952. Herb; flowers purple and yellow. Savannas, edges of forests, savannas, 50–1300 mm; Bolívar (widespread), Amazonas (Cerro Guanay, Puerto Ayacucho, Río Coromoto, Río Sipapo, Sabana Budare in upper Río Ventuari, Santa Bárbara del Orinoco). Anzoátegui, Apure, Carabobo, Cojedes, Distrito Federal, Guárico, Mérida, Miranda, Monagas, Portuguesa, Táchira, Trujillo, Yaracuy, Zulia; U.S.A. (Florida, Louisiana, Missisipi), Mexico, Central America, Cuba, Colombia, Guyana, Suriname, French Guiana, Trinidad, Ecuador, Brazil, Bolivia, Paraguay, Argentina. ◆Fig. 271. Polygala violacea is a common species and is highly variable in the flora area, with much variation in leaf shape, pubescence, and the relative abundance of glandular-ciliation on the outer sepals.

Polygala violacea Aubl., Hist. Pl. Guiane 735, t. 294. 1775; emend. Marques, Rodriguésia 31: 175. 1979. —JuiquiJuiqui, Pata de grillo, Rabo de alacrán, Raíz de cachicamo. Polygala angustifolia Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 405, t. 511. 1821 [1823]. Polygala mollis Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 406. 1821 [1823]. Polygala monticola Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 405. 1821 [1823]. Polygala brizoides A. St.-Hil. in A. St.-Hil. et al., Fl. Bras. Merid. 2: 44, t. 88. 1829.

Fig. 266. Polygala glochidiata var. glochidiata Fig. 265. Polygala microspora

Polygala 333

Fig. 267. Polygala caracasana

Fig. 269. Polygala exigua var. fendleri

Fig. 268. Polygala asperuloides

Fig. 270. Polygala appressa

334

P OLYGALACEAE

Fig. 271. Polygala violacea

Fig. 274. Polygala blakeana

Fig. 272. Polygala sanariapoana

Fig. 273. Polygala paniculata

Fig. 275. Polygala timoutoides var. maguirei

Polygala 335

Fig. 276. Polygala longicaulis

Fig. 277. Polygala savannarum

Fig. 278. Polygala hygrophila

Fig. 279. Polygala tenella

336

P OLYGALACEAE

Fig. 280. Polygala spectabilis

Fig. 281. Polygala spruceana

Fig. 282. Polygala subtilis

Securidaca 337

7. SECURIDACA L., Syst. Nat. ed. 10, 1155. 1759, nom. cons. non L. 1753. Elsota Adans., Fam. Pl. 2: 358. 1763. Lianas, scandent shrubs, rarely small trees. Leaves alternate, entire, shortpetiolate, generally with nodal stipular glands. Inflorescences few- to many-flowered, racemose or paniculate, terminal or axillary. Flowers papilionaceous; sepals 5, ± free from base, deciduous or persistent, arranged in 2 distinct whorls, the outer whorl 3 and herbaceous, the inner 2 (wings) much larger and petal-like, caducous in fruit; petals 3, white, pink, or purple, the lowest (keel) boat-shaped, clawed, with a plicate, fimbriate crest, the upper 2 adnate to the base of the staminal tube but distinct from the keel. Stamens 8; filaments connate nearly to the apex into a sheath split on the upper side, adnate to the keel, and upper petals toward the base; anthers basifixed, confluently 1-locular, opening by a large introrse-apical pore. Ovary (1)2, the ovule solitary, pendulous; disk a low fleshy ring at base of the ovary; style curved; stigma 2-lobed. Fruit a samara, the wing large, entire, sometimes bifurcate, 1-seeded. Seed glabrous, exarillate; endosperm absent; cotyledons thick-fleshy, oily. Pantropics (except Australia), mostly Neotropics; ca. 80 species, ca. 28 in Venezuela, 22 of these in the flora area. Securidaca is much in need of a taxonomic revision. Key to the Species of Securidaca 1. 1. 2(1). 2.

3(2).

3.

4(3).

4.

5(1.) 5. 6(5). 6. 7(6). 7.

Flowers solitary or in short axillary racemes 0.5–1.5 cm long with fewer than 8 flowers ........................................................................................ 2 Inflorescences multiflowered racemes 2.5–45 cm long, axillary or terminal ........................................................................................................... 5 Branchlets glabrous; inflorescence with 1–3 flowers; pedicels 6–12 mm long; flower keel lacking crest; samara wing glabrous ............. S. uniflora Branchlets densely hirsutulous or sparsely strigulose to glabrescent when mature; inflorescences with 4–8 flowers; pedicels 2–5 mm long; flower keel crested ................................................................................. 3 Branchlets, petioles, and pedicels with white pubescence; leaves ovateelliptic to elliptic; margins revolute; ovary sparsely strigulose to glabrescent .......................................................................... S. savannarum Branchlets, petioles, and pedicels without white pubescence; leaves ovate-oblong to oblong-ovate; margins not revolute; ovary glabrous ................................................................................................................ 4 Branchlets densely hirsutulous with trichomes ca. 1 mm long; petiolar glands sessile; leaves chartaceous, base subcordate; lateral petals sparsely ciliate outside ................................................................ S. prancei Branchlets sparsely strigulose to glabrescent when mature with trichomes 0.05 mm long; petiolar glands elevated; leaves coriaceous, base obtuse or rotundate; lateral petals glabrous outside .............. S. fruticans Leaves glabrous on both surfaces .............................................................. 6 Leaves strigulose, pilose, or pubescent at least on one surface ............... 8 Leaf margins crenate and thickened ........................................ S. marginata Leaf margins entire, not thickened ........................................................... 7 Leaves ovate to widely ovate; racemes 3–5 cm long; pedicels 8–10 mm long; wings of the samara glabrous, 4–5 cm long .................... S. scandens Leaves elliptic to ovate; racemes 4–12 cm long; pedicels 5–6 mm long;

338

P OLYGALACEAE

8(5). 8. 9(8). 9. 10(9). 10. 11(10). 11.

12(8). 12. 13(12). 13. 14(13).

14. 15(14). 15.

16(13). 16.

17(16).

17.

18(12). 18. 19(18).

wings of the samara puberulent, ca. 1.5 cm long ......................... S. retusa Samaras 2-winged ...................................................................................... 9 Samaras 1-winged .................................................................................... 12 Racemes 10–30 cm long; wings ca. 1 cm long ................................S. pendula Racemes 4–10 cm long; wings 1–3 cm long ............................................ 10 Leaves oblong, sometimes elliptic or obovate; inflorescence racemes branched (panicles); flowers 5–7 cm long; ovary glabrous ..... S. paniculata Leaves ovate, lanceolate, or lanceolate-ovate; inflorescence racemes unbranched; flowers 8–16 cm long; ovary pubescent ............................. 11 Leaves coriaceous; inner sepals symmetrical, glabrescent outside; keel crest 1.2–1.5 mm long; wings glabrescent .................................. S. bialata Leaves chartaceous to subcoriaceous; inner sepals asymmetric, pilose outside; keel crest 0.5–0.7 mm long; wings densely strigulose .................................................................................................. S. longifolia Lower surface of leaves puberulent, sparsely hirsutulous, strigulose, or pilosulous, more evident along the midrib and secondary veins ....... 13 Lower surface of leaves hirsute or densely appressed-pubescent ......... 18 Inflorescences simple racemes; flower keel lacking crest or crest reduced and < 4 mm long, not reflexed ............................................................. 14 Inflorescences branched panicles; flower keel crested; crest 8–15 mm long, reflexed ........................................................................................ 16 Leaves subcoriaceous, margins not revolute, pilosulous on the upper surface; outer sepals ovate, ca. 1.5 mm wide; keel petal not ciliate ................................................................................................. S. divaricata Leaves coriaceous, margins revolute, glabrous on the upper surface; outer sepals ovate-orbicular, 2–3.5 mm wide; keel petal ciliate ........ 15 Branchlets, petioles, rachis of inflorescence, and pedicels hirsute; petioles 5–7 mm long; dorsal lobe of samara well developed .............. S. cacumina Branchlets, petioles, rachis of inflorescence, and pedicels velutinouspubescent; petioles 1–2 mm long; dorsal lobe of samara obsolete ............................................................................................ S. warmingiana Leaves all one size; outer sepals glabrous outside; samara wing 1–1.5 cm long ......................................................................................... S. paniculata Leaves two different sizes, the ones on inflorescence rachis smaller than ones on main branches; outer sepals strigulose outside; samara wing 2–4 cm long .......................................................................................... 17 Leaves elliptic-ovate, obtuse or subcordate at the base, sparsely hirsutulous to glabrescent on the upper surface; inflorescences 8–45 cm long; petal keel 10–11 mm long .......................................................... S. speciosa Leaves ovate-oblong, sometimes ovate, short-attenuate at the base, strigulose on the lower surface; inflorescences 4–12 cm long; petal keel ca. 8 mm long ........................................................................ S. diversifolia Young branchlets, petioles, inflorescences, and pedicels densely whitepubescent; petioles 10–15 mm long; ovary puberulent ........... S. maguirei Young branchlets, petioles, inflorescences, and pedicels not white-pubescent; petioles 1–7 mm long; ovary glabrous or densely hirsute ......... 19 Leaves two different sizes, the ones on the inflorescence rachis smaller than ones on main branches, with long trichomes (0.7–1.5 mm long) on

Securidaca 339

the lower surface ........................................................................ S. coriacea 19. Leaves all one size, trichomes ca. 0.5 mm long on the lower surface .... 20 20(19). Branches and branchlets furfuraceous-pilose; leaves coriaceous; flowers 3–8 mm long; inner sepals glabrous ................................... S. pyramidalis 20. Branches and branchlets tomentose, densely pilose or densely pubescent; chartaceous to subcoriaceous; flowers 9–13 mm long; inner sepals strigillose or pubescent ........................................................................ 21 21(20). Leaves densely puberulent on the lower surface; petioles 5–9 mm long; ovary glabrous; samara wing ca. 4 cm long ............................ S. pubescens 21. Leaves densely pubescent on the lower surface; petioles 2–4 mm long; ovary pubescent; samara wing 4–6 cm long ....................................... 22 22(21). Leaves elliptic, sometimes ovate, densely pubescent on the upper surface; pedicels 4–5 mm long; outer sepals puberulous; upper petals spatulate-ovate, obtuse at apex .......................................................... S. volubilis 22. Leaves ovate, oblong-ovate, or ovate, sparsely pilose to glabrous on the upper surface; pedicels 5–8 mm long; outer sepals densely pilose; upper petals obovate or obovate-spatulate, rounded at apex .... S. tenuifolia Securidaca bialata Benth., Hooker’s J. Bot. Kew Gard. Misc. 3: 162. 1851. Liana; flowers pink; samaras 2-winged. Evergreen lowland forests, 50–400 m; Delta Amacuro (Caño Corisal, Sacupana), Bolívar (Hato La Vergareña, Río Aro, Río Asa, Río Karún, upper Río Paragua, Río Torono, Temblador), Amazonas (Puerto Ayacucho). Falcón, Guárico; Brazil. ◆Fig. 283. Securidaca cacumina Wurdack, Mem. New York Bot. Gard. 23: 126. 1972. Securidaca ecristata Wurdack, Mem. New York Bot. Gard. 9: 351. 1957, non Kassau 1934. Liana; flowers pink. Montane forests, 1400–2100 m; Bolívar (Cerro Jaua, Cerro Sarisariñama), Amazonas (Cerro Huachamacari, Cerro Marahuaka). Endemic. ◆Fig. 284. Securidaca coriacea Bonpl., Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck. Gesammten Naturk. 2: 47. 1808. —Bejuco cuadrado, Bejuco jabón. Securidaca mollis Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 421. 1821 [1823]. Liana; flowers pink. Scrub savannas, evergreen lowland forests, 50–300 m; Delta Amacuro (east-southeast of Los Castillos de Guayana), widespread in Bolívar and Amazonas. Anzoátegui, Apure, Aragua, Barinas, Guárico, Lara, Mérida, Miranda, Monagas, Portuguesa, Táchira, Trujillo, Zulia; Pa-

nama, Colombia, Trinidad, Guyana, Suriname, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 286. Securidaca divaricata Nees & Mart., Nova Acta Phys.-Med. Acad. Caes. Leop.Carol. Nat. Cur. 12(1): 25. 1824. Securidaca rivinifolia A. St.-Hil. & Moq., Mém. Mus. Hist. Nat. 17: 328. 1828. —Securidaca rivinifolia var. parvifolia A.W. Benn. in Mart., Fl. Bras. 13(3):66. 1874, “rivinaefolia.” Liana; flowers purple-pink. Evergreen lowland forests, 100–200 m; Amazonas (Río Guainía-Banderita). Anzoátegui, Aragua, Miranda; Colombia, Ecuador, Guyana, Suriname, French Guiana, Peru, Amazonian Brazil, Bolivia. In herbaria, specimens of Securidaca divaricata often have been annotated as S. rivinifolia var. parvifolia. Securidaca diversifolia (L.) S.F. Blake, Contr. U.S. Natl. Herb. 23: 594. 1923. —Polygala diversifolia L., Sp. Pl. 703. 1753. —Adájate (Bale), Bejuco cuadrado, Bejuco negro, Guambe (Bale), Ojo de zamuro, Pentro, Requena, Ukánaha (Bale). Securidaca fendleri Chodat, Bull. Herb. Boissier 3: 545. 1895. Securidaca venosa Rusby, Descr. S. Amer. Pl. 41. 1920.

340

P OLYGALACEAE

Liana; flowers pink and yellow. Savannas, semideciduous and evergreen lowland to montane forests, 50–500 m; Delta Amacuro (El Palmar-Río Grande), Bolívar (Cerro Pijiguao, Ciudad Bolívar, Ciudad Piar, Corozal, El Manteco, Jabillal, La Paragua, Maripa, Represa Guri, Río Caura, Río Karún, Río Suapure), Amazonas (Caño Asisa, La Esmeralda, Río Cuao, basin of Río Manapiare, upper Río Orinoco, Río Yatúa, Santa Bárbara, Trapichote, Yavita). Anzoátegui, Apure, Aragua, Barinas, Distrito Federal, Falcón, Guárico, Mérida, Miranda, Portuguesa, Sucre, Trujillo, Yaracuy, Zulia; Mexico, Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil. Securidaca fruticans Wurdack, Mem. New York Bot. Gard. 23: 124. 1972. Liana; flowers pink. Evergreen lowland forests, 100–200 m; Amazonas (Caño Hechimoni on Río Siapa, Río Pasimoni, Río Yatúa). Endemic. Securidaca longifolia Poepp. & Endl., Nov. Gen. Sp. Pl. 3: 66. 1845. —Bejuco jabón. Liana; flowers pink. Evergreen lowland forests, riparian black-water forests, 100– 200 m; Amazonas (Caño Atamoni, Caño San Miguel, Caño Yagua, Laguna Paciba, Maroa, Río Atacavi, Río Baría, Río Emoni, Río Pasimoni, Río Puruname, Río Temi, Río Yatúa, San Carlos de Río Negro, Solano). Colombia, Peru, Brazil, Bolivia. ◆Fig. 287. Securidaca maguirei Wurdack, Mem. New York Bot. Gard. 23: 122. 1972. Liana; flowers pink. Evergreen lowland and montane forests, 100–1000 m; Amazonas (Cerro Sipapo, 35 km southeast of La Esmeralda, 25–30 km southeast of Puerto Ayacucho, upper Río Cunucunuma, Río Matacuni, San Fernando de Atabapo, Trapichote). Brazil (Roraima). Securidaca marginata Benth., J. Bot. (Hooker) 4: 103. 1842. Liana or scandent shrub; flowers pink. Scrub savannas, gallery forests, evergreen lowland to montane forests, 300–1300 m; Bolívar (Amaruay-tepui, Auyán-tepui [Salto Angel], Canaima, Cerro Bolívar, Cerro Guaiquinima, Cusimi, Gran Sabana, Río Asa, Río

Caroní, Río Hacha, Río Paragua, Río Urimán). Guyana, Brazil (Roraima). ◆Fig. 285. Securidaca paniculata Rich., Actes Soc. Hist. Nat. Paris 1: 111. 1792. Guyana, Suriname, French Guiana, Peru, Brazil; 2 varieties, both in the flora area. Key to the Varieties of S. paniculata 1. Midvein on upper surface of leaf ciliate; lower surface of leaf pale yellowish; samaras 3–4 cm long ..... var. lasiocarpa 1. Midvein on upper surface of leaf not ciliate; lower surface of leaf greenish; samaras 1–2 cm long .......... var. paniculata S. paniculata var. lasiocarpa Oort, Meded. Bot. Mus. Herb. Rijks Univ. Utrecht 71: 683. 1939. —Pentro. Securidaca hostmannii Miq., Tijdschr. Wis-Natuurk. Wetensch. Eerste Kl. Kon. Ned. Inst. Wetensch. 1: 153. 1848, “hostmanni.” Liana; flowers pink. Evergreen lowland and montane forests, 50–1000 m; Delta Amacuro (Caño Araguabisi, Caño Araguao, El Palmar, Río Acure, Río Grande), Bolívar (Río Caura, Río Venamo, Salto El Loro), Amazonas (San Carlos de Río Negro, slopes of Sierra de la Neblina). Mérida; Guyana, Suriname, French Guiana, Peru, Brazil. S. paniculata var. paniculata Liana; flowers pink. Evergreen lowland and montane forests, 50–800 m; Delta Amacuro (El Palmar, east-southeast of Los Castillos, Río Amacuro), Bolívar (Altiplanicie de Nuria, lower Río Caura, Río Karún, Río Tabaro, Río Venamo, Santa Elena de Uairén, Tumeremo). Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil. ◆Fig. 288. Securidaca pendula Bonpl., Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck. Gesammten Naturk. 2: 47. 1808. —Bejuco cuarenta días, Bejuco culebrón. Securidaca complicata Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 423. 1821 [1823]. Securidaca orinocensis Rusby, Descr. S. Amer. Pl. 40. 1920.

Securidaca 341

Liana; flowers pink; samaras 2-winged. Evergreen lowland forests, granitic outcrops, 50–200 m; Bolívar (El Dorado, El Temblador, Piedra Marimare on middle Río Orinoco, Río Cuyuní, Río Paragua, Río Tonoro, San Félix, Túriba), Amazonas (Caño Guacamayo, Caño Yagua, Río Atabapo, Río Cataniapo, Río Guayapo, Río Sipapo). Anzoátegui, Apure, Guárico; Colombia, Brazil. ◆Fig. 292. Securidaca prancei Wurdack, Mem. New York Bot. Gard. 23: 124. 1972. Liana; flowers pink. Scrubby savannas, evergreen lowland forests, 100–400 m; Bolívar (Río Canaracuni, upper Río Caura), Amazonas (Caño Chimoni in Río Casiquiare basin, Río Baría). Brazil (Amazonas, Pará), Bolivia. Securidaca pubescens DC., Prodr. 1: 341. 1824. Securidaca major Sagot ex A.W. Benn. in Mart., Fl. Bras. 13(3): 64. 1874. Securidaca densiflora Oort, Meded. Bot. Mus. Herb. Rijks Univ. Utrecht 5: 13. 1933. Liana; flowers pink. Evergreen lowland forests, 100–300 m; Bolívar (Cerro El Sol), Amazonas (Culebra, San Carlos de Río Negro). Cojedes, Guárico, Miranda; Colombia, Guyana, French Guiana, Peru, Brazil. ◆Fig. 293. Securidaca pyramidalis Sprague, Bull. Misc. Inform. Kew 1931: 174. 1931. —Bejuco jabón. Liana; flowers purple. Scrub savannas, evergreen montane forests, gallery forests and thickets, 100–1400 m; Bolívar (lower slopes of Ilú-tepui, Kavanayén, 20–30 km north of Las Bonitas, Luepa, Río Aponguao, Santa Elena de Uairén), Amazonas (upper Río Orinoco, Yavita). Guyana. Securidaca retusa Benth., Hooker’s J. Bot. Kew Misc. 2: 210. 1850. —Bejuco de jabón. Liana; flowers pink. Semideciduous forests, savannas, low Amazon caatinga, evergreen lowland and montane forests, 50–800 m; Bolívar (Puerto Ordaz), Amazonas (La Esmeralda, Maroa, San Carlos de Río NegroSolano area). Colombia, Guyana, French Guiana, Brazil. ◆Fig. 291.

Securidaca savannarum Wurdack, Mem. New York Bot. Gard. 9: 352. 1957. Liana; flowers pink. Scrub savannas on sandy soils, Río Negro caatinga, 100–200 m; Amazonas (Caño Cotúa at base of Cerro Yapacana). Endemic. Securidaca scandens Jacq., Enum. Syst. Pl. 27. 1760. —Bejuco cuarenta días. Securidaca cordata J.R. Johnst., Proc. Amer. Acad. Arts 40: 688. 1905. Liana; flowers pink. Shrubby savannas, deciduous to evergreen forests, 50–300 m; Bolívar (Cerro La Cruz, Ciudad Bolívar, El Manteco, Represa Guri, Río Canaracuni, San Felíx to Puerto Ordaz). Anzoátegui, Aragua, Carabobo, Distrito Federal, Guárico, Falcón, Lara, Mérida, Miranda, Monagas, Nueva Esparta, Sucre, Táchira, Zulia; Colombia. ◆Fig. 295. Securidaca speciosa Wurdack, Mem. New York Bot. Gard. 9: 353. 1957. Liana; flowers black-purple. Evergreen lowland to montane forests, 100–1300 m; Bolívar (Gran Sabana, La Paragua, Raudal Auraima on Río Paragua), Amazonas (Río Matacuni). Endemic. Securidaca tenuifolia Chodat, Bull. Herb. Boissier 36: 545. 1895. —Roblito. Liana; flowers pink. Semideciduous to evergreen lowland forests, 200–500 m; Bolívar (Kavanayén, La Camilera 40 km west of El Manteco, Represa Guri, Santa Elena de Uairén). Aragua, Barinas, Carabobo, Distrito Federal, Mérida, Miranda, Portuguesa, Táchira; Panama, Colombia, Trinidad. Securidaca uniflora Oort, Meded. Bot. Mus. Herb. Rijks Univ. Utrecht 5: 15. 1933. Liana; flowers pink, maroon, or white. Evergreen lowland forests, riparian scrub, 100– 200 m; Amazonas (Maroa, Río Atacavi). Colombia, Guyana, Suriname, French Guiana, Brazil (Amazonas). ◆Fig. 289. Securidaca volubilis L., Sp. Pl. 707. 1753; emend. Oort, Meded. Bot. Mus. Herb. Rijks Univ. Utrecht 71: 679. 1939. —Cashék (Yanomami). Liana; flowers pink. Semideciduous to evergreen lowland forests, 100–900 m; Bolívar

342

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Fig. 283. Securidaca bialata

Fig. 284. Securidaca cacumina

Fig. 285. Securidaca marginata

Securidaca 343

Fig. 286. Securidaca coriacea

Fig. 287. Securidaca longifolia

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Fig. 288. Securidaca paniculata var. paniculata

Fig. 289. Securidaca uniflora

Fig. 290. Securidaca warmingiana

Securidaca 345

Fig. 291. Securidaca retusa

Fig. 292. Securidaca pendula

Fig. 293. Securidaca pubescens

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P OLYGALACEAE

Fig. 294. Securidaca volubilis

(Represa Guri, Río Caura, Salto Pará), Amazonas (Río Ocamo, base of Sierra de la Neblina, Simarawochi). Mérida, Zulia; Panama, Colombia, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 294. The genus Securidaca was first described by Linnaeus in Genera Plantarum, ed. 5: 316, 1754. It was based on the type species S. volubilis from Species Plantarum (Linnaeus 1753). The type specimen consisted of three sheets, two of which contained Polygalaceae flowers and fruits, and the other consisting of fruits of Leguminosae [A.J.P. Oort. Critical remarks on the Suriname species of the genus Securidaca (Polygalaceae). Meded. Bot. Mus. Herb. Rijks Univ. Utrecht 71: 677–685. 1939]. In 1759, Linnaeus made corrections to Securidaca (see: Systema Naturae, ed. 10: 1155). Oort (1939) examined the type speci-

Fig. 295. Securidaca scandens

P O LY G O N A C E A E 347

men, made a complete description, and emended Linnaeus’s work. He concluded that the name Securidaca belongs to the Polygalaceae plants on the type specimen. Adams (Regnum Veg. 127: 87. 1993) generityped Securidaca as Linnaeus 1759, Polygalaceae, nom. cons. against Securidaca L. 1753, Fabaceae. He cited Securidaca volubilis L. (1759) as nom. rej., interpreting it as a later homonym of S. volubilis L. (1753) and designating S. brownei Griseb. as the correct name for it. Eriksen et al. (Bente Eriksen, Bertil Ståhl, and Claes Persson. Polygalaceae. In: Flora of Ecuador No. 65. 2000) and Marques [Mendes Marques. Securidaca L. (Polygalaceae) do Brasil, Arch. Jard. Bot. Rio Janeiro 34: 7–144. 1996] considered S. volubilis to be auct. non and placed this name in the synonymy of S. rivinifolia A. St.-Hil. and S. diversifolia (L.) S.F. Blake respectively. However, we use the name S. volubilis here, in anticipation of a generic revision, because S. brownei is a Ja-

maican endemic; in addition, the specimens of the flora area match neither S. rivinifolia A. St.-Hil. nor S. diversifolia (L.) S.F. Blake. Rather, our specimens match the descriptions of Oort (1939) and A. Jacobs-Brouwer [Polygalaceae. Pp. 585–589, in: S. A. Mori et al. (eds.). Guide to the Vascular Plants of Central French Guiana: part 2 (Dicotyledons). Mem. New York Bot. Gard. 76(2): 2002]. Securidaca warmingiana Chodat, Bull. Herb. Boissier 3: 544. 1895. —Bejuco jabón. Liana; flowers pink. Evergreen lowland forests, 50–600 m; Amazonas (Caño Colorado 55 km southeast of Puerto Ayacucho, Caño Yapacana, Capibara, Río Autana, Río Cuao, upper Río Orinoco at Raudal de Los Guaharibos, Río Siapa, Río Sipapo, Río Yatúa, San Carlos de Río Negro, San Fernando de Atabapo). Mérida; Colombia, Peru, Amazonian Brazil, Bolivia. ◆Fig. 290.

POLYGONACEAE by Gerardo A. Aymard C. and Richard A. Howard Annual or perennial herbs, twining herbaceous or woody vines, shrubs, or trees, occasionally with short shoots, these flowering, leafy, or ending in spines. Stems often striate or sulcate, internodes hollow or solid, sulcate. Leaves mostly alternate, seldom opposite or whorled, simple, pinnately veined; margins usually entire; petiolate; stipules intrapetiolar, united into a tubular, persistent or caducous, usually membranous to hyaline, often bilobed or fringed sheath (ochrea). Inflorescences terminal or axillary, occasionally ending in a tendril, racemose, paniculate, or spikelike in appearance, but basically composed of partial inflorescences, these congested dichasia or helicoid cymes; the partial inflorescences subtended by bracts, each flower subtended by a persistent, membranous, tubular ochreola consisting of the fused bracteoles. Flowers articulated to the pedicel, bisexual or unisexual (the plants then dioecious). Perianth uniseriate or biseriate, of 3–6 free or partially united tepals, occasionally with 3 differentiated petals, greenish to red or white; hypanthium variously developed, either the tube or the lobes or both enlarging in fruit development, or the lobes keeled, the keel wing-like and at times extending onto the pedicel, or the lobes enlarging into 2 or 3 wings. Stamens usually 6–9 (numerous in Symmeria); filaments flattened, usually glabrous, fused near the base and/or partially adnate to the hypanthium, rudimentary or as staminodes in pistillate flowers; anthers 2- or 4-locular, usually versatile, introrse. Ovary superior, triquetrous or lenticular, unilocular, with one erect ovule; styles 1–3, filiform,

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terminally capitate. Fruit an achene or small nut, round, trigonous, or sometimes lenticular, covered by the adherent hypanthium or enclosed by a papery or fleshy perianth or by the perianth lobes, or the perianth lobes extended as wings; pericarp usually shiny, thin or thick. Seeds with endosperm mealy, often ruminate; embryo centric or eccentric. Temperate, subtropical, and tropical regions, mostly in north-temperate areas; 45 genera and ca. 1000 species, 8 genera and 60 species in Venezuela, 6 genera and 45 species in the flora area. Key to the Genera of Polygonaceae 1. 1. 2(1). 2. 3(1). 3. 4(3).

4.

5(3).

5.

Herbs or vines; tepals 5 ............................................................................. 2 Shrubs, trees, or lianas; tepals 6 ............................................................... 3 Vines with tendrils; stigma peltate ............................................ 1. Antigonon Plants erect or prostrate, without tendrils; stigma capitate ....... 3. Polygonum Stipules entirely caducous; outer tepals expanding and extending in fruit to form a winged, helicopter-like fruit .................................................. 4 Stipules wholly or partially persistent; outer tepals not developing into wings in fruit .......................................................................................... 5 Leaves 3–12 cm long, with < 12 pairs of major secondary veins; achene with 3 deep longitudinal sulci separating 3 rounded sides, loosely enclosed within the fruiting perianth ....................................... 4. Ruprechtia Leaves 15–35 cm long, with > 12 pairs of major secondary veins; achene with 3 sharp ridges and 3 flat or concave faces, tightly enclosed within the fruiting perianth .................................................................. 6. Triplaris Stipules with ochreae; perianth segments and stamens 5(7); fruit falsely drupaceous, usually tightly enclosed by the enlarged fleshy hypanthium or the perianth lobes; achene black, brown, or tan .................................................................................................. 2. Coccoloba Stipules without ochreae; perianth segments 6; stamens 20–35; fruit strongly trigonous, tightly enclosed by lobes of the perianth; achene green and red .......................................................................... 5. Symmeria

1. ANTIGONON Endl., Gen. Pl. 310. 1837. Scandent vines with tendrils terminating the inflorescences. Stems sulcate, herbaceous or suffrutescent, pubescent or glabrate. Leaves alternate; blades membranous, broadly ovate to deltoid, the base cordate to deltoid, the apex acute to acuminate; ochreae obsolete; petioles terete or winged. Inflorescence of axillary and terminal racemes or panicle-like pleiothyrsi, the axes pubescent to glabrous, terminated by tendrils. Flowers bisexual, articulated to pedicels, in fascicles of 1–many. Perianth of 5 free unequal tepals, the outer 3 broader than the inner 2, red or greenish white, accrescent in fruit. Stamens 8, equal; filaments united basally into a tube essentially free from the tepals; anthers introrse, versatile. Ovary trigonous; styles 3, arcuate; stigmas peltate. Achenes mostly enclosed by the tepals, bluntly 3-angled, usually brown and lustrous. Mexico, Central America, introduced and widely cultivated in all tropical and subtropical areas for the attractive inflorescences with long-persisting rose red, pink, or white perianth parts; 8 species, 2 in Venezuela, 1 of these in the flora area.

Coccoloba 349

Fig. 296. Antigonon leptopus

Antigonon leptopus Hook. & Arn., Bot. Beechey Voy. 308. 1838. —Bellisima, Coronilla. Vine; flowers pink to purplish. Cultivated and occasionally escaped from cultivation, 100–200 m; Delta Amacuro (Sacupana),

Bolívar (Ciudad Bolívar, Upata), Amazonas (San Carlos de Río Negro). Widespread elsewhere in northern Venezuela; native to Mexico, widely cultivated in tropical and subtropical areas. ◆Fig. 296.

2. COCCOLOBA P. Browne ex L., Syst. Nat. ed. 10, 1007, 1367. 1759, nom. cons. Guaiabara Mill., Gard. Dict. ed. 4, 2. 1754. Coccolobis P. Browne, Civ. Nat. Hist. Jamaica 209, pl. 14, fig. 3. 1756. Campderia Benth., Bot. Voy. Sulphur suppl. 159. 1846, non Lag. 1821. Shrubs, trees with scrambling branches, or lianas. Stems stout or slender, striate or smooth, pith solid or stem hollow in internodal areas; nodes commonly slightly swollen but in some West Indian species enlarged and bulbous. Leaves alternate; ochreae sheathing at first, splitting along 1 or 2 sides or the basal portion coriaceous and persistent, the apical portion membranous and deciduous; petioles stout to slender, borne at the base or well above the base of the ochrea; blades coriaceous or membranaceous, pubescent to glabrous; margins flat to undulate; secondary venation conspicuous and reticulate or inconspicuous. Inflorescence terminal or terminal on lateral short shoots, paniculate, racemose, or spicate; peduncle usually short. Flowers functionally unisexual (plants dioecious). Staminate flowers borne in clusters of 2–7. Pistillate flowers solitary at each nodule on the axis, each cluster ochreolate and each pedicel with a smaller ochrea, these covering the flower in bud

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and splitting, persistent or caducous; pedicels short or well developed, equaling or exceeding the ochreolae; hypanthium well developed or very slight. Perianth lobes 5(–7), imbricate in bud, often reflexed in flower. Stamens usually 5(–7), functional stamens exserted, nonfunctional stamens included; filaments commonly flaring at the base, often forming a short tube, free from hypanthium. Functional pistil exserted, styles 1–3, the nonfunctional pistils rudimentary, included. In fruit, hypanthium and perianth lobes expanded, or the achene surrounded by the expanded hypanthium with the perianth lobes imbricate or coronate, or the hypanthium barely expanded and the achene surrounded by the imbricate expanded perianth lobes; achene trigonous in outline, black, brown, or tan, shiny or dull, outer wall hard, inner layer papery. Seed with ruminate endosperm, the involutions many, the major lobes 3; embryo centrally located; cotyledons orbicular, flat, rarely folded or contorted, the radicle small and terete. U.S.A. (southern Florida), Mexico, Central America, West Indies, south to Argentina; 400 species, ca. 41 species in Venezuela, 29 of these in the flora area. Coccoloba is in need of revision, and some of the taxa included here will likely be modified as a result. Key to the Species of Coccoloba 1. 1. 2(1). 2. 3(2).

Inflorescence paniculate or fasciculate ..................................................... 2 Inflorescence spicate or racemose ............................................................. 5 Plants entirely glabrous ............................................................................ 3 Plants pubescent ........................................................................................ 4 Stems not striate, internodes solid; leaves oblong-lanceolate, smooth; inflorescence short-stalked, appearing fascicled, < 1/2 the length of the leaves, red, pendent ....................................................................... C. fallax 3. Stems strongly striate, ridged, internodes hollow; leaves broadly ovate to suborbicular, strongly bullate; inflorescence with elongate peduncles, > 1/2 the length of the leaves, green, erect ................................. C. latifolia 4(2). Plants sericeous-pubescent; leaves primarily broadly ovate, the apex acute; perianth lobes coronate in fruit ......................................... C. mollis 4. Plants puberulent; leaves primarily oblong to elliptic, the apex rounded, emarginate, or rarely short acute; perianth lobes imbricate in fruit .............................................................................................. C. dugandiana 5(1). Plants scandent .......................................................................................... 6 5. Shrubs or trees, not at all scandent ........................................................ 14 6(5). Inflorescence slender, shorter than the leaves ......................................... 7 6. Inflorescence stout, longer than the leaves .............................................. 8 7(6). Leaves 30–33 × 20–25 cm, with black dots on the lower surface ..... C. sp. D 7. Leaves 10–25 × 7–20 cm, without black dots on the lower surface ................................................................................................. C. ochreolata 8(6). Flowers and fruits pedicellate, pedicels > 8 mm long ............... C. wurdackii 8. Flowers and fruits sessile or pedicels very short, 2–6 mm long .............. 9 9(8). Ochreolae membranaceous, much longer than pedicels, flaring ... C. excelsa 9. Ochreolae short-truncate, much shorter than pedicels .......................... 10 10(9). Leaves broadly oblong or elliptic, 12–20 cm wide; fruit 1.5–2 cm diameter ................................................................................................. C. ascendens

Coccoloba 351

10. 11(10). 11. 12(11). 12. 13(11). 13. 14(5). 14. 15(14). 15. 16(14). 16. 17(16). 17. 18(17).

18.

19(17).

19.

20(19). 20. 21(20).

21.

Leaves oblanceolate to lanceolate or oblong to elliptic-oblong, 5–10 cm wide; fruit 0.5–1 cm diameter ............................................................. 11 Leaves oblong to elliptic-oblong or elliptic-ovate, midrib depressed above, petioles 0.5-1 mm wide ........................................................................ 12 Leaves oblanceolate, lanceolate, or lanceolate-elliptic, midrib elevated above, petioles > 1.5 mm wide ............................................................ 13 Leaves obovate-elliptic, rigid-coriaceous, base cordate, secondary veins 10–14; stamens 1.5–2 mm long ...................................... C. gymnorrhachis Leaves obovate, chartaceous, base acute, secondary veins 7–9; stamens ca. 1 mm long ...................................................................................C. sp. C Leaves thinly coriaceous, the base obtuse; petioles striate, stout ................................................................................................ C. marginata Leaves chartaceous, lustrous on the upper surface; base rounded or subcordate, petioles not striate, slender ......................................... C. lucidula Ochreae of flowering branches large, 1.5–5 cm long, 1–2 cm wide, foliaceous and thinly membranaceous ....................................................... 15 Ochreae of flowering branches small, to 1 cm long, 0.3–0.8 cm wide, the basal portion coriaceous and persistent ............................................. 16 Plants growing in flooded forests; leaves subobovate, 15–27 cm long, the apex acute to acuminate; flowers 1 or 2 per fascicle ............. C. spruceana Plants growing in savannas or dry forests; leaves rounded or cordate, 8– 13 cm long, the apex rounded; flowers 3 or 4 per fascicle ...... C. caracasana Leaves rounded, obtuse, or rarely bluntly acute at the apex ................. 17 Leaves acute to acuminate at apex ......................................................... 22 Leaves pubescent on lower surface ......................................................... 18 Leaves glabrous ....................................................................................... 19 Branches and branchlets smooth; leaves ovate to broadly ovate, glabrous on the upper surface except at the midrib and the secondary veins, black dots on the lower surface .............................................. C. orinocana Branches and branchlets striate; leaves reniform, pubescent on the upper surface, more densely on the midrib and the secondary veins, without black dots on the lower surface ...................................................... C. sp. B Shrub found at elevations above 1000 m; leaves rigidly coriaceous; fruit ovoid, 6–7 × 5 mm; perianth lobes coronate, not enlarged or extended ........................................................................................... C. schomburgkii Small trees found at elevations below 1000 m; leaves membranous or thinly coriaceous, not rigid; fruit generally larger; perianth lobes enlarged and extended ............................................................................ 20 Trees planted as ornamentals, leaves orbicular to reniform; fruit ca. 20 mm long ................................................................................... C. uvifera Trees not planted, leaves ovate, elliptic, or obovate; fruit 4–10 mm long .............................................................................................................. 21 Inflorescence with flowering nodes widely separated, the rachis zigzag; perianth in fruit fleshy, 1–2 cm long, the hypanthium surrounding the achene, the perianth lobes enlarged and extended ....................... C. ovata Inflorescence with flowering nodes proximate on a straight axis; perianth lobes surrounding the achene, imbricate, the hypanthium short ................................................................................................ C. obtusifolia

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22(16). Leaves obovate, clearly narrowed below the middle .............................. 23 22. Leaves broadly ovate, ovate, elliptic, lanceolate, lanceolate-elliptic, or ovate-lanceolate ................................................................................... 24 23(22). Leaves membranaceous, with lepidote scales and gland dots on the lower surface; flowers sessile ............................................................ C. llewelynii 23. Leaves subcoriacerous or coriaceous, the lower surface without lepidote scales or gland dots; flowers pedicellate, pedicels 5–7 mm long ................................................................................................. C. densifrons 24(22). Leaves borne above the base of the ochrea, shiny on the upper surface, drying olive-green ...................................................................... C. lucidula 24. Leaves borne at the base of the ochrea, opaque on the upper surface, drying black, red, or brown ....................................................................... 25 25(24). Leaves broadly ovate, 9–12 cm wide, base cordate; fruit striate ....... C. striata 25. Leaves elliptic, lanceolate, lanceolate-elliptic, or ovate-lanceolate; fruit smooth .................................................................................................. 26 26(25). Leaves shiny when dry, strongly reticulate on both surfaces ............ C. sp. A 26. Leaves opaque, not reticulate to discretely veined on both surfaces .... 27 27(26). Leaves lanceolate or lanceolate-elliptic; fruit with perianth lobes imbricate ................................................................................ C. acuminata 27. Leaves ovate or elliptic-obovate; fruit with perianth lobes valvate ...... 28 28(27). Leaves with base acute; fruit pyriform or ovoid ..................................... 29 28. Leaves with base cordate; fruit rounded ................................................ 30 29(28). Leaves obovate or elliptic, the base narrowly cordate; petiole hirsute; fruit pyriform ............................................................................ C. coronata 29. Leaves ovate, the base acute; petiole glabrescent; fruit ovoid ...... C. declinata 30(28). Leaves obovate to elliptic-obovate ........................................................... 31 30. Leaves ovate, broadly oblong or elliptic .................................................. 32 31(30). Leaves slightly rugose, secondary veins 5–8; flowers sessile ............... .......................................................................................... C. charitostachya 31. Leaves smooth, secondary veins 9–14; flowers pedicellate ...... C. marginata 32(30). Leaves ovate; fruit 0.6–1.2 cm diameter ......................................... C. excelsa 32. Leaves broadly oblong or elliptic; fruit 1.5–2 cm diameter ...... C. ascendens Coccoloba acuminata Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 176. 1817. Shrub or small slender tree; flowers white. Riparian forests, 200–300 m; Bolívar (middle Río Botanamo). Apure, Barinas, Distrito Federal, Mérida, Miranda, Portuguesa, Yaracuy, Zulia; Central America, Colombia, Guyana, Ecuador, Peru, Brazil, Bolivia. Coccoloba ascendens Duss ex Lindau, Bot. Jahrb. Syst. 13: 154. 1890. —Guayapón, Poa-wip (Piaroa). Liana or tree with heavily coriaceous leaves and large fruits for the genus; flowers white. Gallery forests, evergreen lowland and montane forests, 200–1200 m; Delta

Amacuro (Río Amacuro, Río Cuyubini), Bolívar (Altiplanicie de Nuria, Gran Sabana, upper Río Caura, Río Uairén), Amazonas (Río Baría, Río Cataniapo, Río Cunucunuma, Río Jenita on Río Ocamo, Río Mawarinuma). Apure, Barinas, Miranda, Sucre, Táchira, Yaracuy, Zulia; West Indies, Colombia, Trinidad, Guyana, Suriname, French Guiana, Peru, Brazil. ◆Fig. 300. Coccoloba ascendens is similar to the poorly known Coccoloba nutans Kunth of Peru. Coccoloba caracasana Meisn. in A. DC., Prodr. 14: 157. 1856. —Alerón. Coccoloba cyclophylla S.F. Blake, Contr. U.S. Natl. Herb. 20: 238. 1919.

Coccoloba 353

Small to medium tree to 15 m tall; flowers white. Edges of savannas, semideciduous forests, 100–200 m; Amazonas (Cacurí savannas, Isla Ratón, upper Río Orinoco). Anzoátegui, Apure, Aragua, Barinas, Carabobo, Cojedes, Distrito Federal, Guárico, Lara, Mérida, Miranda, Monagas, Portuguesa, Yaracuy, Zulia; Mexico, Central America, Colombia. Coccoloba charitostachya Standl., Lloydia 2: 176. 1939. Shrub to liana; flowers white. Edge of savannas, gallery forests, 200–1000 m; Bolívar (Kurún-tepui near Cerro Venado, Río Abacapá, Río Torono). Barinas; Guyana, Brazil. Coccoloba coronata Jacq., Enum. Syst. Pl. 19. 1760. —Lata. Coccoloba novogranatensis Lindau, Bot. Jahrb. Syst. 13: 192. 1890. Coccoloba caribaea Urb., Symb. Antill. 5: 337. 1907. Small tree 3–10 m tall; flowers white. Deciduous forests, 100–300 m; Bolívar (Cerro La Cruz, El Manganeso, El Manteco, east of El Miamo, Los Testigos, Represa Guri). Anzoátegui, Barinas, Dependencias Federales (Islas Los Testigos). Distrito Federal, Falcón, Guárico, Lara, Mérida, Miranda, Nueva Esparta, Portuguesa, Sucre, Zulia; Central America, West Indies, Colombia, Ecuador, Peru, Brazil, Paraguay, Argentina. Coccoloba declinata (Vell.) Mart., Flora 20(Beibl.): 20. 1837. —Polygonum declinatum Vell., Fl. Flumin. 162. 1825 [1829]; Fl. Flumin. Icon. 4: 41. 1827 [1831]. Liana with slender stems or understory tree to 4 m tall; flowers pale green. Evergreen lowland to montane forests, 200–900 m; Delta Amacuro (Los Castillos), Bolívar (Altiplanicie de Nuria), Amazonas (Río Baría, upper Río Matacuni). Guárico, Miranda, Sucre; Suriname, French Guiana, Brazil. ◆Fig. 297. Coccoloba densifrons Mart. ex Meisn. in Mart., Fl. Bras. 5(1): 26, pl. 7. 1855. Small tree; flowers white. Evergreen lowland forests, 100–200 m; Bolívar (middle Río Botanamo). Apure, Barinas, Falcón, Guárico, Miranda, Portuguesa, Táchira, Yaracuy,

Zulia; Colombia, Guyana, Ecuador, Peru, Brazil, Bolivia. Coccoloba dugandiana A. Fernández, Mutisia 5: 1. 1952. —Karaw-karaw (Pemón), Karaw-yek (Pemón), Uvero. Tree to 16 m tall; flowers white. Granitic outcrops, gallery forests, evergreen lowland forests, 100–400 m; Bolívar (Los Pijiguaos, Río Caroní, Río Ikabarú, Río Karún, Río Parguaza), Amazonas (Puerto Ayacucho, Río Cataniapo, Río Manapiare, Río Sipapo, Río Ventuari, Samariapo, San Antonio del Orinoco, Yutajé). Apure, Miranda, Monagas, Táchira, Yaracuy; Colombia. ◆Fig. 301. Coccoloba excelsa Benth., London J. Bot. 4: 624. 1825. —Chinay-yek (Pemón), Palo perro de agua, Uña de murciélago. Coccoloba parimensis Benth., London J. Bot. 4: 626. 1845. Coccoloba bracteolosa Meisn. in Mart., Fl. Bras. 5(1): 30. 1855. Coccoloba parimensis var. schomburgkii Meisn. in Mart., Fl. Bras. 5(1): 35. 1855. Coccoloba paraensis Meisn. in Mart., Fl. Bras. 5(1): 38. 1855. Coccoloba micropunctata Eyma, Recueil Trav. Bot. Néerl. 4: 1. 1932. Decumbent shrub, small tree, or highclimbing vine; flowers white. Gallery forests, savannas, bana (white-sand shrubland), evergreen lowland forests, near sea level to 800 m; Delta Amacuro (Altiplanicie de Nuria, Río Amacuro, Río Grande), Bolívar (Canaima, El Dorado, Gran Sabana, Los Pijiguaos, Represa Guri, Río Acanán, Río Caroni, Río Chicanán, upper Río Cuyuní, Río Torono, Urimán), Amazonas (widespread). Aragua, Distrito Federal, Miranda, Sucre, Táchira, Yaracuy, Zulia; Panama, Colombia, Trinidad, Guyana, Suriname, French Guiana, Peru, Brazil. ◆Fig. 302. We are treating Coccoloba excelsa here as a variable species, particularly in growth habit, leaf pubescence, and punctations. Further field work will help determine if this is a wise approach, or if a finer species circumscription should instead be used. Coccoloba fallax Lindau, Bot. Jahrb. Syst. 13: 172. 1890. —Aragueque, Arahueque, Uverillo.

354

P OLYGONACEAE

Coccoloba caurana Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 22: 73. 1940. Multi-stemmed tree to 15 m tall; flowers red and white. Semideciduous to evergreen lowland forests, 100–500 m; Delta Amacuro (Los Castillos de Guayana, Río Grande), northern Bolívar, Amazonas (Río Manapiare, Yutajé). Barinas, Mérida, Miranda, Sucre, Yaracuy; Colombia, Trinidad, Ecuador. ◆Fig. 305. Coccoloba gymnorrhachis Sandw., Bull. Misc. Inform. Kew 1932: 221. 1932. Liana; flowers greenish white. Evergreen slope forests, 400–500 m; Bolívar (Macizo del Chimantá [Abacapá-tepui]). Guyana, Suriname, French Guiana, Brazil. Coccoloba latifolia Poir. in Lam., Encycl. 6: 61, illus. 316, fig. 4. 1804. —Coccoloba grandis Benth., London J. Bot. 4: 624. 1845. —Nairu-cuaja (Warao), Uvero. Multi-stemmed tree to 15 m tall; flowers white. Semideciduous to evergreen forests, 50–1000 m; Delta Amacuro (Caño Angosturita east of Pedernales, Cãno Atoiba north of Caño Araquao, Cãno Capurito, Río Cuyubini), Bolívar (Altiplanicie de Nuria, El Palmar, Represa Guri, middle Río Chiguao, Río Cuyuní, upper Río Paragua, San Francisco de Yuruaní, Amazonas (Raudal Arata on Río Ocamo, Salto Salas on upper Río Orinoco). Anzoátegui, Monagas, Sucre; Trinidad, Guyana, Suriname, French Guiana, Brazil. Coccoloba llewelynii R.A. Howard, J. Arnold Arbor. 42: 89. 1961. —Brusquillo blanco, Guayapapón, Potoruco, Uvero blanco. Coccoloba bolivarana Ll. Williams, Trop. Woods 68: 39. 1941, nom. nud. Shrub or small tree to 6 m tall; flowers white. Deciduous to evergreen forests, edges of savannas, 50–300 m; Bolívar (El Manteco, Puerto Ordaz, Represa Guri, Río Botanamo, Río Cuchivero, San Felíx), Amazonas (lower Río Baría). Anzoátegui, Barinas, Distrito Federal, Monagas, Nueva Esparta, Portuguesa, Sucre, Zulia. ◆Fig. 304. Coccoloba lucidula Benth., London J. Bot. 4: 627. 1845. —Guaya-papón sabanero, Guayapapón. Coccoloba sagotii Lindau, Bot. Jahrb. Syst. 13: 184. 1890.

Liana to small tree 3–7 m tall; upper surface of leaves shiny; flowers white; fruits black. Moist forests and deciduous forest on quarzite-ferruginous outcrops, 50–400 m; Bolívar (Isla Anacoco, Represa Guri, middle Río Botanamo, Tumeremo). Guyana, Suriname, French Guiana. Coccoloba marginata Benth., London J. Bot. 4: 626. 1845. —Arahuequito, Baquerito, Dukuadi-jodu (Yekwana), Etuburrucuaja (Warao), Guayapapón, Uña de gato. Coccoloba guianensis Meisn., Linnaea 21: 264. 1848. Coccoloba trinitatis Lindau, Bot. Jahrb. Syst. 13: 182. 1890. Liana or tree with scrambling branches to 20 m tall; flowers white. Gallery forests, semideciduous to evergreen forests, 50–1400 m; Delta Amacuro (Caño Araguao, Caño Atoiba, Caño Güiniquina, Río Grande, Río Orocoima), Bolívar (Canaima, Represa Guri, Río Asa, Río Botanamo, Río Carrao, Río Caura, Río Chicanán, Río Paragua, upper Río Supamo, Upata), Amazonas (Caño San Miguel, La Esmeralda, Maroa, Río Cariche, Río Casiquiare, Río Cataniapo, Río Matacuni, Río Puruname, lower Río Siapa, San Carlos de Río Negro to Solano Road, Tamatama). Monagas; Colombia, Trinidad, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia. ◆Fig. 303. Coccoloba mollis Casar., Nov. Stirp. Bras. 72. 1844. —Cowicowinae (Guahibo), Karaw-karaw (Pemón). Coccoloba polystachya Wedd., Ann. Sci. Nat. Bot. sér. 3, 13: 261. 1850. Small tree to 15 m tall; flowers white. Lowland seasonally flooded to evergreen montane forests, 200–600 m; Bolívar (Ikabarú, upper Río Caroní), Amazonas (near Canaripó, Caño Topochito 60 km northeast of Puerto Ayacucho). Miranda, Monagas, Táchira, Yaracuy; Colombia, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. Coccoloba obtusifolia Jacq., Enum. Syst. Pl. 19. 1760; Select. Stirp. Amer. Hist. 11. 1763. —Jarizo, Mangle. Shrub to 3 m tall; flowers white. Gallery forests, 100–300 m; Bolívar (lower Río Caura), Amazonas (middle Río Orinoco, San Fernando de Atabapo). Anzoátegui, Apure,

Coccoloba 355

Barinas, Falcón, Guárico, Miranda, Portuguesa, Zulia; Colombia, Peru. ◆Fig. 306.

tama). Monagas; Ecuador, Peru, Brazil, Bolivia. ◆Fig. 308.

Coccoloba ochreolata Wedd., Ann. Sci. Nat. Bot. sér. 3, 13: 259. 1850. Liana; flowers white; fruit green. Gallery forests, 300–500 m; Bolívar (Represa Guri, Río Uiri-yuk on upper Río Cuyuní), Amazonas (Río Orinoco near Puerto Ayacucho). Brazil.

Coccoloba striata Benth., London J. Bot. 4: 626. 1845. —Coccoloba pittieri R. Knuth ex Pittier, Man. Pl. Usual. Venez. 355. 1926. —Arajueque montañero, Arizo, Canilla de venado, Guayapapón, Lata, Melero, Uvero, Uvero negro. Small tree; flowers white. Edges of deciduous forests to wet forests, 200–1200 m; Bolívar (Altiplanicie de Nuria, El Manganeso, El Palmar, El Pao, Represa Guri, San Felíx, San Francisco de Yuruaní), Amazonas (slopes of Cerro Marahuaka, Puerto Ayacucho, Río Cataniapo, middle Río Ventuari, San Carlos de Río Negro). Anzoátegui, Apure, Carabobo, Distrito Federal, Guárico, Lara, Miranda, Monagas, Nueva Esparta, Sucre; Trinidad, Guyana, Brazil. ◆Fig. 298.

Coccoloba orinocana R.A. Howard, J. Arnold Arbor. 42: 90. 1961. —Aledodö (Yekwana), Pankecho. Shrub or tree to 5 m tall; flowers white. Semideciduous to wet forests, edges of savannas, 50–500 m; Bolívar (Los Pijiguaos, Maniapure, Río Paragua, Río Parguaza, Río Parhueña), Amazonas (Cacurí, base of Cerro Aracamuni, Puerto Ayacucho, Salto Yureba on Río Yureba). Barinas, Guárico, Portuguesa. ◆Fig. 299. Coccoloba ovata Benth., London J. Bot. 4: 627. 1845. —Arizo, Jariso, Mangle. Shrub to small tree forming dense thickets; flowers white; fruits edible. Flooded and gallery forests, 50–200 m; Bolívar (Ciudad Bolívar, Río Caura, Río Cuyuni, middle Río Orinoco, Río Parhueña), Amazonas (Caño Marueta affluent of Río Ventuari, Isla Ratón, Puerto Ayacucho, Río Atabapo). Anzoátegui, Apure, Barinas, Guárico, Miranda, Portuguesa, Zulia; Colombia, Guyana, Suriname, French Guiana, Brazil. Coccoloba schomburgkii Meisn., Linnaea 21: 265. 1848. —Huonay-key (Pemón). Shrub; flowers white. Gallery forests, montane forests, 100–2000 m; Bolívar (Amaruay-tepui, Cerro Sarisariñama, Gran Sabana, Ilú-tepui, Murisipán-tepui, Ptaritepui, Uei-tepui), Amazonas (Cerro Marahuaka). Guyana, Brazil. ◆Fig. 307. Coccoloba spruceana Lindau, Bot. Jahrb. Syst. 13: 162. 1891. —Chaparro rebalsero. Shrub to small tree; flowers white. Flooded forests, near sea level to 200 m; Delta Amacuro (Río Grande), Amazonas (Caño Marueta off Río Ventuari, Caño Yagual above mouth of Río Atabapo, Cariche, Río Atabapo, Río Baría, Río Casiquiare, Río Yatúa, Santa Bárbara del Orinoco, Tama-

Coccoloba uvifera Salzm. ex Lindau, Bot. Jahrb. Syst. 13: 186. 1890. Small tree with rounded, coriaceous leaves; flowers white. Cultivated and locally escaped, near sea level to 200 m; Amazonas (Puerto Ayacucho). Aragua, Carabobo, Dependencias Federales, Distrito Federal, Falcón, Mérida, Portuguesa, Sucre, Vargas, Zulia. U.S.A.(southern Florida), Mexico, Central America, Colombia, Trinidad, Guyana, French Guiana, Ecuador, Peru, Brazil; introduced and cultivated in the paleotropics. The fruits of Coccoloba uvifera are edible and have an acid flavor. They are used to make jelly and beverages. Coccoloba wurdackii R.A. Howard, J. Arnold Arbor. 13: 162. 1980. —Pene del diablo. Liana or shrub; flowers white. Edges of white-sand savannas and shrublands, whitesand scrub (bana), seasonally flooded blackwater forests, 100–200 m; Amazonas (Caño San Miguel, Maroa, Río Baría, San Carlos de Río Negro). Apure, Táchira; Colombia. Coccoloba sp. A Small shrub; flowers greenish white. Edges of forests, 100–200 m; Amazonas (Pimichín to Yavita road). This distinctive taxon is known from a single collection, Maguire & Wurdack 35651 (A, NY).

356

P OLYGONACEAE

Coccoloba sp. B Shrub; flowers white. Semideciduous forests, 200–300 m; Amazonas (Raudal Arata on Río Ocamo). This taxon is known from a single collection, A. Fernández 6647 (A, MO, PORT). Coccoloba sp. C. —Uña de gato. Liana; flowers white. Evergreen lowland forests, 100–200 m; Amazonas (Río Cunu-

cunuma, mouth of Río Matacuni, Río Mavaca, Río Ocamo, San Carlos de Río Negro). Coccoloba sp. D Liana; fruits green. Evergreen lowland forests, 300–400 m; Bolívar (El Cácaro near Río Caura, Río Paragua). This taxon is known from two collections, Marin 218 (MO, PORT) and Stergios et al. 15220 (MO, PORT, TFAV, VEN).

Fig. 297. Coccoloba declinata

Fig. 298. Coccoloba striata

Coccoloba 357

Fig. 299. Coccoloba orinocana

Fig. 300. Coccoloba ascendens

358

P OLYGONACEAE

Fig. 301. Coccoloba dugandiana

Coccoloba 359

Fig. 302. Coccoloba excelsa

Fig. 303. Coccoloba marginata

360

P OLYGONACEAE

Fig. 304. Coccoloba llewelynii

Fig. 305. Coccoloba fallax

Coccoloba 361

Fig. 306. Coccoloba obtusifolia

Fig. 307. Coccoloba schomburgkii

362

P OLYGONACEAE

Fig. 308. Coccoloba spruceana

Polygonum 363

3. POLYGONUM L., Sp. Pl. 359. 1753. Persicaria (L.) Mill., Gard. Dict. abr. ed. 4. 1754. Herbs, annual or perennial, sometimes scandent, stems glabrous to densely tomentose, hispid, or with stipitate-glandular hairs. Ochreae usually conspicuous, membranaceous or sometimes scarious, often fringed with strigose cilia, the margin sometimes recurved and green. Leaves alternate; blades mostly linear to lanceolate or ovate, occasionally sagittate, cordate or hastate, always longer than wide; petioles slender, terete. Inflorescence terminal, spicate or of spicate panicles, racemes, or cymes, sometimes with solitary flowers or congested partial inflorescences in leaf axils. Flowers small, bisexual, articulated to pedicels, usually clustered within an ochreola; perianth of 4 or 5 subequal tepals, these white, green, red, or pink; stamens 3–9; filaments often unequal in length, occasionally adnate to the tepals. Ovary lenticular or trigonous; styles 2 or 3; stigma capitate. Achene usually included in a membranaceous perianth, lenticular or trigonous or plano-convex, often beaked, shining. Cosmopolitan, mainly north-temperate areas; ca. 300 species, 9 in Venezuela, 5 of these in the flora area. Key to the Species of Polygonum 1. 1. 2(1). 2. 3(2). 3.

4(2).

4.

Tepals, leaves, and ochreae conspicuously dark-punctate ....... P. punctatum Tepals, leaves, and ochreae inconspicuously pellucid-punctate .............. 2 Ochreae and ochreolae long-ciliate at the apex when mature ................. 3 Ochreae and ochreolae usually without cilia when mature, these, if present when young, weak and poorly developed ................................ 4 Stems slender, glabrous; leaf margin minutely ciliate; ochreae with strigose cilia to 8 mm long............................................. P. hydropiperoides Stems robust, often conspicuously whitish gray-strigose; leaf margin slightly to densely appressed-strigose; ochreae with coarse strigosewhitish cilia to 15 mm long ................................................ P. acuminatum Stems slender, the internodes not conspicuously hollow; achenes no longer than 2.5 mm long, oblong to obovoid, one face flat and the other convex ......................................................................................... P. glabrum Stems stout, the internodes hollow; achenes 3–3.5 mm long, orbicular, the faces slightly to conspicuously concave near the apex ............... .............................................................................................. P. ferrugineum

Polygonum acuminatum Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 178. 1817. —Camuco-anajoru (Warao). Polygonum acuminatum var. glabrecens Meisn. in A. DC., Prodr. 14: 114. 1856. Polygonum acuminatum var. weddelii Meisn. in A. DC., Prodr. 14: 114. 1856. Perennial herb to 1 m tall; flowers white. Abandoned fields, flooded savannas, 50–200 m; Delta Amacuro (Caño Araguao, Caño Iburama, Caño Jobure near the mouth of Río Amacuro, Caño Macareo, Tucupita), Bolívar

(La Paragua, Santa Elena de Uairén), Amazonas (Río Casiquiare, Río Manapiare, lower Río Putaco). Apure, Aragua, Lara, Miranda, Monagas, Sucre; Mexico, Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Chile, Paraguay, Argentina, Uruguay. ◆Fig. 309. Polygonum ferrugineum Wedd., Ann. Sci. Nat. Bot. sér. 3, 13: 252. 1849. Floating or rooted plant to 1.5 m tall; flow-

364

P OLYGONACEAE

Fig. 309. Polygonum acuminatum

Ruprechtia 365

ers white and pink. Abandoned wet fields, ca. 50 m; Delta Amacuro (Caño Jota-Sabuca near Caño Mariusa, Caño Mánamo). Apure, Aragua, Carabobo, Cojedes, Miranda, Portuguesa, Yaracuy; El Salvador, Hispaniola, Colombia, French Guiana, Peru, Brazil, Bolivia, Paraguay, Argentina, Uruguay. Polygonum glabrum Willd., Sp. Pl. 2: 447. 1799. Polygonum densiflorum Meisn. in Mart., Fl. Bras. 5(1): 13. 1855. Polygonum portoricense Bertero ex Small, Mem. Dept. Bot. Columbia Coll. 1: 46. 1895, nom. illeg. Herb to 60 cm tall; flowers white. Abandoned fields, 50–200 m; Delta Amacuro, Amazonas (Raudal Arata on Río Ocamo). Aragua, Guárico, Miranda, Portuguesa, Zulia; Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Chile, Paraguay, Argentina, Uruguay.

Polygonum hydropiperoides Michx., Fl. Bor.-Amer. 1: 239. 1803. Perennial herb to 1 m tall; flowers pink. Abandoned fields, 50–200 m; Delta Amacuro (between Tucupita and La Horqueta), Amazonas (Río Matacuni). Monagas; Canada, U.S.A., Mexico, Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Chile, Paraguay, Argentina, Uruguay. Polygonum punctatum Elliott, Sketch Bot. S. Carol. 1: 455. 1821 [1817]. —Ají de morocoto, Osíbu-ajuca (Warao). Annual or perennial herb to 1 m tall; flowers pink and white. Abandoned fields, 50– 200 m; Delta Amacuro (Río Acure), Amazonas (Río Ocamo). Apure, Aragua, Barinas, Carabobo, Distrito Federal, Lara, Mérida, Miranda, Portuguesa, Sucre, Táchira, Trujillo; Central America, West Indies, Colombia, French Guiana, Ecuador, Peru, Brazil, Bolivia, Chile, Paraguay, Argentina, Uruguay.

4. RUPRECHTIA C.A. Mey., Mém. Acad. Imp. Sci. St. Pétersbourg Sér. 6, Sci. Math. 6: 148. 1840. Trees or shrubs, unarmed, evergreen or occasionally deciduous. Twigs striate, glabrous to puberulous or pilose, solid or occasionally hollow. Ochrea (stipules) soon caducous, leaving a ring-like scar on the twigs, intrapetiolar, glabrous to puberulous outside. Leaves alternate, deciduous, simple, pinnately veined; petioles short, often canaliculate above. Inflorescences open pleiothyrsi, variously branched from the base, mostly axillary, sometimes terminal or occasionally very short and borne on lateral short shoots, partial inflorescences monochasia, subtended by small ovatetriangular bracts. Flowers bisexual (plants dioecious). Staminate flowers with articulate, slender pedicels; tepals 6, subequal, ovate-lanceolate to oblanceolate, acute to rounded apically, connate for 1/3 their length or free almost to the base; petals free; stamens 9, in 2 whorls, the outer of 6, the inner of 3; anthers versatile. Gynoecium rudimentary; pistillate flowers with 3 sepals, united at the base; petals 3, much smaller than the sepals; staminodes 9; ovary tricarpellate, unilocular, styles 3, short; stigmas globose or linear; ovule 1. Achene protected by perianth tube, sepals accrescent, reflexed or inflexed, achene oblong, endosperm ruminate, cotyledons foliaceous. Neotropics (mostly drier areas from Mexico and Trinidad to Argentina); 20 species, 6 in Venezuela, 5 of these in the flora area. Key to the Species of Ruprechtia 1. 1.

Leaves and inflorescence persistently ferruginous-hirsute, midrib on upper surface of leaves depressed ........................................... R. howardiana Leaves and inflorescence glabrous, or if pubescent never ferruginoushirsute, midrib on upper surface of leaves elevated ............................ 2

366

2(1). 2. 3(2). 3. 4(3). 4.

P OLYGONACEAE

Leaf margin undulate; sepals spatulate or linear-spatulate, 3-veined; fruiting perianth wings broadly obovate-oblong .................... R. ramiflora Leaf margin flat; sepals lanceolate or linear, 1-veined or without veins; fruiting perianth wings linear to obovate ............................................. 3 Twigs and internodes hollow; sepals lanceolate, acute, 3.5–5 mm long ................................................................................................ R. tangarana Twigs and internodes solid; sepals linear, ca. 0.6 mm long ..................... 4 Leaves ovate to ovate-lanceolate; sepals spatulate, 3-veined, 1–3 mm long ............................................................................................ R. laxiflora Leaves lanceolate; sepals linear, 1-veined, 12–17 mm long ...... R. tenuiflora

Ruprechtia howardiana Aymard & P.E. Berry, Novon 9: 313. 1999. Small tree to 3 m tall; flowers white. Deciduous forests, 100–300 m; Bolívar (Serranía Baraguán). Endemic. ◆Fig. 311. Ruprechtia laxiflora Meisn. in Mart., Fl. Bras. 5(1): 56. 1855. Small tree; flowers white. Seasonally flooded forests, 100–200 m; Amazonas

(around Puerto Ayacucho). Colombia, Peru, Brazil, Paraguay, Argentina, Uruguay. Ruprechtia ramiflora (Jacq.) C.A. Mey., Mém. Acad. Imp. Sci. St. Pétersbourg Sér. 6, Sci. Math. 6: 150. 1840. —Triplaris ramiflora Jacq., Select. Stirp. Amer. Hist. 14. 1763. —Palo de agua. Triplaris coriacea H. Karst., Fl. Columb. 2: 131, pl 169. 1862 [1866]. —Ruprechtia

Fig. 310. Ruprechtia tenuiflora

Ruprechtia 367

Fig. 311. Ruprechtia howardiana

coriacea (H. Karst.) S.F. Blake, Contr. U.S. Natl. Herb. 20: 239. 1919. Ruprechtia hamanii S.F. Blake, Contr. Gray Herb. n.s. 52: 31. 1918. Ruprechtia concina Pittier, Bol. Minist. RR. EE. no. 8/9, reprinted in Arb. Arbust. Venez. 6–8: 1. 1927. Shrub or tree to 20 m tall; flowers pale green. Deciduous to semideciduous and seasonally flooded forests, 50–200 m; Bolívar (Ciudad Bolívar, Mapire, Puerto Ordaz, San Felíx). Anzoátegui, Apure, Aragua, Cojedes, Distrito Federal, Falcón, Guárico, Lara,

Miranda, Portuguesa, Sucre, Táchira, Zulia; Colombia, Guyana, Brazil. Ruprechtia tangarana Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 22(2): 74. 1940. Small tree to 10 m tall; flowers white. Evergreen forests, 100–200 m; Bolívar (Río Oris in upper Río Paragua basin), Amazonas (Puerto Ayacucho, Río Cuao basin). Colombia, Peru, Brazil. Ruprechtia tenuiflora Benth., London J. Bot. 4: 629. 1845. —Ruprechtia amen-

368

P OLYGONACEAE

tacea Meisn. in Mart., Fl. Bras. 5(1): 56. 1855. —Guácimo negro, Para-pájaro, Sinátomo (Yekwana). Shrub or tree to 20 m tall; flowers white. Semideciduous and flooded forests, 50–200 m; Delta Amacuro (Santa Catalina), Bolívar

(Ciudad Bolívar, Jabillal, La Paragua, Puerto Ordaz, lower Río Caroni, Río Chiguao, Río Cuyuní, Río Nichare). Anzoátegui, Apure, Guárico, Zulia; Colombia, Guyana, Brazil. ◆Fig. 310.

5. SYMMERIA Benth., London J. Bot. 4: 630. 1845. Shrubs or small trees; occasionally branched from the base, twigs solid, somewhat ferruginous-pilose to tomentulose or glabrous when mature. Ochreae not evident, apparently soon caducous. Leaves alternate, coriaceous; blades entire, pinnately veined, glabrous; petiole short. Staminate inflorescences large open pleiothyrsi with slender branches; pistillate inflorescences with shorter, somewhat stouter branches. Flowers unisexual (plants dioecious). Staminate flowers small, subsessile, glomerate; sepals orbicular, the outer 3 smaller. Stamens numerous; filaments very short. Pistillate flowers pedicellate, calyx 6-parted, outer segments small, oblong, slightly accrescent, inner tepals cordate-ovate, erect, connivent, accrescent and closely investing the triangulate achene. Achenes elongated, pyramidal, green, reddish when mature. Seeds with endosperm ruminate; cotyledons foliaceous. Panama, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, tropical West Africa (Sierra Leone); 1 species.

Fig. 312. Symmeria paniculata

Symmeria paniculata Benth., London J. Bot. 4: 630. 1845. —Chaparro de agua, Manteco rebalsero, Palo perro de agua, Tin-tin. Shrub or tree to 30 m tall; flowers white. Flooded evergreen forests, gallery forests, 50–200 m; Delta Amacuro (Corisal, Sacupana), Bolívar (Carichana, Ciudad Bolívar, El Dorado, lower and middle Río Caura, Río Cuyuní, Río Oris, Río Paragua), Amazonas (Caño Ucata, La Esmeralda, Marueta, Río Casiquiare, Río Manapiare, middle Río Ventuari, San Antonio del Orinoco). Anzoátegui,

Apure, Barinas, Guárico, Zulia; other distribution as in genus. ◆Fig. 312. The bark of Symmeria paniculata is used locally to treat diarrhea and dysentery.

Triplaris 369

6. TRIPLARIS Loefl. ex L., Syst. Nat. ed. 10, 2: 881, 1360. 1759. Trees, bark often peeling, twigs striate. Leaves alternate, petiolate, stipules (ochreae) intrapetiolar, to 32 cm long, deciduous, pilose to tomentose; blades entire, oblong or lanceolate to ovate, acute to acuminate, base acute or rounded, glabrous or tomentose to strigose, glandular-punctate on the lower surface. Inflorescence axillary or terminal, enclosed in a deciduous spathe-like stipule, tomentose, bracteoles ovate, acute to acuminate, fissured on two sides. Flowers unisexual (plants dioecious). Staminate flowers 3–5 in each partial inflorescence, sessile or subsessile; tepals 6, subequal, ovate, obtuse or lanceolate, pilose or strigose; stamens 9, exserted. Pistillate flowers one in each partial inflorescence, pedicellate; outer tepals basally connate, forming an urceolate-campanulate to globose tube, lobes lanceolate, acute, spreading, enlarged into wings in fruit; inner tepals smaller, linear to ovate, free or united to perianth tube; ovary trigonous; styles 3; stigmas linear extending down styles. Achenes enclosed in the persistent 3-winged perianth tube. Neotropics (mostly western Amazon region); 17 species, 4 species in Venezuela, all in the flora area. Several species are cultivated for the attractive and unusual fruits. Key to the Species of Triplaris 1. 1. 2(1). 2. 3(2). 3.

Inflorescence axis puberulous or glabrate ............................... T. weigeltiana Inflorescence axis densely gray-yellow- to brownish yellow-pubescent ................................................................................................................ 2 Staminate perianth 2–3 mm long; stamens 3–4 mm long; achenes slightly trigonous to terete basally, sulcate, not veined ..................... T. americana Staminate perianth 5–7 mm long; stamens 5–8 mm long; achene triquetrous to 3-winged, not sulcate, veined .................................................. 3 Petals basally adnate to fruiting perianth; achene punctate-granulate, with a short beak .................................................................. T. cumingiana Petals not basally adnate to fruiting perianth; achene not punctategranulate, without a beak .................................................... T. caracasana

Triplaris americana L., Syst. Nat. ed. 10, 2: 881, 1360. 1759. —Biyw mamo ka hi, Santa Maria. Triplaris schomburgkiana Benth., London J. Bot. 4: 628. 1845. Triplaris felipensis Wedd., Ann. Sci. Nat. Bot., sér 3, 3(13): 263. 1849. Triplaris pavonii Meisn. in Mart., Fl. Bras. 5(1): 172. 1856. Tree to 27 m tall; fruiting perianth pale yellow to reddish when mature. Gallery forests, flooded evergreen forests, granitic outcrops, 100–200 m; Bolívar (La Urbana, Pararuma, Río Villacoa), Amazonas (La Esmeralda, Puerto Ayacucho, Río Mavaca, Río Ocamo, Río Puruname, Tamatama). Apure, Barinas, Carabobo, Falcón, Mérida, Portuguesa, Táchira, Yaracuy, Zulia; Panama, Co-

lombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. Triplaris caracasana Cham., Linnaea 8: 137. 1833. —Triplaris caracasana var. genuina Meisn. in A. DC., Prodr. 14: 172. 1856. Triplaris caracasana var. vargasii Meisn. in A. DC., Prodr. 14: 172. 1856. Tree to 25 m tall; fruiting perianth pale yellow to reddish when mature. Seasonally flooded lowland forests, 100–200 m; Bolívar (Río Pamoni tributary of the Río Casiquiare). Aragua, Carabobo, Distrito Federal, Falcón, Mérida, Miranda, Portuguesa, Yaracuy, Zulia. Triplaris cumingiana Fisch. & C.A. Mey. ex C.A. Mey., Mém. Acad. Imp. Sci. St.

370

P OLYGONACEAE

Pétersbourg Sér. 6, Sci. Math. 7: 148. 1845. Tree to 20 m tall; fruiting perianth pale yellow to reddish when mature. Flooded evergreen forests, 100–200 m; Amazonas (lower Río Siapa). Barinas, Mérida, Portuguesa, Táchira, Zulia; Panama, Colombia, Ecuador, Peru. Triplaris weigeltiana (Rchb.) Kuntze, Revis. Gen. Pl. 3(2): 271. 1898. —Blochmannia weigeltiana Rchb., Consp. Regn. Veg. 1828. —Maria, Palo Maria, Santa Maria. Triplaris surinamensis Cham., Linnaea 8: 130. 1833. Tree to 25 m tall; fruiting perianth pale yellow to reddish when mature. Evergreen lowland forests, 50–200 m; Delta Amacuro (Río Grande, Caño Curiapito, Caño Jana-

Fig. 313. Triplaris weigeltiana

mana), Bolívar (Cerro Coroba, El Dorado, Río Botanamo, Río Paragua, Río Supamo). Apure, Barinas, Miranda; Colombia, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 313.

Eichhornia 371

PONTEDERIACEAE by Charles N. Horn Herbaceous aquatic annuals or perennials, rooted in mud or free-floating. Stems of 2 types, vegetative and flowering; vegetative stem indeterminate, bearing many leaves, glabrous; flowering stem determinate, bearing a single leaf, a spathe, and a terminal inflorescence. Leaves of 2 types, sessile and petiolate; sessile leaves submersed, simple, linear, glabrous, entire, acuminate to obtuse at apex; petiolate leaves simple, glabrous, entire, obtuse to acuminate at apex. Stipules when present marcescent, transparent, truncate at apex, sheathing. Inflorescence a panicle, spike, or reduced to a single flower; peduncle glabrous to glandular-pubescent, the subtending spathe bract-like, folded, glabrous, with an acute to caudate apex. Flowers bisexual, perigynous, sessile. Perianth tubular or funnelform, mauve, blue, or white, glandular-pubescent, zygomorphic, 6-lobed, the lobes oblong to oblanceolate, obtuse to acuminate at apex. Stamens 1, 3, 4, or 6 (3 or 6 in the flora area), adnate to perianth; filaments pubescent with multicellular or glandular hairs; anthers rounded. Carpels 3, fused, with 1 or 3 developing to maturity, placentation parietal or basal; ovules 1–many; style pubescent with multicellular hairs or glabrous; stigma with single-celled hairs. Fruit a capsule with persistent unmodified perianth or a utricle with thickened, ridged perianth. Seeds longitudinally winged or smooth. Pantropics, extending into temperate regions; 7 genera and ca. 33 species, 3 genera and 7 species in the flora area. Key to the Genera of Pontederiaceae 1. 1. 2(1). 2.

Inflorescences commonly with > 50 flowers; fruit a utricle, with 1 smooth seed .......................................................................................... 3. Pontederia Inflorescences with < 30 flowers; fruit a capsule with 10–200 seeds; seeds with longitudinal ridges ........................................................................ 2 Petiolate leaf blade coriaceous; perianth limb lobes ovate, > 2 cm long; stamens 6 ............................................................................... 1. Eichhornia Petiolate leaf blade membranous or absent; perianth limb lobes linear to oblanceolate, < 2 cm long; stamens 3 ................................ 2. Heteranthera

1. EICHHORNIA Kunth, Eichhornia 3. 1842, nom. cons. Annuals or perennials, rooting in mud or free-floating. Vegetative stem submersed and growing to surface, or emersed and short; flowering stem emersed, glabrous or glandular-pubescent. Sessile leaves forming a basal rosette or on an elongate stem, acuminate at apex; petiolate leaves floating or emersed, cordate to oblong, acuminate to obtuse at apex. Inflorescence a panicle or spike developing over 1–several days, individual flowers open one day only; subtending spathe folded longitudinally, with an acuminate to caudate apex and sometimes with a leaf-like extension. Perianth funnelform, blue, mauve, or white, the lobes oblanceolate, glandular-pubescent on adaxial surface, obtuse to acute at apex. Stamens 6; filaments purple, glandular-pubescent, curved upwards toward apex; anthers yellow, rounded to oblong. Ovary 1- or 3-locular, containing many ovules, heterostylous or homostylous. Fruit a capsule. Seeds many, ovoid; testa with longitudinal wings. Neotropics, Africa; 7 species, 5 in Venezuela, 4 of these in the flora area.

372

P ONTEDERIACEAE

Key to the Species of Eichhornia 1. 1. 2(1). 2. 3(2).

3.

Plants free-floating; vegetative stem highly condensed; petioles usually swollen ...................................................................................... E. crassipes Plants submersed with stem rooted in mud and leaves typically floating; vegetative stem elongate; petioles not swollen .................................... 2 Leaf base cordate; flowers < 2.5 cm across .............................. E. diversifolia Leaf base truncate to obtuse; flowers > 2.5 cm across ............................. 3 Inflorescence generally 2–3 times longer than spathe (excluding perianth), entire perianth 3–4.5 cm long, margin of lobes distinctly erose ....................................................................................................... E. azurea Inflorescence generally shorter than spathe such that only the perianth extends beyond the spathe apex, entire perianth 2–3 cm long, margin of perianth lobes minutely erose ....................................... E. heterosperma

Eichhornia azurea (Sw.) Kunth, Eichhornia 4. 1842. —Pontederia azurea Sw., Prodr. 57. 1788. —Bora, Lirio de agua, Masure, Mosori. Robust floating aquatic with stem rooted in mud; submersed sessile leaves linear, ribbon-like; petiolate leaves ovoid; perianth blue or white with a yellow spot on upper central lobe, heterostylous. Calm backwaters, at or near sea level to 100 m; Río Orinoco and its major tributaries in Delta Amacuro and Bolívar. Anzoátegui, Apure, Guárico, Mérida, Monagas, Zulia; Central America, South America except Chile. ◆Fig. 314. Eichhornia crassipes (Mart.) Solms in A. DC. & C. DC., Monogr. Phan. 4: 527. 1883. —Pontederia crassipes Mart., Nov. Gen. Sp. Pl. 1: 9, t. 4. 1824 [1823]. —Bora, Lechuga de agua. Free-floating plant with emersed petiolate leaves; perianth blue with a yellow spot on the upper central lobe, heterostylous. Calm backwaters of rivers, marshes, lakes, and large ponds, at or near sea level to 300 m; Río Orinoco and tributaries in Delta Amacuro, Bolívar, and Amazonas. Common throughout Venezuela; Pantropics and subtropics. ◆Fig. 315. Eichhornia crassipes is a native of northeastern South America (Amazon basin) and introduced to all other parts of the world for its large showy flowers. It is now a worldwide aquatic weed. The extent of petiole swelling is highly variable; the thinner petioles are

found on plants of dense populations in nutrient-rich water. Eichhornia diversifolia (Vahl) Urb., Symb. Antill. 4: 147. 1903. —Heteranthera diversifolia Vahl, Enum. Pl. 2: 44. 1805. Small aquatic plant with stem rooted in mud; submersed sessile leaves linear; petiolate floating leaves ovoid to cordate; perianth mauve with 2 yellow spots on the upper central lobe, homostylous. Small streams, shallow marshes, ephemeral pools, ponds, Mauritia palm swamps, at or near sea level to 1300 m; tributaries of Río Orinoco in Delta Amacuro, Bolívar, and Amazonas. Venezuelan states along Río Orinoco and Caribbean coast; coastal Central America (south from Nicaragua), coastal Colombia, Guyana, Suriname, French Guiana, Ecuador, and Brazil (north of 15°S). Eichhornia heterosperma Alexander, Lloydia 2: 170. 1939. —Bora. Eichhornia venezuelensis Velásquez, Acta Bot. Venez. 6: 367. 1972. Moderate-sized floating aquatic plant with stem rooted in mud; submersed sessile leaves linear; emersed petiolate leaves ovoid; perianth blue or white with a violet region near base of the upper central lobe, heterostylous. Calm backwaters of rivers and marshes, 50–700 m; tributaries of Río Orinoco in Bolívar and Amazonas. Scattered in northern Venezuela; coastal regions of Central America (south from Nicaragua), coastal Colombia, Guyana, Suriname, French Guiana, Ecuador, northern Brazil.

Eichhornia 373

Fig. 314. Eichhornia azurea

Fig. 315. Eichhornia crassipes

374

P ONTEDERIACEAE

Fig. 316. Heteranthera multiflora

2. HETERANTHERA Ruiz & Pav., Fl. Peruv. Prodr. 9, pl. 2. 1794, nom. cons. Annuals or perennials, rooting in mud. Vegetative stem submersed and growing to surface or emersed and short; flowering stem submersed or emersed. Sessile leaves submersed, forming a basal rosette, or emersed on an elongate stem, acuminate to acute at apex; petiolate leaves floating or emersed, cordate, reniform, or oblong, obtuse to acute at apex. Inflorescence a spike or flower solitary, 1–30-flowered, developing over 1–several days; spathes folded or clasping, with acute to caudate apex. Perianth with tepals connate 1/2 or more of its length, yellow, blue-mauve, mauve, or white, tubular or salverform, limb lobes linear to oblanceolate or narrowly elliptic, < 2 cm long, apex obtuse to acuminate. Stamens 3, unequal; filaments yellow or purple, glabrous, glandular-pubescent, or pilose; anthers yellow or purple, rounded, oblong, or sagittate. Ovary incompletely 3-locular; ovules 10–many; style 3-lobed. Fruit an elongate capusle. Seeds 10–200, ovoid, testa with longitudinal wings. Western Hemisphere, Africa; 12 species, 5 in Venezuela, 1 of these in the flora area. Heteranthera multiflora (Griseb.) C.N. Horn, Phytologia 59: 290. 1986. —Heteranthera reniformis var. multiflora Griseb., Abh. Königl. Ges. Wiss. Göttingen 24: 323. 1879.

Annual aquatic herb. Along river edges, near sea level to 100 m; Delta Amacuro (Caño Tucupita). Scattered in northern Venezuela; southeastern U.S.A., Mexico, Central America, Brazil, Paraguay, Argentina. ◆Fig. 316.

3. PONTEDERIA L., Sp. Pl. 288. 1753. Reussia Endl., Gen. Pl. 139. 1836. Annuals or perennials rooting in mud. Vegetative stem submersed and growing to surface or emersed and short; flowering stem glabrous, slightly constricted just below the first node. Sessile leaves submersed, forming a basal rosette, acuminate to acute at apex; petiolate leaves floating or emersed, cordate to reniform, obtuse to acuminate at apex. Inflorescence a spike, developing over several days;

Pontederia 375

emersed peduncle glandular-pubescent or pilose, the subtending spathe folded longitudinally, with an acute apex. Perianth funnelform, blue or white, the lobes oblong, glandular-pubescent or pilose on adaxial surface, obtuse to acuminate at apex, with a bilobed yellow spot on the upper central lobe. Stamens 6; filaments purple, glandular-pubescent; anthers yellow, ovoid to oblong. Carpels 3, only 1 developing to maturity, containing a single ovule, placentation basal, heterostylous or homostylous. Fruit a utricle, with spinulose, toothed, or smooth longitudinal ridges. Seeds ovoid, smooth. Southern Canada to Argentina; 6 species, 5 in Venezuela, 2 of these in the flora area.

Fig. 317. Pontederia rotundifolia

Fig. 318. Pontederia triflora

376

P ONTEDERIACEAE

Key to the Species of Pontederia 1.

1.

Leaves 3–22 cm wide, the base sagittate to subovate; inflorescence usually with > 20 flowers; tristylous (three different kinds of plants, each with different style and stamen lengths) ............................ P. rotundifolia Leaves 0.7–2.8 cm wide, the base truncate to cuneate; inflorescence with 2 or 3 flowers; homostylous (all plants with styles and stamens of same length) ........................................................................................... P. triflora

Pontederia rotundifolia L., Pl. Surin. 8. 1775. Pontederia cordifolia Mart. ex Schult. & Schult. f., Syst. Veg. 7(2): 1142. 1830. Robust aquatic plant. Calm waters of rivers, streams, and marshes, at or near sea level to 1000 m; tributaries of Río Orinoco in Delta Amacuro, Bolívar, and Amazonas. Northern Venezuelan states and states bordering Río Orinoco and its tributaries; larger river systems of Central and South America. ◆Fig. 317.

Pontederia triflora (Endl. ex Seub.) G. Agostini, D. Velásquez & Velásquez, Ernstia 27: 9. 1984. —Reussia triflora Endl. ex Seub. in Mart., Fl. Bras. 3(1): 94. 1847. Small floating aquatic plant. Calm waters of savannas, 100–1000 m; Bolívar (tributaries of Río Orinoco). Apure; Guyana, northern Brazil. ◆Fig. 318.

PORTULACACEAE by Julian A. Steyermark Annual or perennial herbaceous plants, rarely shrubs, generally succulent with mucilaginous cells in stems and leaves, glabrous or pubescent, especially at the nodes. Leaves alternate, opposite, sometimes verticillate around the flowers, basal, entire; stipules scarious, lacerate, or modified into hairy tufts, or absent. Inflorescences solitary, racemose, cymose, or paniculate, sometimes subcapitate terminal or axillary. Flowers small or large, usually actinomorphic, bisexual, usually hypogynous or semi-epigynous. Sepals (or modified bracteoles) usually 2, sometimes unequal, imbricate, persistent or deciduous; petaloid sepals mostly 5 but also (2–)4– 12, distinct, sometimes slightly united at base, often ephemeral or marcescent. Stamens as many as and usually opposite the petals, sometimes more numerous or fewer; filaments filiform, free, or sometimes adnate to the base of the petals or short corolla tube; anthers 2-locular, longitudinally dehiscent. Ovary with 2–5 carpels whose partitions disappear to become 1-locular, superior or partly to completely inferior; ovules usually 2–many, rarely 1, the placenta central or basal; styles usually 2–7, the styles more or less united, rarely a single cleft or lobed style. Fruit capsular, loculicidally dehiscent or circumscissile, the valves the same number as the styles, or rarely hard and nut-like. Seeds 3–many, or 1 or 2 by abortion, compressed, lenticular, often smooth and shining, tuberculate, rugulose, with stellulate or irregular patterns, sometimes strophiolate (with an appendage on the hilum); testa fre-

Portulaca 377

quently crustaceous; embryo horseshoe-shaped, enclosing the abundant starchy perisperm; true endosperm absent. Cosmopolitan; ca. 25 genera and ca. 400 species; 2 genera and 13 species in the flora area. Key to the Genera of Portulacaceae 1.

1.

Ovary partly to completely inferior; capsule circumscissile, the lower part adnate to the calyx tube; plants often pubescent, especially around the leaf axils ................................................................................... 1. Portulaca Ovary superior; capsule 3-valvate, the lower part free, not adnate to the calyx tube; plants glabrous ....................................................... 2. Talinum

1. PORTULACA L., Sp. Pl. 445. 1753. Annual or perennial succulent, herbaceous plants, rarely subligneous at base, glabrous or pubescent, with low, often prostrate, to ascending or erect stems. Root fibrous to sometimes tuberous. Leaves alternate, subopposite, or in whorls around the flowers, flat or terete, entire; stipules scarious or absent, frequently reduced to hairy tufts. Flowers bisexual, terminal, solitary or crowded at the ends of the stems, mainly sessile, small or large. Calyx 2-cleft, deciduous; petaloid sepals (4)5(6), white, yellow, orange, rose, red, scarlet, or rose-purple. Stamens (4–)8–100, inserted at the base of the petals. Ovary partly or rarely wholly inferior, at first several-locular toward the base and 1-locular toward the apex, eventually entirely 1-locular; ovules mostly numerous (1–few), amphitropous, the placenta free and central; style usually 2-lobed, sometimes unlobed, or rarely styles many. Capsule membranaceous, subglobose, globose, or obovoid, often with the persistent base of the style, circumscissile at, below, or above the middle, 1-locular, mostly many-seeded; operculum mostly cupuliform. Seeds black, gray, brown, or rufous, shining or opaque, smooth or variously tuberculate, granulate, foveolate, or variously stellulate. Pantropical and subtropical, but mostly Old World; ca. 150 species, 13 in Venezuela, 11 of these in the flora area. Key to the Species of Portulaca 1.

1. 2(1). 2. 3(2).

3.

Stem glabrous or nearly so to the summit, the axillary trichomes either lacking or scarcely manifest; terminal leaf whorl not concealing or exceeding flower or fruit nor with hairy tufts .......................................... 2 Stem, including the summit, with evident trichomes; terminal leaf whorl concealing or exceeding flower or fruit and bearing hairy tufts ......... 5 Leaves very small, 1.5–4 × 0.5–2.3 mm; entire plant diminutive with stems 2–4 cm long or high ..................................................................... 3 Leaves well developed, 5–40 × 2–15 mm; plant with stems 5–30 cm long or high .................................................................................................... 4 Roots elongated, not tuberoid; petaloid sepals white to pink to rose-purplish; leaves 0.5–1.5 mm wide; capsule subovoid, narrowed at apex, dehiscent below the middle .......................................................... P. pusilla Roots tuberoid; petaloid sepals yellow; leaves 1.7–2.3 mm wide; capsule depressed-hemispheric, rounded at summit, dehiscent near base .................................................................................................... P. pygmaea

378

4(2).

P ORTULACACEAE

Cauline and apical leaves oblong-obovate or broadly spatulate, truncate or broadly rounded at apex, broadest above the middle; petaloid sepals yellow .......................................................................................... P. oleracea 4. Cauline and apical leaves broadly linear or sublinear, narrowly obtuse; petaloid sepals roseate, purplish, or sometimes yellow ..... P. umbraticola 5(1). Seed smooth or nearly so, etuberculate, with or without a thin, pale, membranous covering overlying a finely striolate or reticulate pattern; leaves (living) 10–15 × 4–9 mm, flat or plane, those subtending the flowers 2–4 mm wide; sepals 11–12 mm long; summit of peduncle, including hairy mass, mainly 8–15 mm diameter; stem leafless for (1.5–) 4–10 cm below summit of inflorescence; petaloid sepals yellow, showy, ca. 10 mm long ........................................................................ P. mucronata 5. Seed tuberculate with manifest verrucose, muricate protuberances or strongly rugulose with a stellate ornamentation; leaves 1–4 mm wide, mainly terete or subterete (flattened in P. sedifolia), those subtending the flowers 0.5–1.5 mm wide; sepals 10 mm or less long; summit of peduncle, including hairy mass, 4–11 mm diameter (to 15 mm diameter in P. elatior); stem chiefly foliose to the summit; petaloid sepals smaller, or, if larger in P. elatior, then rose, red, or lavender-red ........ 6 6(5). Entire plant covered with a dense mass of elongated, intertwined trichomes which equal, exceed, surround, or conceal the leaves and flowers; stems short, 4–7 cm tall, procumbent to ascending ........... P. insignis 6. Plant not as above, but if manifesting abundant or elongate pubescence, then the cauline leaves and those surrounding the flowers 8–20 mm long or the stems erect and elongated to 30–40 cm ............................. 7 7(6). Seeds ash gray with prominent stellulate flat ornamentation throughout; axillary trichomes at junction of leaf and stem short, scarce .................................................................................................. P. teretifolia 7. Seeds black when mature, verrucose, muricate, or tuberculate, or with a stellate or tuberculate pattern on the lateral faces; axillary trichomes longer or more numerous ...................................................................... 8 8(7). Uppermost or involucral leaves conspicuously exceeding the flowers; cauline leaves 10–40 mm long; stem usually erect, simple, elongated, with branching, if any, in upper portion, to 30–40 cm long ......... P. elatior 8. Without the above combination, the uppermost or involucral leaves at most slightly exceeding the flowers; cauline leaves and stems usually shorter, more branched, or not erect ..................................................... 9 9(8). Leaves flat or flattish; plants very small with stems chiefly 3–8 cm long; leaves 3–6 × 1–1.5 mm; stamens 5–7(–13); seeds mainly with tuberculate ornamentation .................................................................... P. sedifolia 9. Leaves terete or subterete; plants more robust and taller, with stems to 20 cm; leaves 5–27 × 1–2 mm; stamens (8–)12–30; seeds with stellate ornamentation on the lateral faces ..................................................... 10 10(9). Petaloid sepals yellow; capsule circumscissile below the middle ................................................................................................ P. halimoides 10. Petaloid sepals purple or pink; capsule circumscissile at the middle ......................................................................................................... P. pilosa

Portulaca 379

Portulaca elatior Mart. ex Rohrb. in Mart., Fl. Bras. 14(2): 302. t. 69. 1872. Annual herb with slender roots; leaves alternate, linear, terete, 1–2 mm wide; pubescence of numerous, elongate, fasciculate trichomes in the leaf axils; flowers 2.2 cm across; stamens numerous, 5 mm long; exposed igneous outcrops, 50–300 m; Bolívar (Piedra Elefante and other lajas between Cuidad Piar and Puerto Ordaz). Distrito Federal, Sucre, Táchira; Trinidad-Tobago, eastern Brazil. Portulaca halimoides L., Sp. Pl. ed. 2. 639. 1762. Annual herb with branching, often fleshy roots; stems suberect or prostrate; leaves alternate, oblong-linear, convex; axillary trichomes numerous; flowers 2–6, surrounded by 4–8 involucral leaves. Disturbed, seasonally dry evergreen forests, ca. 200 m; Bolívar (Caño Aliviadero near Tumeremo). Falcón, Guárico, Nueva Esparta, Sucre; U.S.A. (Florida), Central America, West Indies, western South America. Potulaca halimoides is close to P. pilosa, from which it may be distinguished by the color of the flowers and the capsule circumscissile below the middle. Portulaca insignis Steyerm., Ann. Missouri Bot. Gard. 75: 1058, fig. 1A–E. 1988. Herbaceous annual with elongated tap root; leaves lance-linear, 4.5–6 × 0.5–1 mm; sepals 4–5 mm long; stamens 13–15; capsule circumscissile near the middle. Shrubby savannas and exposed areas of granitic outcrops, 50–300 m; Bolívar (Piedra Elefante between Ciudad Piar and Puerto Ordaz, near Río Aro between Cuidad Bolívar and Maripa, near Upata). Endemic. The entire plant of Portulaca insignis is covered with a dense mass of elongated intertwined trichomes which equal, exceed, surround, or conceal the leaves and flowers, making it the species with the greatest indument covering of any of the Venezuelan taxa. Portulaca mucronata Link, Enum. Hort. Berol. Alt. 2: 2. 1822. —Botón de soldado.

Perennial herb with a thick somewhat hardened taproot; stems erect, 15–25 cm tall, leafless below the inflorescence; flowers 2 or 3 aggregated at the summit, surrounded by 4–8 involucral leaves; stamens numerous; capsule obovoid, circumscissile at the middle; seed lead-black, iridescent. Rocky outcrops in dry forests and streams, 100–300 m; Bolívar (Altiplanicie de Nuria), Amazonas (region of San Carlos de Río Negro). Brazil, Paraguay, Argentina. ◆Fig. 319. Portulaca oleracea L., Sp. Pl. 445. 1753. —Verdolaga. Portulaca marginata Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 72. 1823. Annual herb with prostrate or ascending stems forming mats; stems green or red; leaves alternate; petaloid sepals variable in size; flowers terminal, solitary or few and congested; stamens 6–15; capsule circumscissile at the middle; seeds black, opaque, obtusely verruculose-granulate. Open, disturbed, and cultivated sites, along stream banks, cut-over forests, 50–400 m; Delta Amacuro (Tucupita), Bolívar (region of El Dorado, Hato La Vergareña, Río Paramichi), Amazonas (region of Puerto Ayacucho). Cosmopolitan weed, native in the Old World and perhaps in the New World as well. ◆Fig. 321. Leaves of Portulaca oleracea are sometimes eaten in salads or used as a green vegetable, similar to spinach. It is also reputed to serve as a vermifuge. Numerous varieties have been described for this highly variable species. Portulaca pilosa L., Sp. Pl. 445. 1753. Annual herb with simple or branched roots; stem usually prostrate and much branched; trichomes numerous, usually shorter than the leaves; flowers surrounded by a dense hairy tuft; petaloid sepals 4–8 mm long; stamens 15–25; capsule obovoid or subglobose. Sandy soils along river banks, rocky igneous outcrops along streams, open disturbed sites, 50–300 m; Delta Amacuro (near Río Grande, Tucupita), Bolívar (Río Botanamo), Amazonas (near Puerto Ayacucho, Río Atabapo, Río Orinoco). Aragua, Anzoátegui, Carabobo, Zulia; southern U.S.A., Mexico, Central America, West Indies, South America.

380

P ORTULACACEAE

Fig. 319. Portulaca mucronata

Fig. 320. Portulaca pusilla

Fig. 321. Portulaca oleracea

Portulaca pusilla Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 71. 1823. Diminutive herbaceous perennial 2–4 cm tall; stems prostrate or procumbent; leaves elliptic, rounded; flowers solitary at the end of the short stem, the surrounding involucral leaves inconspicuous; capsule pedicellate, 1– 2.5 × 1.5–1.6 mm; seeds black or steel gray, fuscous when immature, tuberculate with rounded, non-protruding projections. Exposed granitic outcrops and small sandy depression of igneous rocks, 50–400 m; Delta Amacuro (Tucupita), Bolívar (Cerro Platanal, near Ciudar Piar and Puerto Ordaz, Hato La Vergareña), Amazonas (near Puerto Ayacucho). Colombia (Guaviare). ◆Fig. 320.

Portulaca pusilla is a dwarf species found associated with the endemic taxon P. pygmaea, and the more common, taller, and more branched P. pilosa. Portulaca pygmaea Steyerm., Ann. Missouri Bot. Gard. 75: 1058, fig. 2A–I. 1988. Dwarf herbaceous perennial; leaves suborbicular, suborbicular-obovate, oval, or obovate, rounded at apex; petaloid sepals 5 mm long; stamens 5; capsule pedicellate, 1.5–1.8 × 1.7–2.3 mm; seeds steel gray or grayish, with flattened, non-projecting contiguous, broader than long tubercles. Moist depressions and sandy pockets on exposed igneous outcrops, sometimes near gallery forests,

Talinum 381

100–200 m; Amazonas (north of El Burro, near Puerto Ayacucho). Endemic. Portulaca pygmaea, together with P. sedifolia and P. pusilla, are the smallest of the Guayana Portulaca and have the shortest leaves and flowers. Portulaca sedifolia N.E. Br., Trans. Linn. Soc. London, Bot., ser. 2, 6: 19. 1901. Herbaceous annual growing in rosettes or tufts with spreading often red-purple stems, shortly pubescent; leaves linear-oblong or oblong-lanceolate; calyx often wine red; petaloid sepals white, rose, or pale lavender, 3– 3.5 mm long; capsule circumscissile below the middle; mature seeds black, bronzeblack, steel gray, or silvery gray, varying from mainly uniform, large, rounded, to flattened and scarcely projecting. Exposed igneous outcrops often near streams, sandy soils of savannas subject to flooding, open disturbed sites, 50–300 m; Delta Amacuro (Tucupita), Bolívar (Altiplanicie de Nuria, Río Aro, Río Parguaza), Amazonas (near Puerto Ayacucho, Río Atabapo, confluence of Río Ventuari and Río Orinoco, Samariapo). Sucre; Colombia, Guyana, Suriname, French Guiana, adjacent Brazil. Portulaca teretifolia Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 73. 1823. Portulaca cinerea Poelln., Repert. Spec. Nov. Regni Veg. 33: 158. 1933.

Herbaceous annual 7.5–12 cm tall, the spreading or prostrate stems shortly pilose; leaf whorl at summit of stem 5–13 mm long, the cauline leaves 4–5 mm long; petaloid sepals purplish or pink; stamens numerous; capsule somewhat conic, circumscissile at or slightly above the middle. Dry savannas, igneous outcrops along streams, open sandy soils, 50–900 m; Delta Amacuro (between Piacoa and Los Castillos de Guayana), Bolívar (Río Cuchivero, along Río Orinoco at Ciudad Bolívar and Altagracia, between Upata and Río Caroni, Brazilian border near Santa Elena), Amazonas (Maypures near Puerto Ayacucho, above confluence of Río Orinoco and Río Ventuari, San Carlos de Río Negro). West Indies, adjacent Colombia, Brazil. Portulaca umbraticola Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 72. 1823. Perennial herbaceous plant with prostrate, unbranched, or branched stems 10–13 cm long; leaves 15–17 × 2–3 mm; petiole 1– 1.5 mm long; apical leaves 8–12, 8–18 mm long; petaloid sepals and filaments to 9 mm long; capsule conic, 4–5 mm long, circumscissile at the middle. Wet savannas around igneous outcrops, ca. 100 m; Amazonas (near Puerto Ayacucho). Sucre; southern U.S.A., Mexico, Central America, West Indies, Guyana, Suriname, French Guiana, Brazil, Paraguay, Argentina.

2. TALINUM Adans., Fam. Pl. 2: 245. 1763. Annual or perennial herbaceous plants or low shrubs, the stems very short or elongate. Leaves fleshy, alternate, or subopposite, flat or terete, entire. Flowers small or conspicuous, in long- or short-peduncled cymes, paniculate, sometimes solitary in the leaf axils. Sepals 2, deciduous; petals 5 or more, soon withering. Stamens few to numerous; filaments filiform. Ovary superior, the 3 styles more or less united; ovules numerous. Capsule 1-locular, 3-valved. Seeds compressed, roundreniform; embryo incompletely annular. U.S.A., Mexico, Central America, West Indies, South America, a few species in Africa and Asia; ca. 40 species, 2 in Venezuela, both in the flora area. Key to the Species of Talinum 1.

Inflorescence cymose or racemose with only 2–4(5) principal axes with a total of 2–20(–25) flowers, the inflorescence mainly broader than long; stem perennial, mainly 15–45(–60) cm tall; leaves chiefly < 2 cm or less wide; petals 7–10 mm long, stamens ca. 30........................... T. fruticosum

382

1.

P ORTULACACEAE

Inflorescence an elongate panicle with 7–25 principal axes with a total of usually 20 to over 100 flowers, the inflorescence much longer than broad; stem annual, mainly (40–)50–100 cm tall; leaves chiefly 2–4 cm wide; petals 3.5–5 mm long; stamens 15–20 ..................... T. paniculatum

Talinum fruticosum (L.) Juss., Gen. Pl. 312. 1789. —Portulaca fruticosa L., Syst. Nat. ed. 10, 2: 1045. 1759.—Verdolaga de cabra. Portulaca triangularis Jacq., Enum. Syst. Pl. 22. 1760. —Talinum triangulare (Jacq.) Willd., Sp. Pl. ed. 4, 2: 862. 1799. Stem erect from a thickened, stout, fleshy root; leaves narrowly oblanceolate to obovate, rounded to obtusely acute at apex, attenuate to a subpetiolate base; petals rose, purple, yellow, or white; capsule 4.5–6 mm diameter; seeds black. Open dry ground and waste places, ca. 50 m; Bolívar (Ciudad

Bolívar, San Félix). Common in coastal and northern Venezuela; U.S.A. (Florida), Mexico, Central America, West Indies, South America, naturalized in Africa. ◆Fig. 322. Talinum fruticosum is sometimes eaten as a pot herb similar to spinach.

Fig. 323. Talinum paniculatum

Fig. 322. Talinum fruticosum

Anagallis 383

Talinum paniculatum (Jacq.) Gaertn., Fruct. Sem. Pl. 2: 219, pl. 128, fig. 13. 1791. —Portulaca paniculata Jacq., Enum. Syst. Pl. 22. 1760. —Verdolaga de cabra. Stem erect from fleshy or tuberous roots, simple or branched, glabrous throughout; leaves obovate or elliptic-obovate, rounded at summit, attenuate to a subsessile base; pet-

als pink, yellow, or white; capsule 3–4.5 mm diameter; seeds black. On rocks in deciduous forests, bases of rocky escarpments in montane forests, 300–800 m; Bolívar (Altiplanicie de Nuria, Cerro Altamira, Cerro Uroi, Río Paragua). Common elsewhere in Venezuela; southern U.S.A., Mexico, Central America, South America. ◆Fig. 323.

PRIMULACEAE by James S. Miller and Paul E. Berry Perennial or annual herbs, rarely subshrubs. Secretory cells or glandular trichomes common. Leaves opposite or whorled, basal, sometimes alternate, usually simple and entire, less often lobed or toothed, rarely very finely pinnately divided (Hottonia), without stipules. Inflorescences paniculate, umbellate, racemose, capitate, or the flowers solitary, bracteate. Flowers bisexual, often distylous, actinomorphic or rarely somewhat zygomorphic (Coris), hypogynous or rarely half-epigynous (Samolus). Calyx synsepalous, usually 5-lobed, persistent. Corolla sympetalous, rarely with the petals free (Pelletiera) or absent (Glaux), the lobes usually 5, imbricate. Stamens the same number as and opposite the corolla lobes, rarely with alternating staminodia; filaments adnate to the corolla tube; anthers bithecal, dehiscing longitudinally or rarely by terminal pores. Ovary superior or rarely half-inferior (Samolus), usually 5-carpellate, unilocular, sometimes with 5 basal partitions, the placentation free central; style terminal, simple, the stigma capitate; ovules usually numerous, hemitropous or anatropous. Fruits capsular, usually 5-valved. Seeds numerous, rarely few, endosperm abundant. Subcosmopolitan, mostly Northern Hemisphere; ca. 22 genera and 1000 species, 1 species in the flora area. There is still some uncertainty whether Samolus belongs in Primulaceae or Theophrastaceae, and recent molecular studies indicate that Myrsinaceae may be phylogenetically embedded within the Primulaceae (L. Martins, C. Oberprieler, and F.H. Hellwig. A phylogenetic analysis of Primulaceae s.l. based on internal transcribed spacer (ITS) DNA sequence data. Plant Syst. Evol. 237: 75–85. 2003). 1. ANAGALLIS L., Sp. Pl. 148. 1753. Annual herbs, the stems erect to prostrate, sometimes angled or winged. Leaves opposite, alternate, or whorled, simple, entire, sessile or short petiolate. Inflorescences terminal, racemose, or the flowers solitary and axillary. Flowers bisexual, actinomorphic, 5- or rarely 4-merous, sessile or pedicellate. Calyx deeply lobed or split nearly to the base. Corolla campanulate to rotate, blue, scarlet, or white, delicate, parted almost to the base, the lobes contorted in bud. Ovary superior, globose. Capsule circumscissile; seeds ± trigonous, papillose.

384

P RIMULACEAE

Canada, U.S.A., Mexico, Central America, West Indies, South America, Europe, Africa, Madagascar, southern Asia, Australia; 15–20 species, 2 or 3 in Venezuela, 1 of these in the flora area. Centunculus is sometimes included in Anagallis due to its similarly circumscissile fruit, but molecular data (cited above) indicate that they are distinct genera. Anagallis pumila Sw., Prodr. 1: 40. 1788. —Micropyxis pumila (Sw.) Duby in A. DC., Prodr. 8: 82. 1844. —Centunculus pumilus (Sw.) Kuntze, Revis. Gen. Pl. 3: 193. 1891. Centunculus pentandrus R. Br., Prodr. 427. 1810. Decumbent or scandent herb with stems to 30(–50) cm long; leaves sparse, alternate, sessile, blade 5–8 × 1–3.5 mm; flowers solitary from leaf axils, calyx green, corolla white, to 5 mm diameter; pedicels 6–10 mm long. Swampy savannas, 100–900 m; Bolívar (Altiplanicie de Nuria, Gran Sabana), Amazonas (lower and middle Río Parucito). Portuguesa; U.S.A. (Florida), Mexico, Central America, West Indies, Colombia, Venezuela, Ecuador, Bolivia, Paraguay, Argentina, Africa, southern Asia, Australia. ◆Fig. 324. Numerous varietal names have been proposed for this species, but we follow Ståhl (B. Ståhl. 150. Primulaceae. Flora of Ecuador 39: 32. 1990) in not recognizing them.

Fig. 324. Anagallis pumila

PROTEACEAE by Julian A. Steyermark Evergreen trees, shrubs, or very rarely herbaceous plants without stellate trichomes or peltate scales, often acumulating aluminum. Leaves alternate, rarely opposite or verticillate, exstipulate, simple or compound, entire or toothed; sometimes heteromorphic. Inflorescence spicate, racemose, capitate, umbellate, paniculate, pseudoracemose (raceme of flower pairs) in the flora area, terminal or axillary, bracteate with persistent or caducous bracts subtending one or more flowers. Flowers bisexual or, by abortion, unisexual (plant polygamous or dioecious). Perianth apetalous, actinomorphic or zygomorphic, 4-merous, hypogynous or rarely slightly perigynous; sepals usually petaloid, valvate, free or united below to form a tube, often separating in anthesis and recurving; corolla represented apparently by the annular or horseshoe-shaped, frequently 4-lobed, hypogynous disk or by 2–4 separate or variously connate hypogynous scales or glands alternating with the sepals or completely absent. Stamens 4, opposite the sepals, and attached to their base, shorter than the sepals, or rarely free; anthers longitudinally dehiscing, 2-locular or one of them abortive, the connective continuous with the filament often prolonged as an

Euplassa 385

appendage. Pollen (2)3(–8)-porate, or sometimes -colpoidate. Pistil 1; style terminal, usually somewhat thickened toward the apex; stigma terminal or lateral (in the flora area small and situated on a pollen-presenter, a usually swollen, modified style tip); ovary superior, 1-locular, 1-carpellate, sessile or stipitate, often oblique; ovules usually 1 or 2, less often numerous and biseriate, marginal, usually pendulous. Fruit sometimes an indehiscent or tardily dehiscent nut, achene, or drupe or a dehiscent follicle. Seeds 1(2–20), the testa membranous or coriaceous, or rarely thick and horny, often winged; endosperm scanty or none. Neotropics, Africa, Australia; ca. 75 genera and 1400 species, 3 genera and 16 species in the flora area. Key to the Genera of Proteaceae 1. 1. 2(1).

2.

Adult leaves even-pinnate; perianth in bud incurved at the apex; hypogynous scales connate into an entire or lobed disk ................. 1. Euplassa Adult leaves simple, juvenile leaves often pinnate; perianth straight in bud; hypogynous scales separate or united .......................................... 2 Fruit dehiscent, 2-valved with 2 winged seeds; hypogynous disk of 4 separate scales; leaves sometimes dimorphic with the juvenile ones often pinnate, if reticulate, usually coarsely so without elevated veinlets, or rarely finely reticulate in R. suaveolens ................................... 3. Roupala Fruit indehiscent or tardily dehiscent with a thick, fleshy, or hard pericarp, the single seed unwinged; disk cupuliform, united, 4-lobed; leaves simple, usually with a fine elevated reticulum on one or both surfaces, or coarsely areolate in P. sessilifolia ........................ 2. Panopsis

1. EUPLASSA Salisb. in Knight, Cult. Prot. 101. 1809. Adenostephanus Klotsch, Linnaea 15: 51. 1841. Trees or shrubs. Leaves alternate, pinnately compound with (2)3–6 pairs of entire or dentate leaflets. Inflorescence many-flowered, consisting of solitary or paired pseudoracemes, axillary or rarely terminal. Flowers zygomorphic, in pairs, obscurely 1-bracteate. Calyx somewhat irregular, somewhat oblique in bud, subclavate-cylindric, curved, the sepals eventually free, revolute, deciduous, bearing anthers at the concave apex. Anthers oval, subsessile, the connective shortly protruding at the apex. Hypogynous glands 4, connate or nearly free in a persistent, entire or lobed annulus. Ovary glabrous or hairy, stipitate; ovules 2, collateral, pendulous; style curving, persistent; pollen-presenter lateral, compressed, convex, bearing a small nipple-shaped projection. Fruit indehiscent. Seeds unwinged. South America, principally Brazil; ca. 30 species, 2 in Venezuela, both in the flora area. Key to the Species of Euplassa 1.

1.

Rachis of mature leaves glabrous; lower surface of mature leaflets completely glabrous; petiolules glabrous, pedicels 4.5–6 mm long; hypogynous disk subentire or undulate on one side .................... E. chimantensis Rachis of mature leaves with brown appressed pubescence; lower surface of mature leaflets minutely appressed-pilosulous; petiolules with brown appressed pubescence; pedicels 1.5–3 mm long; hypogynous disk lobed or undulate-margined ........................................ E. venezuelana

386

P ROTEACEAE

Fig. 325. Euplassa venezuelana

Euplassa chimantensis Steyerm., Bol. Soc. Venez. Ci. Nat. 25: 77, fig. 1. 1963. Tree to 3 m tall; leaves with 2 or 3 pairs of subopposite, coriaceous leaflets; leaflets broadly oblong or obovate, broadly rounded, or emarginate at apex, 6–9.5 × 3.5–6 cm; peduncle ferruginous; flower buds yellowish; ovary ferruginous-hirtellous. Forested tepui slopes, 1900–2100 m; Bolívar (Macizo del Chimantá [Toronó-tepui]). Endemic. Euplassa venezuelana Steyerm., Fieldiana, Bot. 28: 217. 1951. —Wakinapaye (Arekuna).

Shrub or tree 2–12 m tall; leaves with 2–5 pairs of subopposite or opposite coriaceous leaflets; rachis ferruginous or red-brown; leaflets oblong, ovate-oblong, or obovate, rounded to subacute, often mucronulate at apex, 7–13 × 3.5–7 cm; flowers very fragrant, creamy, yellow, or tinged reddish, the outer surface of buds buff-brownish or olive green; style greenish white; stigma orange-yellow. Gallery forests bordering savannas, 400– 1300 m; Bolívar (Gran Sabana, basins of Río Aponguao, Río Caruay, and Río Kukenán). Endemic. ◆Fig. 325.

2. PANOPSIS Salisb. in Knight, Cult. Prot. 104. 1809. Andriapetalum Pohl, Pl. Bras. Icon. Descr. 113. 1827, spelling variant: Andripetalum. Trees or shrubs. Leaves alternate, opposite, or verticillate, simple, entire, usually with conspicuous reticulate venation. Inflorescence simply pseudoracemose or rarely branched, axillary or terminal, solitary or fasciculate, many-flowered; flowers actinomorphic, in pairs, 1-bracteate. Sepals 4, free, linear, outer surface pubescent, inner surface glabrous, revolute, deciduous. Stamens inserted at the base or middle of the sepals; anthers oval or oblong, borne on partially free filaments, the

Panopsis 387

connective shortly protruding at the apex. Hypogynous scales united into a cupuliform or urceolate, 4-dentate or lobed disk. Ovary pubescent, sessile; ovules 2, collateral; style filiform, deciduous. Fruit indehiscent to tardily dehiscent, hard, 1seeded, globose or ellipsoid, with a thick woody exocarp. Seed unwinged. Central America, South America; ca. 20 species, 10 in Venezuela, 6 of these in the flora area. Key to the Species of Panopsis 1.

1.

2(1). 2. 3(2). 3. 4(3).

4.

5(3).

5.

Leaves sessile or subsessile; reticulation coarsely areolate with areoles 1–5 mm diameter; leaves crowded on the stem, pseudoverticillate; fruit subglobose, 3–5 cm diameter ......................................... P. sessilifolia Leaves petiolate; reticulation generally of more minute areoles, the areoles 0.5–1 mm diameter; leaves scattered on the stem, mainly alternate, opposite, or subopposite; fruit fusiform or longer than broad, or the fruit < 3 cm diameter ...................................................................... 2 Stems, leaf blades, and petioles glabrous or essentially so ......... P. ptariana Young stems, portions of the leaf blades, and petioles pubescent ........... 3 Trees 4–22 m tall; leaves mainly 10–21 cm long ...................................... 4 Small shrubs 1–1.5 m tall; leaves 2–9 cm long ......................................... 5 Inflorescence 15–22 cm long; pedicels filiform, elongated, 7–13 mm long; lower surface of leaves with pale sericeous pubescence or subglabrescent; plants of low elevations at 50–500 m ....................... P. rubescens Inflorescence 5–8 cm long; pedicels 2 mm long; lower surface of leaves rufous-tomentose; plants of high elevations at 1500–1600 m .................................................................................................... P. tepuiana Pedicels to 2 mm long; style glabrous throughout; reticulation of lower surface of leaves with larger areoles than that on upper surface, but manifest and impressed on both sides; leaves 4.5–9 cm long; plants of the Sierra Parima, Amazonas ............................................... P. parimensis Pedicels 2.5–7 mm long; style strigillose below the middle and toward the base; reticulation of lower surface of leaves subelevated, scarcely evident or obscure on upper surface; leaves 2–7 cm long; plants of sandstone table mountains of eastern Bolívar ........................... P. ornatinervia

Panopsis ornatinervia Steyerm., Bol. Soc. Venez. Ci. Nat. 25: 79, fig. 2. 1963. Shrub 1–1.5 m tall with decumbent ferruginous-pubescent stems; leaves alternate or subopposite, small, ferruginous brown on lower surface, obovate-oblong or oblong, broadly rounded at summit, 2–4 × 1.5–2.5 cm; calyx ferruginous without, pale green within; style pale green. Open dwarf forests on sandstone and diabase, 2000–2200 m; Bolívar (Macizo del Chimantá [summit of Toronó-tepui]). Endemic. Panopsis parimensis Steyerm., Ann. Missouri Bot. Gard. 74: 612. 1987. Shrub to 1.5 m tall; branchlets densely

ferruginous-pubescent; leaves alternate or subopposite, elliptic-obovate or elliptic-oblong, 2–4 cm wide, subacutely obtuse at apex. Fern-covered, previously burnt-over, and disturbed slopes, ca. 1100 m; Amazonas (Sierra Parima). Endemic. Panopsis ptariana Steyerm., Fieldiana, Bot. 28: 218. 1951. Shrub 0.5–1.8 m tall; stems, petioles, and leaf blades glabrous; leaves opposite, obovate-oblong, 3.5–6 × 1.7–4 cm; pedicels, calyx, and ovary ferruginous; style and stigma yellow. Dry sandy and rocky sandstone exposures adjacent to swampy places on slopes of table mountains and shrubby savannas near

388

P ROTEACEAE

Fig. 326. Panopsis rubescens

Fig. 327. Panopsis sessilifolia

Roupala 389

streams, 1400–1600 m; Bolívar (Gran Sabana, Ptari-tepui, near Salto Torón in basin of Río Aponguao). Endemic. Panopsis rubescens (Pohl) Pittier, Bol. Comerc. Industr. Venez. 13: 417. 1921; reprinted as Arb. Arbust. Venez. 1: 21. 1923. —Andriapetalum rubescens Pohl, Pl. Bras. Icon. Descr. 1: 114, t. 91. 1927 [1928]. —Camilla de locho, Chaparro de agua, Kawadi-jodedü (Yekwana). Panopsis rubescens var. simulans J.F. Macbr., Field Mus. Nat. Hist., Bot. Ser. 11: 67. 1931. Panopsis cuaensis Steyerm., Ann. Missouri Bot. Gard. 74: 612. 1987. Tree (3–)7–15 m tall; young branchlets, leaves, and inflorescence ferruginous-pubescent; leaves alternate, opposite, or subopposite, oblanceolate-oblong or broadly oblanceolate, obtuse to acute at apex, (7–)10–15 cm long, mainly (1.5–)2.5–7 cm wide; flower buds silvery green; flowers fragrant, buff-yellow; fruit fusiform, elongated, densely brown-pubescent, at maturity 4.5–5 × 1.5–3 cm. Gallery and flooded riparian forests, wet open areas, Mauritia palm swamps, 50–500 m; Bolívar (Maripa, tributaries of Río Aponguao, Río Caroni, Río Carrao, Río Caura between Entrerios and El Paují, middle Río Orinoco, Río Paragua), Amazonas (Río Atabapo, Río Casiquiare, Río Cuao, Río Guainía, upper Río Orinoco, Samariapo). Anzoátegui, Apure; Colombia, Guyana, Suriname, French Guiana, Peru, Brazil. ◆Fig. 326. Panopsis sessilifolia Rich., Mém. Soc. Hist. Nat. Paris 1: 106. 1792. —Andria-

petalum sessilifolium (Rich.) Klotsch, Linnaea 15: 43. 1841, “Andripetalum.” Tree (6–)10–12 m tall, only sparsely pubescent on branches and leaves; leaves pseudoverticillate, broadly elliptic or oblongobovate, acute to acuminate at apex, obtuse to acute at base, 10–26 × 5–8 cm; inflorescence terminal, simple or with 1 or 2 slender elongated axes, relatively sparsely cinerouspubescent; fruit subglobose, 4–5 cm diameter. Riparian forests and gallery forests in savannas, 400–1200 m; Bolívar (Gran Sabana along Río Yuruani, affluents of Río Aponguao, Río Caruay, and Río Kukenán, [along Río Apacará at base of Apacarátepui]). Guyana, Suriname, French Guiana, Brazil. ◆Fig. 327. Vegetatively, Panopsis sessilifolia is similar to Roupala suaveolens, but differs in the more conspicuously elevated reticulate venation of the upper surface of leaves. Panopsis tepuiana Steyerm., Fieldiana, Bot. 28: 219. 1951. —Para-ma-yek (Arekuna). Tree 20–22 m tall, the branches, young leaves, petioles, and inflorescence densely ferruginous-pubescent; leaves alternate or opposite, oblanceolate or elliptic-oblanceolate, subobtusely apiculate at apex, 8.5–15 × 2–5 cm; rachis, pedicels, and sepals orangeferruginous; sepals creamy yellow within; style and stigma pale green. Tepui slope forests, 1500–1600 m; Bolívar (Ptari-tepui). Endemic. The fruit of Panopsis tepuiana is reputed to be edible.

3. ROUPALA Aubl., Hist. Pl. Guiane 83, t. 32. 1775, spelling variant: Rhopala. Shrubs or trees. Leaves alternate, heteromorphic, the adult leaves usually simple, entire or dentate, those of sterile branches or of young plants often incised or odd-pinnate. Inflorescence axillary or terminal, pseudoracemose or pseudospicate. Flowers actinomorphic, in pairs, often divaricate, subtended by a common bract, shortly pedicellate, bracteate, the bracts persistent or caducous. Calyx subclavate-cylindric, straight, the sepals linear, free, recurved, with stamens attached above the middle. Stamens exserted; free part of filaments very short; anthers linear or oblong, the connective shortly protruding at the apex. Hypogynous scales 4, distinct, obtuse or acute. Ovary sessile; ovules 2, collateral, pendulous from the apex of the locule; style filiform, persistent; pollen-presenter terminal, clavate, obtuse; stigma minute, terminal. Fruit a hard, somewhat ligneous, oblique follicle, shortly stipitate and bearing 2 winged, compressed seeds. Neotropics; ca. 55 species, 12 in Venezuela, 8 in the flora area.

390

P ROTEACEAE

Key to the Species of Roupala 1. 1. 2(1). 2. 3(2). 3.

4(3). 4. 5(3).

5.

6(2).

6.

7(6).

7.

Dwarf shrubs 0.8–1.5 m tall; leaves 1.3–3 × 0.7–2.5 cm; petioles 1–2 mm long; ovary glabrous ................................................................... R. minima Shrubs or trees 3–20 m tall, leaves 3.5–17 × (2–)2.5–10 cm; petioles 10– 50 mm long; ovary pubescent ................................................................ 2 Leaves rounded, obtuse, or obtusely acute at apex .................................. 3 Leaves acute to acuminate at apex ........................................................... 6 Pedicels glabrous; rachis of inflorescence glabrous; lower surface of leaves glabrous ...................................................................................... 4 Pedicels tomentose or furfuraceous-puberulent; rachis minutely ferruginous-puberulent or furfuraceous-castaneous-puberulent; leaves minutely puberulent or furfuraceous ........................................................ 5 Petioles 3–5 mm long; leaf blades 3.5–7.5 cm long, obtuse at base .................................................................................................. R. paruensis Petioles 12–20(30) mm long; leaf blades (5–)7–11(–17) cm long, acute at base ............................................................................................ R. obtusata Mature perianth 7–10 mm long; ovary hirsutulous; leaf blades elliptic or ovate oblong, obtuse to subacute at the base, 2.5–7 cm wide ............................................................................................... R. sororopana Mature perianth (10–)12–13 mm long; ovary shortly appressed-pubescent; leaf blades ovate to suborbicular-ovate, truncate or broadly rounded at base, 7.5–10 cm wide ...................................... R. chimantensis Stems, petioles, and lower surface of leaf blades glabrous or glabrescent; perianth strigulose to glabrescent toward apex; fruit (dried state) minutely pale strigulose to glabrescent ................................... R. montana Stems, petioles, and lower surface of leaf blades manifestly pubescent; perianth densely tomentulose to villous; fruit (dried state) densely ferruginous-tomentose ........................................................................... 7 Lower surface of leaf blades with fine, minute, elevated tertiary reticulation; main lateral veins 7–15 each side; fruit body obliquely ovate, elliptic to lance-oblong, 12–25 × 7–14 mm, 1.2–2 times longer than broad; stipe of fruit 4–5 mm long .......................................... R. suaveolens Lower surface of leaf blades ± plane, the tertiary venation obsolete or at least not forming an elevated network; main lateral veins 5–7 each side; fruit body obliquely ovate, 15–19 × 13–14 mm, 1.3–1.4 times longer than broad; stipe of fruit 2 mm long ................................. R. griotii

Roupala chimantensis Steyerm., Bol. Soc. Venez. Ci. Nat. 25: 81, fig. 3. 1963. Tree 3–4 m tall; leaves stiff-coriaceous; flowers cream-colored. Dwarf tepui forests and scrub, 1900–2000 m; Bolívar (summit of Chimantá-tepui). Endemic. ◆Fig. 328. Roupala chimantensis is very close to R. sororopana. Additional collections are needed to determine if it is distinct from the earlier published R. sororopana. Roupala griotii Steyerm., Phytologia 44: 321. 1979.

Tree to 3 m tall; leaves broadly ovate or broadly elliptic, 8–15 × 3–8.5 cm; rachis of inflorescence, pedicels, and sepals with spreading tawny-brown pubescence; sepals pale brown, 10–13 mm long, pubescent without, glabrous within; style green. Gallery forests bordering tree savannas with Platycarpum orinocense, ca. 200 m; Amazonas (Río Coro Coro at base of Cerro Yutajé). Endemic. Roupala minima Steyerm., Fieldiana, Bot. 28: 220. 1951. Spindly ligneous plant or shrub 0.8–3 m

Roupala 391

tall; branchlets slender, wand-like; leaves small, elliptic-oblong or suborbicular, subsessile, glabrous; inflorescence glabrous; sepals 2–3 mm long; follicles obliquely oblongelliptic, 2–2.8 × 0.8–1 cm. Savannas with shrubs on white sand, sandy valleys following streams in savannas and sandy plateaus, 900–1600 m; Bolívar (Gran Sabana in area between Ptari-tepui and Guyana border). Endemic. ◆Fig. 331. Roupala montana Aubl., Hist. Pl. Guiane 83, t. 32. 1775. —Carne asada, Horca, Horca mandingo, Mandingo.

Roupala dentata R. Br., Trans. Linn. Soc. London 10: 192. 1810. —Roupala montana var. dentata (R. Br.) Sleumer, Bot. Jahrb. Syst. 76: 173. 1954. Roupala complicata Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 153, t. 119. 1817. —Roupala montana ß complicata (Kunth) Griseb., Fl. Brit. W. I. 277. 1864. Roupala macropoda Klotzsch & H. Karst., Linnaea 20: 473. 1847. Roupala dissimilis Pittier, Bol. Soc. Venez. Ci. Nat. 5: 303. 1939. Shrub or tree to 15 m tall; leaves often heteromorphic, the adult leaves simple,

Fig. 328. Roupala chimantensis

Fig. 329. Roupala obtusata

392

P ROTEACEAE

Fig. 330. Roupala sororopana

Fig. 331. Roupala minima

Fig. 332. Roupala suaveolens

Q U I I N A C E A E 393

ovate, 4–14 × 2–9 cm, the juvenile leaves oddpinnately compound, margins entire to coarsely serrate; flowers white, cream, or greenish white, fragrant, with sepals 7–8.5 mm long; follicles 2.5–4 cm long. Dry open or shrubby savannas, often with Curatella, Bowdichia, and Byrsonima crassa, borders of granitic outcrops, gallery forests, 50–1300 m; Delta Amacuro (eastern Los Castillos), Bolívar (widespread except the eastern part), Amazonas (Auyán-tepui, Cerro Aratitiyope, Cerro Duida, Cerro Mahedi, Cerro Parú, Puerto Ayacucho, Río Ventuari, Roraimatepui, Sierra de la Neblina, Sierra Parima). Common in Coastal Cordillera, Llanos, and base of the Andes; widely distributed in Neotropics. When cut, the young branches of Roupala montana have an odor similar to canned corned beef. The wood is hard and the reddish color resembles that of smoked beef. Roupala obtusata Klotzsch, Linnaea 15: 54. 1841. Shrub or tree to 7 m tall; leaves oblong, narrowly obovate-oblong, oblong-elliptic, or lance-elliptic, glabrous; sepals 10–11 mm long, white, greenish white, or cream-colored; follicles gray-green, 3–4 cm long, glabrate. Riparian forests mostly along black-water rivers, 100–200 m; Amazonas (Caño San Miguel, base of Cerro Sipapo, Río Atabapo, Río Guainía, Río Sipapo, Río Ventuari). Colombia (Guainía: Río Guainía), Guyana, French Guiana. ◆Fig. 329. Roupala paruensis Steyerm., Ann. Missouri Bot. Gard. 74: 613. 1987. Tree to 7 m tall; leaves and inflorescence glabrous; leaves ovate-oblong; follicles ob-

liquely obovoid, 1.5–2.5 cm long, glabrous. Rocky savannas on tepui summits, ca. 2000 m; Amazonas (Cerro Parú). Endemic. Roupala sororopana Steyerm., Fieldiana, Bot. 28: 220. 1951. Shrub 3–4 m tall; leaves stiff-coriaceous, the lower surface dull green and ferruginous; flowers with dark maroon-furfuraceous indument; sepals greenish without, yellowish white within. Rocky open lateritic substrate, scrub forests on diabase on tepui summits, 2100–2300 m; Bolívar (Macizo del Chimantá [Toronó-tepui], Serra do Sol on the border with Brazil, Sororopán-tepui). Brazil (Roraima). ◆Fig. 330. Roupala sororopana and R. chimantensis are very close. More intensive collecting is needed before definite conclusions can be reached as to their validity. Roupala suaveolens Klotzsch, Linnaea 20: 473. 1847. Roupala schomburgkii Klotzsch, Linnaea 20: 474. 1847. Shrub or tree 2.5–10(–20) tall; leaves heteromorphic, both simple and compound and entire to coarsely dentate, (5–)7–15 × (3.5–) 4.5–7.5 cm; lower leaf surface finely reticulate; flowers fragrant; sepals pale ferruginous or yellow, 10–11 mm long. Gallery forests, river margins, around rapids, flooded savannas, tepui slopes, 100–1200 m; Bolívar (Cerro Arepuchi in upper Río Caroní basin opposite mouth of Río Icabarú, Río Apacara, Río Aponguao, Río Caroni, Río Caura, Roraima-tepui, Uaipán-tepui), Amazonas (Cerro Yutajé, Río Cunucunuma). Guyana, northeastern Amazonian Brazil. ◆Fig. 332.

QUIINACEAE by Julio V. Schneider and Georg Zizka Trees or shrubs, rarely vines, to 25(–30) m tall. Stems sometimes unbranched, with adventitious roots. Leaves opposite or whorled, very rarely alternate (juvenile plants), simple or pinnately compound, evergreen, chartaceous to coriaceous, petiolate or ± sessile; margins entire, crenate, or serrate, flat to ± crisp; venation craspedodromous to brochidodromous, tertiary veinlets parallel (plumose-reticulate in

394

Q UIINACEAE

Quiina); stipules interpetiolar (usually not fused in Quiina), foliaceous or setose, entire or divided into several setose parts (Froesia). Inflorescence axillary or terminal, thyrsoid or botryoid; rachis usually shortly pilose, in Quiina also glabrous or tomentose; bracts triangular to ovate. Flowers ± pedicellate, ± actinomorphic, unisexual or bisexual (plants hermaphroditic, androdioecious, or dioecious); receptacle ± inconspicuous. Sepals (3)4 or 5(6), imbricate, ± ciliate, highly variable in shape; petals (3)4–8. Stamens 12–many; filaments filiform; anthers subglobose to oblong in outline, longitudinally dehiscent. Gynoecium apocarpous or syncarpous, superior; if apocarpous comprising 3 carpels; if syncarpous then ovary 2–14-locular, ± globose to pyriform; ovules (1?)2–4 per locule, placentation basal-axile(?); styles terminal; stigma ± peltate. Fruit berry-like, globose, ovoid to pyriform, elliptic, or of 1–3 follicles (Froesia); pericarp hard or fleshy, fibrous. Seeds 1 to more than 20 per fruit, villous or glabrous (Froesia); endosperm present or absent(?); embryo straight, cotyledons thick. Central America (Belize to Panama), West Indies, Colombia, Venezuela, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 4 genera and ca. 55 species, 4 genera and 22 species in the flora area. Key to the Genera of the Quiinaceae 1. 1. 2(1). 2. 3(1).

3.

Leaves of adult plants pinnately compound ............................................. 2 Leaves of adult plants simple .................................................................... 3 Gynoecium apocarpous; seeds glabrous .......................................... 1. Froesia Gynoecium syncarpous; seeds villous ......................................... 4. Touroulia Flowers staminate or bisexual, the plants androdioecious; leaves opposite and decussate, rarely whorled (Q. pteridophylla), with intersecondary veins; ovary 2(3)-locular ............................................................... 3. Quiina Flowers unisexual, the plants dioecious; leaves in whorls of 4, rarely opposite (L. oppositifolia), without intersecondary veins; ovary 4–14locular ...................................................................................... 2. Lacunaria

1. FROESIA Pires, Bol. Técn. Inst. Agron. N. 15: 22. 1948. by Georg Zizka Erect trees to 20 m tall or treelets to 5 m (F. tricarpa) tall, unbranched. Leaves large, to 140 cm long, crowded at stem apex, opposite, sometimes in whorls of three (F. venezuelensis), odd-pinnately compound, 11–27-foliolate, petiolate; stipules interpetiolate, divided into 2–5 setaceous parts, united at base, to 2(–5) cm long; petiole 15–40 cm long; leaflets opposite to alternate, chartaceous to coriaceous, ovate to oblong, 10–45 × 5–14 cm, petiolulate, ± rounded at base, acute; margins entire, crenate or serrate. Inflorescence terminal, thyrsoid, paniculately branched, many-flowered; primary and floral bracts triangular, acute, abaxially pilose. Flowers bisexual, 9–25 mm diameter, pedicellate, the pedicel densely pilose, articulating basally. Sepals (4)5, unequal, coriaceous, brownish, suborbicular, abaxially densely pilose, adaxially inconspicuously pilose, persistent and reflexed in fruit; petals 5, coriaceous, glabrous, obovate, bilobed at apex, the lobes broadly rounded. Stamens many (to > 300). Gynoecium apocarpous, carpels 3, glabrous to densely pilose, each

Froesia 395

Fig. 333. Froesia tricarpa

396

Q UIINACEAE

with 1 or 2 ovules. Fruit composed of 1–3 free follicles (1 or 2 carpels often aborting), ± globose or pyriform, opening irregularly by a longitudinal slit, glabrous or very shortly pilose. Seeds globose, black, glabrous, 1 per follicle; embryo small, with two large cotyledons. Colombia, Venezuela, Peru, Brazil; 5 species, 3 in Venezuela, 2 of these in the flora area. The third species in Venezuela, Froesia venezuelensis Steyerm. & G.S. Bunting, is restricted to cloud forests in Yaracuy state. Key to the Species of Froesia 1.

1.

Margins of leaflets entire at base, crenulate-serrulate near apex, with (27–)40–57 pairs of secondary veins, distance between secondary veins 4–6(–7) mm; leaflets coriaceous; follicles minutely pilose; trees to 12 m tall ........................................................................................... F. gereauana Margins of leaflets conspicuously serrate, with 19–30 pairs of secondary veins, distance between secondary veins 7–10 mm; leaflets chartaceous to coriaceous; follicles glabrous; small trees to 5 m tall ..................................................................................................... F. tricarpa

Froesia gereauana J.V. Schneid. & Zizka, Novon 7: 408. 1997. Tree to 9–12 m tall; petiolules 5–18 mm long. Granitic outcrops, ca. 1500 m; Amazonas (Caño Piedra in Serranía Sipapo). Endemic.

Froesia tricarpa Pires, Bol. Técn. Inst. Agron. N. 15: 22. 1948 Small tree to 5 m tall; petiolules (15–)20– 45 mm long. White-sand savannas, 100–200 m, Amazonas (near Maroa). Colombia, Brazil. ◆Fig. 333.

2. LACUNARIA Ducke, Arch. Jard. Bot. Rio de Janeiro 4: 139. 1925. by Georg Zizka Erect trees or shrubs, to 15(–20) m tall. Leaves verticillate in whorls of 4 (rarely 3), opposite in L. oppositifolia, simple (early leaves of seedlings can be pinnatifid), ± petiolate; blade chartaceous to coriaceous, lanceolate to elliptic, ovate or obovate; margin of blade entire, ± crenate or serrate, often ± crisp; venation craspedodromous, secondary veins conspicuous, ± arching upward near margin, intersecondary veins absent; stipules interpetiolar, setaceous to foliaceous (L. minor), entire, acute, caducous. Inflorescence axillary or in some species terminal, thyrsoid, the pistillate inflorescences often less branched than the staminate; bracts triangular, abaxially shortly pilose; pedicels articulating basally or at about the middle. Flowers pedicellate, unisexual, the plants dioecious; floral bracts triangular, equaling the primary bracts but slightly smaller. Sepals 4, imbricate, ± unequal, ovate to (sub)orbicular, rounded, coriaceous, ciliolate on the margin, minutely pilose abaxially, persisting in fruit. Petals 4–8, often ± unequal, ligulate to obovate, rounded, glabrous, coriaceous, whitish to yellowish. Stamens many in staminate flowers; anthers longitudinally dehiscent. Pistillate flowers with ovary globose to pyriform, longitudinally striate (except in L. oppositifolia), with 4–14 locules and terminal styles. Fruit ± globose, usually notched and with persistent style bases at apex, longitudinally striate; pericarp with latex-containing depressions. Seeds 4 to

Lacunaria 397

more than 20, (1)2–4 per locule, densely villous with brownish hairs; endosperm absent; embryo straight; cotyledons thick. Costa Rica, Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 9–11 species, 4 in Venezuela, all in the flora area. Key to the Species of Lacunaria 1. 1. 2(1).

2.

3(2). 3.

Leaves opposite ........................................................................ L. oppositifolia Leaves whorled, 4 or rarely 3 per node ..................................................... 2 Leaf margin conspicuously serrate-crenate; petals (6–)8; petioles very short, 2–8 mm long; leaf blades coriaceous, (8–)10–19(–25) × (2–)3– 5(–6) cm, with 9–16 pairs of secondary veins; fruits 1.8–3(–6) cm long, 7–9-locular, 1 seed per locule ...................................................... L. crenata Leaf margin entire to inconspicuously crenate, petals 4–6(7); petioles 5– 70 mm long; leaf blade chartaceous to subcoriaceous, 9.5–30(–52) × 3– 12 cm, with 7–25(–29) pairs of secondary veins; fruits (2.5–)3–7 cm long, 9–14-locular, (1)2–4 seeds per locule ........................................... 3 Leaves with 7–12 pairs of secondary veins; blades 9.5–19(–21) × 3– 7.5 cm; stipules 4–6 mm long ........................................... L. macrostachya Leaves with 14–25(–29) pairs of secondary veins; blades 14–30(–52) × 4.2–12 cm; stipules 8–11 mm long ........................................... L. jenmanii

Lacunaria crenata (Tul.) A.C. Sm., Trop. Woods 58: 31. 1939. —Quiina crenata Tul., Ann. Sci. Nat. Bot. sér. 3, 11: 163. 1849. Tree to 15 m tall; leaves coriaceous, the lower surface shining and differing in color from the upper; stipules 8–14 mm long. Evergreen lowland to lower montane forests, expected to occur in the lowlands of Bolívar or Amazonas. Apure; Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil. ◆Fig. 334. This species is one of the most widespread of the genus, and its occurrence in the flora area is probable. Lacunaria jenmanii (Oliv.) Ducke, Arch. Jard. Bot. Rio de Janeiro 5: 171. 1930. —Touroulia jenmanii Oliv. in Hook., Icon. Pl. 20: pl. 1998. 1891. —Molleja de gallineta, “jenmani.” Tree to 18 m tall; leaves chartaceous to subcoriaceous; stipules 8–11 mm long; fruit reportedly edible. Evergreen lowland forests, 100–400 m; Amazonas (La Esmeralda, Río Cuao, San Carlos de Río Negro). Apure; Colombia, Suriname, French Guiana, Peru, Brazil. ◆Fig. 335.

Small-leaved specimens of this widespread and variable species can be difficult to separate from Lacunaria macrostachya. The number of secondary veins and stipule length are the most reliable characters to separate them. Lacunaria macrostachya (Tul.) A.C. Sm., Trop. Woods 58: 31. 1939. —Quiina macrostachya Tul., Ann. Sci. Nat. Bot. sér. 3, 11: 162. 1849. —Coco de mono, Molleja de gallineta banera, Molleja de paují, Oroma-Jijote (Yekwana). Tree to 20 m tall; leaves subcoriaceous; stipules lanceolate, 4–6 mm long; fruit pulp reportedly edible. Evergreen lowland forests, 100–200 m; Bolívar (Aripao, Río Caura), Amazonas (Río Cunucunuma, San Carlos de Río Negro, Tamatama). Peru, Brazil, Bolivia. Lacunaria oppositifolia Pires, Bol. Técn. Inst. Agron. N. 28: 45. 1954. —Coco de mono pequeño, Jakí-ko, Shinatü. Quiina spruceana Engl. in Mart., Fl. Bras. 12(1): 481. 1888. —Lacunaria spruceana (Engl.) Pires, Bol. Técn. Inst. Agron. N. 15: 29. 1948.

398

Q UIINACEAE

Fig. 334. Lacunaria crenata

Fig. 335. Lacunaria jenmanii

Quiina 399

Tree to 15 m tall; leaves chartaceous; stipules narrowly lanceolate, 7–10 mm long; fruits 4- or 5-locular, 1 seed per locule. Evergreen lowland forests, 100–200 m; Amazonas (Culebra, Río Orinoco, Río Padamo, Río Ventuari). Brazil. In this species, the leaves have the charac-

teristic venation of Lacunaria (no intersecondary veins) but are opposite as in Quiina. It also differs from other species in the genus in its fruit morphology. Because of these characters, it has been considered to belong to a new genus, but this has so far not been formally proposed.

3. QUIINA Aubl., Hist. Pl. Guiane 2(suppl.): 19, pl. 379. 1775. by Julio V. Schneider Trees and shrubs, rarely vines, to 25 m tall. Stems sometimes with adventitious roots, terminal internodes ± laterally compressed. Leaves decussate, simple, petiolate or sessile; blades entire or minutely serrate; secondary veins ± arching upward near margin, intersecondary veins present; stipules setaceous or foliaceous, entire. Inflorescence axillary, thyrsoid or botryoid; bracts triangular to ovate; pedicels articulating basally or at about the middle. Flowers unisexual or bisexual, plants androdioecious; floral bracts triangular to ovate, abaxially ± hairy. Sepals 3– 5(6), ciliolate, highly variable in shape; petals 3–7, white or yellow, rarely roseate, mostly obovate, rounded. Stamens 12–21 (bisexual) or 14–62 (staminate). Ovary 2(3)-locular; ovules 2 per locule, basal-axile(?); style terminal; stigma ± peltate. Fruit berry-like, (1)2(3)-locular, pericarp fibrous. Seeds 1 or 2 per fruit, villous; endosperm absent; embryo straight, cotyledons thick. Central America (Belize, Honduras, Panama), West Indies, Colombia, Venezuela, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; ca. 40 species, 14 in Venezuela, all of these in the flora area. In addition to the species listed below, there is another undescribed species from Bolívar and Amazonas states, so far with five specimens from Salto Salas in the upper Río Orinoco (Croizat 536, NY) and Los Pijiguaos (Fernández & Delgado 5813, MO, NY, PORT; Fernández & Sanoja 5892, MO, NY, PORT; Fernández 6789, NY, PORT; Sanoja & Fernández 2904, MO). Key to the Species of Quiina 1. 1. 2(1). 2.

3(2). 3. 4(2).

Leaves in whorls of (3)4 or 5, pinnatifid when juvenile to simple when adult .................................................................................. Q. pteridophylla Leaves opposite and simple ....................................................................... 2 Largest stipules > 4 mm wide, ovate, rarely elliptic; fruits oblong-elliptic to ovoid, never globose ........................................................................... 3 Largest stipules < 4 mm wide (rarely > 4 mm in Q. florida, but then fruits globose), subulate to narrowly lanceolate; fruits oblong-ovoid or globose ......................................................................................................... 4 Sepals and fruit densely velvety-pubescent; fruit ± ovoid; leaves ± obovate; petiole 2.5–9 cm long ................................................... Q. oiapocensis Sepals abaxially ± glabrous; fruit glabrous, oblong to elliptic in outline; leaves ± elliptic; petiole < 2 cm long ....................................... Q. longifolia Stipules adaxially densely brownish-tomentose, narrowly triangular; lower surface of leaves with secondary veins very prominent, rising

400

Q UIINACEAE

4.

5(4). 5. 6(5). 6. 7(6).

7. 8(6).

8.

9(8). 9. 10(9). 10. 11(10). 11. 12(11). 12. 13(11). 13.

ridge-like from blade; fruit oblong-ovoid, (2–)3–4.5 cm long; leaves ± obovate, even when dried mostly deep bluish green .......... Q. indigofera Stipules adaxially glabrous or rarely pilose, not densely brownish-tomentose, setose, lanceolate, triangular, elliptic, or ovate; secondary veins ± prominent, not ridge-like; fruits < 2 cm long (rarely to 2.5 cm long in Q. florida); leaves of various shape, not bluish green .............................. 5 Rachis of inflorescence densely tomentose, > 20-flowered; pedicels ≤ 2(–2.5) mm long ............................................................................... Q. macrophylla Rachis of inflorescence not densely tomentose, if so, < 15-flowered; pedicels > (2–)2.5 mm long .................................................................... 6 Leaves sessile, subcordate or rounded at base ......................................... 7 Leaves petiolate, rarely shortly petiolate, not subcordate or rounded at base ......................................................................................................... 8 Leaf blade chartaceous, acuminate, margin minutely serrulate (use hand lens); petals 6 or 7; inflorescence branched at base or with several axes ................................................................................................ Q. guianensis Leaf blade coriaceous, subacuminate to acute, margin entire; petals 3 or 4; inflorescence not branched at base, rachis 1 per leaf axil ... Q. tinifolia Lateral veins of leaf blades more prominent on upper surface than lower surface; stipules curving upward at base; leaves obovate (rarely elliptic), minutely serrulate (use hand lens); stipules subulate, > 1.5 mm wide ............................................................................................. Q. obovata Lateral veins not more prominent on upper surface than lower surface; stipules ± straight at base; leaves mostly not obovate, with entire margin, when minutely serrulate, then stipules < 1 mm wide .................. 9 Secondary veins (18–)20–30; leaves 15–40 × 5–13 cm, ± elliptic; terminal internodes (densely) pubescent ........................................... Q. cruegeriana Secondary veins < 18; leaf blade smaller than 20 × 6 cm ...................... 10 Leaf margin minutely serrulate (use hand lens); leaves obovate to elliptic, < 10 cm long ....................................................................... Q. attenuata Leaf margin entire; leaves ovate to elliptic (rarely obovate in Q. wurdackii) ............................................................................................ 11 Stipules ≤ 5 mm long, narrowly ovate .................................................... 12 Stipules 5–15 mm long, elliptic, ovate, or subulate. .............................. 13 Leaves 1–3 cm wide; petiole 1–3 mm long ................................. Q. wurdackii Leaves 3.5–6 cm wide; petiole > 8 mm long ................................ Q. yatuensis Fruit ± elliptic, 0.9–1.5 × 0.4–0.7 cm; basal part of pedicel (below articulation) < 1 mm long; stipules 0.3–1.2(–2.5) mm wide ............. Q. rhytidopus Fruit globose, 1.1–2.5 × 0.9–2 cm; basal part of pedicel at least at base of rachis > 1 mm long; stipules 0.8–5.5 mm wide .......................... Q. florida

Quiina attenuata J.V. Schneid & Zizka, Candollea 58: 462. 2003. —Simiuri. Tree to 14 m tall; leaves 2.5–9.5 × 1–3 cm, elliptic to obovate, shortly acuminate, the base ± attenuate; margin minutely serrulate (use hand lens); midvein conspicuously prominent above, secondary veins 7–12; stipules ± subulate; staminate inflorescences with up to 70 flowers per rachis. Seasonally

black-water flooded lowland forests, 100–200 m; Amazonas (Caño San Miguel, San Carlos de Río Negro). Peru. Quiina cruegeriana Griseb., Fl. Brit. W. I.: 106. 1864. —Caspadillo, Murundek (Arekuna). Tree to 15 m tall. Evergreen, moist or seasonally dry, (riparian) lowland and montane

Quiina 401

forests, 300–2000 m; Bolívar (Cerro Venamo, Kavanayén, Mata de Cuchillo 16 km west of Santa Elena de Uairén), Amazonas (Sierra de la Neblina, Sierra Parima). Venezuelan Coastal Cordillera (Sucre); West Indies, Trinidad, Suriname, French Guiana, Ecuador, Peru, Brazil. Some specimens of Quiina cruegeriana are difficult to distinguish from Quiina guianensis. The 5-lobed calyx, longer petiole, smaller fruits, and the pubescence of the terminal internodes distinguish it from this species. Quiina florida Tul., Ann. Sci. Nat. Bot. sér. 3, 11: 167. 1849. Quiina poeppigiana Tul., Ann. Sci. Nat Bot. sér. 3, 11: 161. 1849. Tree to 19 m tall. Lowland evergreen forests, riparian forests, 100–200 m; Amazonas (Los Pijiguaos, Río Cunucunuma, upper Río Orinoco, Río Padamo). Ecuador, Peru, Brazil, Bolivia. ◆Fig. 336. Quiina florida intergrades in several characters with Quiina rhytidopus. They are best separated by the microstructure of the leaf surface. In gross morphological characters Q. florida tends to have more elliptic rather than ovate leaves. The mature fruits are globose, and the basal part of the pedicels (below articulation) is more than 1 mm long. Quiina guianensis Aubl., Hist. Pl. Guiane 2(suppl.): 19, pl. 379. 1775. —Cola de pava, Cola de pava chiquita, Coloradito, Gaspadillo negro, Mulunyek (Arekuna). Understory tree or shrub to 12 m tall; leaves ± sessile, chartaceous, minutely serrulate; flowers with 6 or 7 petals. Moist and riparian lowland and montane forests, 50– 1000 m; Delta Amacuro (Río Amacuro, Serranía de Imataca), Bolívar (widespread). Trinidad, Guyana, Suriname, French Guiana. ◆Fig. 337. Quiina indigofera Sandwith, Bull. Misc. Inform. Kew 1931: 178. 1931. —Cola de pava, Icayek (Arekuna). Tree to 26 m tall; leaves deep green to bluish green (even when dried), minutely serrulate, with prominent, ridge-like secondary veins on lower surface; stipules densely brown-tomentose on adaxial side; fruits to 5.2 cm long. Evergreen moist lowland to lower montane forests, riparian forests, 100– 500 m; Delta Amacuro (north of Altiplanicie

de Nuria), Bolívar (La Escalera). Guyana. ◆Fig. 338. Quiina longifolia Spruce ex Planch. & Triana, Ann. Sci. Nat. Bot. sér. 4, 15: 314. 1861. —Ají de agojó, Hiel de pescado, Macho de puya, Simiyurí (= Palo de paloma), Tripa de pollo, Tripa de sardina. Understory tree or shrub to 10(–15) m tall; stipules foliaceous, ovate; inflorescences densely flowered. Evergreen riparian, gallery, and seasonally flooded forests, secondary forests, scrublands, frequently along black-water streams, 50–200(–300) m; Amazonas (widespread). Colombia, Brazil. ◆Fig. 340. Quiina macrophylla Tul., Ann. Sci. Nat. Bot. sér. 3, 11: 164. 1849. Shrub or tree to 15 m tall; rachis of inflorescence tomentose; pedicels short, to 2 mm long; sepals and petals 3 or 4. Evergreen lowland forests, gallery and seasonally flooded forests, 50–300 m; Amazonas (Caño Yureba, Río Cataniapo, Río Siapa). Apure; Colombia, Ecuador, Peru, Brazil. Quiina macrophylla has often been misidentified as Q. integrifolia Pulle, a species with longer petioles and pedicels, larger fruits, and inflorescences with apparently fewer flowers. Quiina schippii Standl., from Central America, might be conspecific with Q. macrophylla, and differs only in its slightly smaller leaves. Quiina obovata Tul., Ann. Sci. Nat. Bot. sér. 3., 11: 157. 1849. —Cola de pava negra, Coloradito, Waiye enuda (Yekwana). Quiina albiflora A.C. Sm., Trop. Woods 58: 30. 1939. Quiina oblanceolata Sandwith, Bull. Misc. Inform. Kew 1939/1940: 546. 1940. Tree to 20 m tall; secondary veins more prominent on the upper surface than the lower one; leaves minutely serrulate; stipules fused (visible at least at terminal nodes), curving slightly upward at the base. Evergreen lowland to lower montane forests, riparian forests, on granitic outcrops, 100– 500 m; Bolívar (Canaima, near Cerro Abismo, La Escalera, 35 km south of Manteco, Río Asa, Río Caura, Sierra de Maigualida). Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil.

402

Q UIINACEAE

Fig. 336. Quiina florida

Fig. 337. Quiina guianensis

Quiina 403

Fig. 338. Quiina indigofera

Fig. 339. Quiina pteridophylla

404

Q UIINACEAE

Fig. 340. Quiina longifolia

Fig. 341. Quiina tinifolia

Fig. 342. Quiina wurdackii

Touroulia 405

Quiina oiapocensis Pires, Bol. Técn. Inst. Agron. N. 38: 32. 1960. —Let’dété rowe (Piaroa), Manteco de agua. Tree to 16 m tall; pedicels, sepals, and fruits velvety-pubescent; leaves to 1 m long, glossy; stipules foliaceous, ovate; petioles 2.5–9 cm long. Evergreen lowland to montane forests, 100–800 m; Bolívar (Amaruaytepui, Serranía de los Pijiguaos), Amazonas (Caño Marieta in Río Mawarinuma basin). Guyana, French Guiana, Brazil. Quiina pteridophylla (Radlk.) Pires, Bol. Técn. Inst. Agron. N. 20: 48. 1950. —Touroulia pteridophylla Radlk., Sitzungsber. Math.-Phys. Cl. Königl. Bayer. Akad. Wiss. München 19: 218. 1889 [1890]. —Ojo de gavilán. Quiina acutangula Ducke, Arch. Jard. Bot. Rio de Janeiro 4: 143. 1925. Tree to 12 m tall; leaves in whorls of (3)4 or 5, pinnatifid (juvenile) or entire (adult), chartaceous; stipules setaceous to foliaceous, ± separate to completely fused. Evergreen lowland forests, 50–400 m; Bolívar (San Pedro), Amazonas (Cerro Duida, Cerro Yapacana, Huachica near San Carlos de Río Negro, Río Cataniapo basin). Zulia; Colombia, French Guiana, Peru, Brazil, Bolivia. ◆Fig. 339. Quiina rhytidopus Tul., Ann. Sci. Nat. Bot. sér. 3, 11: 166. 1849. Shrub or tree to 10 m tall. Gallery forests, seasonally flooded black-water riparian forests, Mauritia palm swamps, 50–300 m; Bolívar (Río Nichare, Río Paragua), Amazonas (Isla Ratón). Anzoátegui, Apure; Colombia, Guyana, Brazil, Bolivia. Apart from leaf microstructure, Quiina rhytidopus differs from the similar Q. florida in having ovate rather than elliptic leaves. The stipules are < 1.5 mm wide, the fruits are elliptic, and the lower part of the pedicels (below articulation) is < 1 mm long.

Quiina tinifolia Planch. & Triana, Ann. Sci. Nat. Bot. sér. 4, 15: 311. 1861. —Perro de agua. Quiina gracilis A.C. Sm., Trop. Woods 58: 26. 1939. Tree to 13 m tall; leaves ± sessile, rounded or subcordate at the base, with entire margin; sepals and petals 3 or 4. Riparian and seasonally flooded forests, white-sand scrub (bana), shrublands, 100–200 m; Amazonas (Río Negro, lower Río Orinoco). Brazil. ◆Fig. 341. Small-leaved forms of Quiina tinifolia are similar to Q. wurdackii, but they can be distinguished by their inflorescences with > 10 flowers. Quiina wurdackii Pires, Bol. Técn. Inst. Agron. N. 38: 30. 1960. —Wondaiyek (Arekuna). Tree to 17 m tall; leaves small, 2–7 × 1–3 cm, mostly attenuate at base; secondary veins 5–10; inflorescences short, with < 10 flowers; sepals and petals 3 or 4. Evergreen lowland to montane (riparian) forests, 100– 1300 m; Bolívar (Ptari-tepui), Amazonas (Laja Catipán on Río Yatúa). Endemic. ◆Fig. 342. Quiina wurdackii is a poorly documented species, whose known collections are from widely separated localities differing considerably in elevation. Quiina yatuensis J.V. Schneid. & Zizka, Novon 7: 406. 1997. Tree to 8 m tall; leaves 8–12 × 3.5–6 cm, elliptic, shortly acuminate, the very apex rounded; secondary veins 8–10; fruit 0.8–0.9 cm long, subglobose. Lower montane forests, 200–300 m; Amazonas (Río Yatúa). Endemic. Quiina yatuensis is similar to Q. rigidifolia Pires but differs in its smaller fruits and inflorescences with a less conspicuously thickened rachis and fewer flowers.

4. TOUROULIA Aubl., Hist. Pl. Guiane 492. 1775. by Georg Zizka Erect trees to 15(–30) m tall. Leaves opposite (in young plants ± alternate and then leaves usually larger than adult leaves), odd-pinnate, petiolate; leaf rachis sometimes (especially in young plants) winged toward the apex; stipules

406

Q UIINACEAE

interpetiolate, subulate to ± lanceolate, acute, glabrous; leaflets sessile, opposite to alternate, ovate to oblong, acute, base ± asymmetrically obtuse, margin serrate, the teeth spinose (secondary veins projecting beyond margin); intersecondary veins absent; terminal leaflets usually slightly larger than the lateral ones. Inflorescence terminal, thyrsoid; bracts ± broadly triangular, rounded, 1–9 × 0.8–3 mm, densely pilose abaxially, minutely pilose adaxially. Flowers unisexual or bisexual (plants androdioecious), pedicellate, the pedicel shortly pilose, articulating close to the middle. Sepals 5, unequal, broadly triangular, coriaceous, free or basally connate; Fig. 343. Touroulia guianensis

Fig. 344. Touroulia amazonica

R A F F L E S I A C E A E 407

petals (4)5, obovate, broadly rounded, glabrous. Stamens 50 to more than 100 in staminate flowers, fewer in bisexual flowers. Ovary syncarpous, (5)6–8-locular, with (5)6–8 styles. Fruit ± globose, longitudinally striate, glabrous, (5)6–8-locular, 1.5– 2.5 cm diameter, 1–4-seeded. Seeds 1 per locule, densely villous, ± ovate in outline. Venezuela, Guyana, Suriname, French Guiana, Brazil; 2 species, both in the flora area. Touroulia guianensis is widespread and well collected, whereas T. amazonica is much rarer. Separate generic status has been suggested for T. amazonica by Pires, but similarities in flower, fruit, and leaf morphology support maintaining one genus. Key to the Species of Touroulia 1.

1.

Secondary veins of leaflets arching upward near the margin, tertiary veins in about middle of intercostal areas geniculately bent; leaflet margin ± crisp; inflorescence 10–15 cm long; flowers ≤ 10 mm diameter ................................................................................................. T. guianensis Secondary veins of leaflets ± straight near the margin, not arching upward, tertiary veins not geniculately bent; leaflet margin flat; inflorescence 24–30 cm long; flowers 13–25 mm diameter ............... T. amazonica

Touroulia amazonica Pires & A.S. Foster, Bol. Técn. Inst. Agron. N. 20: 49. 1950. Tree to 15 m tall. Evergreen lowland to lower montane forests, 100–400 m; Amazonas (Río Negro, Sierra de la Neblina). Brazil. ◆Fig. 344.

Touroulia guianensis Aubl., Hist. Pl. Guiane 492, pl. 194. 1775. Tree to 10(–30) m tall; fruits reported to be edible. Evergreen lowland to lower montane, primary and secondary forests, usually on sandy soil, 200–700 m; Bolívar (Macizo del Chimantá, Soledad, Upata), Amazonas (Cerro Duida). Guyana, Suriname, French Guiana, Brazil. ◆Fig. 343.

RAFFLESIACEAE by George Yatskievych and Willem Meijer Achlorophyllous stem parasites (root parasites elsewhere), rootless, the endophytic system forming cord-like strands or plates of cells in association with the host cambium. Leaves absent. Inflorescences of solitary flowers terminating very short stems or (elsewhere) short racemes or spikes. Flowers unisexual (bisexual elsewhere), deeply perigynous to ± epigynous, subtended closely by 1 or more series of bracts. Perianth of 4 or 5 petaloid or sepaloid tepals (to 16 elsewhere), these often basally connate. Stamens 8–many; filaments and usually also the anthers adnate around a thickened central column; anthers opening by slits or pores. Pollen single or in tetrads. Stigmatic region at or near tip of central column, which is sometimes

408

R AFFLESIACEAE

expanded apically (clavate to discoid). Ovary completely or partially inferior (appearing superior in Mitrastemon), unilocular, at maturity with 4–8 parietal placentae (numerous irregular placentae elsewhere). Fruits berry-like. Seeds numerous, tiny, with oily endosperm; embryo undifferentiated. Nearly worldwide (except Antarctica), but most diverse in the tropics; 8 or 9 genera and ca. 55 species, 1 species in the flora area. The family is described above and treated here in the broad sense of Meijer (1993). The two small-flowered genera present in the flora area and adjacent Brazil (Apodanthes and Pilostyles), along with the two East African species of Berlinianche (Harms) Vattimo-Gil, traditionally have been classified in the tribe Apodantheae R. Br. Preliminary molecular data suggest that these small-flowered plants may not be very closely related to the rest of the family, and some botanists prefer to treat them in the segregate family Apodanthaceae (Nickrent and Duff 1996; Angiosperm Phylogeny Group 2003). Pilostyles Guill. is a poorly understood genus taxonomically comprising 9–25 species. It has a highly disjunct distribution mainly from the southwestern U.S.A. to South America but also in southwestern Asia and southwestern Australia. All of the species parasitize various genera of legumes (Fabales). The genus differs from Apodanthes in having flowers with the tepals sepaloid, not differentiated from the bracts, attached at the broad nonpeltate base, and persistent (even at fruiting). Pilostyles has not been collected yet in the flora area, but two species have been collected immediately to the south in Estado Roraima, Brazil [P. galactiae Uhl and P. blanchetii (Gardn.) R. Br.]. Rafflesiaceae sensu stricto comprise three genera endemic to the IndoMalesian region that parasitize stems or roots of Vitaceae (mostly Tetrastigma). They are characterized by large (often very large) flowers. Recent molecular studies have suggested that these genera are related to the Malpighiales (Barkmann et al. 2004). The New World Bdallophyton Eichler is of uncertain classification, but may be more closely related to the Apodanthes group than to other genera of Rafflesiaceae sensu lato. It contains two species parasitic on roots of Burseraceae from Mexico, Central America, and Colombia, and has spiciform inflorescences with medium-sized flowers. The other American genus, Mitrastemon Makino, which parasitizes mainly roots of Fagaceae, consists of two taxa, one in eastern Asia and the other known from Mexico, Central America, and Colombia. Molecular analyses suggest that this genus is misplaced in the Rafflesiaceae and should be reclassified in the Ericales (Barkmann et al. 2004). The whole complex of genera described above is very poorly collected. Specimens in herbaria often are in poor condition and difficult to interpret because of distortion and/or shattering of the flowers and fruits. Thus our knowledge of distribution, taxonomy, morphological variation, and host ranges is deficient, especially for the small-flowered genera. It is not even clear whether plants are monoecious or dioecious.

References Cited Angiosperm Phylogeny Group. 2003. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG II. Bot. J. Linn. Soc. 141: 399–436.

Apodanthes 409

Barkmann, T. J., S.-H. Lim, K. M. Salleh, and J. Nais. 2004. Mitochondrial DNA sequences reveal the photosynthetic relatives of Rafflesia, the world’s largest flower. Proc. Natl. Acad. Sci. U.S.A. 101: 787–792. Croat, T. B. 1978. Flora of Barro Colorado Island. Stanford, CA: Stanford University Press. Gómez, L. D. 1983. Rafflesiaceae. In Flora Costaricensis, edited by W. Burger. Fieldiana, Bot. n.s. 13: 89–93. Meijer, W. 1993. Rafflesiaceae. In The Families and Genera of Vascular Plants, Vol. 2, edited by K. Kubitzi, J. G. Rohwer, and V. Bittrich, pp. 557–563. Berlin: Springer-Verlag. Nickrent, D. L., and R. J. Duff. 1996. Molecular studies of parasitic plants using ribosomal RNA. In Advances in Parasitic Plant Research. Junta de Andalucia, edited by M. T. Moreno, J. I. Cubero, D. Berner, D. Joel, L. J. Musselman, and C. Parker, pp. 28–52. Cordoba, Spain: Dirección General de Investigación Agraria. Vattimo-Gil, I. de. [1970] 1971. Contribucão ao conhecimento da tribo Apodanthea R. Br. Parte I—conspecto das espécies. (Rafflesiaceae). Rodriguésia 26: 37– 62. 1971. Vattimo-Gil, I. de. 1973. Notas sobre o gênero Apodanthes Poit. com descrição de duas novas espécies. Rev. Brasil. Biol. 33: 135–141. Vattimo-Gil, I. de. 1978. Uma nova espécie de Apodanthes Pot. (Rafflesiaceae). Rodriguésia 29: 47–51. 1978. 1. APODANTHES Poit., Ann. Sci. Nat. (Paris) 3: 422, t. 26. 1824. Apparently dioecious, parasitic on stems of Flacourtiaceae. Staminate and pistillate flowers similar in appearance, bursting individually through the host bark and leaving long-persistent, circular, crateriform, sometimes irregularly lobed scars of host tissue after abscission, oblong-ovoid to nearly globose, the expanded proximal portion tapered to a short stout column with a slightly expanded hemispherical apex. Floral bracts persistent, in 2 series, scale-like; basal series of 2(3) bracts, these loosely ascending to spreading, broadly obovate to nearly circular, slightly imbricate proximally, connate at the base, usually light yellowish brown; distal series of 4 bracts, these calyculate, strongly imbricate, broadly obovate to oval, rounded at the tip, the basal 1/4–1/3 adnate to the ovary, usually light yellow, drying brown. Perianth caducous, leaving small slightly impressed circular attachment scars on the distal 1/3 of the ovary; tepals 4, petaloid, broadly spatulate to broadly obovate or nearly circular, slightly peltate, attached just above the often somewhat asymmetric base, appressed or more commonly spreading at the tip, imbricate, white to cream-colored or light yellow, sometimes reddish-tinged. Staminate flowers with ca. 20 sessile anthers under the glandular rim of the dilated column tip. Pistillate flowers with the stigmatic region terminal or subterminal on the column tip; ovary with 4 placentae. Fruits somewhat larger than the flowers, ellipsoid to nearly globose, yellow to yellowish orange. Seeds ± globose, with a finely reticulate, chartaceous outer testa covering a harder inner layer. Belize, Honduras, Costa Rica, Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 1–7 species, 1 in Venezuela.

410

R AFFLESIACEAE

The genus Apodanthes is badly in need of further taxonomic study. Because these mostly endophytic parasites are character-poor and few good specimens or photographs exist, it is difficult to evaluate the importance of the subtle differences in floral morphology that have led some authors to recognize as many as seven species (Vattimo-Gil 1971; 1973; 1978). According to Gómez (1983), the fruits are eaten by birds (tanagers of the genus Thraupis) and the flowers are visited by bees in the genus Trigona. Croat (1978) cited notes on the label of a collection made by Robin Foster in Central America that the flowers also are visited by small butterflies and mosquitoes. Apodanthes caseariae Poit., Ann. Sci. Nat. (Paris) 3: 422, t. 26. 1824. Apodanthes flacourtiae H. Karst., Linnaea 28: 413. 1856. Apodanthes roraimae Vattimo-Gil, Rodriguésia 29: 48, figs. 1–5. 1978. Parasitic on trunks and branches of Flacourtiaceae (mainly Casearia and Xylosma), appearing as patches of dense flowers, fruits, and/or scars on the host bark; subtending cup-like bases of host tissue 1.5–3.0 mm diameter, 0.5–2 mm tall; flowers 3.5–5 mm long (pistillate flowers enlarging to 6–10 mm in fruit), oblong-ovoid to nearly globose. Riparian and lower montane forests, 600–1200 m; Bolívar (northeast of Serranía Marutaní, Salto Pulpul on Uaipán-tepui). Belize, Honduras, Costa Rica, Panama, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 345.

Fig. 345. Apodanthes caseariae

RANUNCULACEAE by Paul E. Berry Herbs, shrubs, or woody climbers. Leaves alternate or opposite, simple or compound, petiolate, often sheathing at the base, with or without stipules. Flowers bisexual or unisexual, solitary or aggregated in cymes, racemes, or panicles, the terminal unit cymose. Perianth either with distinct calyx and corolla, intergrading from sepals to petals, or all sepaloid or petaloid. Calyx when present with sepals (3–)5–8 (or more, often becoming petaloid), distinct to (partially) gamosepalous, sometimes spurred, imbricate or valvate. Corolla when definable of 3–50 petals, distinct to gamopetalous, plane to spurred, imbricate. Androecium of (5–)15–100 free, centripetal stamens, sometimes staminodial and nectariferous; staminodes when present petaloid or nonpetaloid; anthers adnate, nonversatile, dehiscing longitudinally. Gynoecium of (1–)3–100 carpels, generally apocarpous; each carpel with 1–

Clematis 411

100 ovules, placentation marginal or basal; styles, when present, usually persistent in fruit as a beak or plume. Fruits usually dry, achenes or follicles, rarely a berry or a capsule, aggregate or rarely compound. Cosmopolitan; about 62 genera and 2500 species, 1 genus and 2 species in the flora area. 1. CLEMATIS L., Sp. Pl. 543. 1753. Woody vines or lianas, less often shrubs or herbs. Leaves opposite, simple to compound, evergreen to deciduous; petioles sometimes twining; stems often grooved or ridged. Infloresences axillary or terminal, paniculate-cymose or rarely flowers solitary; flowers bisexual or unisexual (plants then monoecious or dioecious), erect to hanging or divergent, radially symmetrical. Sepals 4 or 5(–8), mostly valvate; petals absent; stamens numerous; filaments slender to thick-flattened; carpels 5– 150, free, each flattened, with a single ovule; styles elongate, usually densely plumose, elongating in fruit. Fruit a head-like aggregate of achenes with persistent plumose styles. Cosmopolitan, most diverse in eastern Asia; about 300 species, 2 in Venezuela, both in the flora area. Both species occurring in Venezuela belong to the same group, section Clematis subsection Dioicae, a group of 18 North and South American species (C. GreyWilson. Clematis the Genus. Timber Press, Portland, OR. 2000). Alternatively, they have been treated simply as belonging to subgenus Clematis (N. P. Moreno. Taxonomic Revision of Clematis L. subgenus Clematis (Ranunculaceae) for Latin America and the Caribbean. Ph.D. dissertation, 255 pp., Rice University, Houston, TX. 1993). The species taxonomy below is based on Moreno’s thesis. Key to the Species of Clematis 1.

1.

Mature leaves with 5 or more leaflets, the margins entire or with shallow teeth cut to < 1/4 the distance to the midvein, glabrate to sparsely sericeous on lower surface; carpels > 25 per flower; achenes long and narrow, 4–5 mm long ................................................................ C. guadeloupae Mature leaves with 3 leaflets, the margins entire, unlobed, sparsely to densely sericeous to tomentose on lower surface, with white to yellow trichomes 0.5–1.0 mm long; carpels < 25 per flower; achenes broadly ovate to suborbicular in outline, 3–4 mm long, 2.5–3.0 mm wide ................................................................................................ C. populifolia

Clematis guadeloupae Pers., Syn. Pl. 2: 99. 1806. Dioecious woody vine to 10 m long; stems ribbed, brown to reddish brown, glabrate to sparsely strigulose, with short, tortuous, white trichomes; leaves with 5 or 7 primary leaflets; ultimate leaflet blades ovate to lanceolate, 3.0–12.5 × 3.0–6.5 cm, apically acute to acuminate, rounded to subcordate at the base, margin entire or with 1 or 2 acute teeth per side; petiole to 6 cm long, petiolules 1.5– 3.5 cm long; inflorescences axillary or termi-

nal, bearing 40–200 or more flowers, peduncle 3.5–17 cm long, rachis 5–50 cm long, with 5–10 nodes bearing pairs of lateral branches; pedicels 9–35 mm long; sepals white, cream, or greenish, obovate, 5–9 × 2.0–2.5 mm; stamens 36–50(–80); filaments 3–5 mm long; anthers ca. 0.75 mm long; carpels 25–60; achenes brown to reddish brown when dry, flattened, fusiform to narrowly falcate, 4–5 mm × 1.25–2.0 mm, sericeous to hirtellous, persistent style 4–6.5 cm long with trichomes 3–4 mm long. Panama, West Indies,

412

R ANUNCULACEAE

Colombia, Venezuela, Ecuador, Peru, Bolivia, western Brazil; 2 varieties, both in Venezuela, 1 of these in the flora area. Clematis guadeloupae is identified by the presence of entire, glabrate to sparsely pubescent leaflets, strictly cymose inflorescences, long robust fruiting pedicels, and numerous fusiform achenes. Variety medusaea is distinguished from var. guadeloupae by its bipinnate leaves (or at least the basal pair of leaflets 3-foliolate versus simply pinnate), membranous (versus thickened) leaves, and pubescent (versus glabrate) vegetative growth.

C. guadeloupae var. guadeloupae Clematis caracasana Kunth ex DC., Syst. Nat. 1: 141. 1818. Clematis caripensis Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 5: 36. 1821. Lower montane forest edges, 200–400 m; Bolívar (Río Supamo). Panama, Puerto Rico, Lesser Antilles (Guadeloupe, Martinique), Colombia, Trinidad, Tobago, Ecuador, eastern Peru, Bolivia, western Brazil; Aragua, Monagas, Sucre, Táchira, Vargas. ◆Fig. 346. Plants treated here as Clematis guadaloupae var. guadaloupae were previously identified as C. brasiliana DC. [= C. dioica

Fig. 346. Clematis guadeloupae var. guadeloupae

R A PAT E A C E A E 413

var. brasiliana (DC.) Eichl.], but these two taxa are distinct and can be distinguished as follows: carpels > 25 per flower, achenes long and narrow, 4–5 mm long, and distribution in the Antilles and northern and western South America in var. guadaloupae (versus carpels < 25, achenes ovate to elliptic in outline and 3–4 mm long, and distribution restricted to eastern and southern Brazil and Paraguay in C. brasiliana). Clematis populifolia Turcz., Bull. Soc. Imp. Naturalistes Moscou 27: 272. 1854. Clematis floribunda Triana & Planch., Ann. Sci. Nat. Bot. sér. 4, 17: 2. 1862. Clematis goudotiana Triana & Planch., Ann. Sci. Nat. Bot. sér. 4, 17: 10. 1862. Dioecious woody vine; stems reddish to yellow-brown, sulcate, sparsely pilose to strigose, with white to yellow hairs; leaves 3foliolate, leaflets ovate to occasionally lanceolate, membranous to slightly succulent,

5.5–13.0 × 3.0–9.5 cm, acute to short acuminate at the apex, rounded to cordate at the base, margin entire; petioles 2.5–10.5 cm long; terminal petiolule 1–4 cm long, lateral ones 0.5–2.6 cm long; inflorescences usually axillary, compound with a central rachis, bearing 10–70 or more flowers; pedicels 10– 26 mm long; sepals white to greenish white, oblanceolate to obovate, 7–10 × 2.0–3.5 mm; stamens 35–50, filaments 3–6 mm long, anthers ca. 1 mm long; carpels 8–25; achenes broadly ovate to suborbicular, 3–4 mm × 2.5– 3.5 mm, with a distinct border to 0.3 mm wide, persistent styles 4–6 cm long with trichomes 3–4 mm long. Lowland to montane forest edges and disturbed areas, 200–2200 m; Bolívar (Cerro Bolívar, Sororopán-tepui). Mérida, Táchira, Vargas; Colombia, Ecuador. Clematis populifolia was formerly included in C. haenkeana C. Presl, but that species has more carpels and 5-foliolate leaves.

RAPATEACEAE by Paul E. Berry Perennial herbs with a stout rhizome, commonly with copious mucilage in the leaf bases and young inflorescences. Leaves basal, generally distichous with a folded, conduplicate sheath, usually fan-shaped but sometimes spirally arranged; blade parallel-veined, normally flattened and rotated at a right angle to the sheaths to become parallel with them, but sometimes compressed or cylindrical. Inflorescence scapose (scape absent in the West African Maschalocephalus), with 1–many spikelets arranged in a head, spike, or corymb, the spikelets usually sessile, sometimes short-pedicellate, rarely longer-pedicellate and then with pedicels of differing lengths in the same inflorescence; each spikelet with a series of imbricate bracteoles and a single, terminal flower; inflorescence usually subtended by an involucre of 2 or more bracts (absent in most Stegolepis). Flowers bisexual, trimerous, actinomorphic or slightly zygomorphic. Sepals 3, free or shortly connate at the base, the lobes lanceolate, imbricate, often rather stiff and translucent, navicular, and sharptipped; petals 3, mostly ephemeral, yellow, red, or white, usually connate at the base, the free lobes imbricate, generally emergent and strongly spreading at anthesis, but in some taxa largely enveloped by the sepals and remaining tightly imbricate. Stamens 6; filaments short, generally connate at the base and adnate to the corolla tube; anthers basifixed, usually shorter than the corolla, 2- or rarely 1-thecal, opening by 1, 2, or 4 apical pores or slits. Ovary superior, syncarpous, 3-carpellate; ovules anatropous, several to numerous per locule on axile placentas or 1 or 2 per locule and sub-basally attached, sometimes only 1 locule fertile; style solitary, terminal; stigma simple. Septal nectaries present at least in Guacamaya,

414

R APATEACEAE

Kunhardtia, and Schoenocephalium. Fruit a loculicidal capsule. Seeds 1–15 per fruit, with a small, lenticular embryo, the testa smooth to striate, furrowed, or muriculate, sometimes surrounded by an arillate structure or with a terminal appendage; endosperm copious and starchy. Panama, Colombia, Venezuela, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, with 1 monotypic genus in tropical West Africa; 17 genera and 100 species; 14 genera and 72 species in the flora area. Two monotypic genera, Windsorina and Potarophytum, occur in the lowlands of adjacent Guyana, and the monotypic Maschalocephalus occurs in Liberia and Sierra Leone in western Africa. Many species of Rapateaceae produce copious amounts of mucilage in the leaf sheaths and inflorescence bracts through specialized trichomes. All species have poricidal anthers. Species with yellow, spreading corollas are generally buzz-pollinated by bees, but the three genera of Schoenocephalieae (Guacamaya, Kunhardtia, and Schoenocephalium) have reddish coloration in the flowers or inflorescence and have tightly imbricate, ± tubular corollas. These flowers produce nectar, and visits by hummingbirds have been observed. Visits by birds to Kunhardtia are frequent, because the flowers remain open for several days. In Guacamaya and Schoenocephalium, however, flowers open very sparingly and only for a few hours at a time, and visits may be limited to early morning hours or other times of ± synchronous anthesis. Hummingbird mites (Tropicoseius, Rhinoseius, and Proctolaelaps species) have been collected by the author in flowers of each of the three genera of Schoenocephalieae, which supports the idea that they are regularly visited by hummingbirds. Key to the Genera of Rapateaceae 1. 1.

2(1). 2.

3(2).

3.

4(2).

Spikelets numerous on one side of a spike that is adnate to the basal portion of a large spathaceous bract ..................................... 13. Spathanthus Spikelets 1–many, variously arranged at the end of a scape, but not on a spike adnate to a single large bract; inflorescence either lacking subtending bracts or with 2–several subtending or enveloping bracts .... 2 Inflorescence without evident bracts, or if bracts are present, then the inflorescence with just 1–3 spikelets ....................................................... 3 Inflorescence with involucral bracts present, these free or connate, sometimes with 2 opposing valves, or saccate and enveloping the inflorescence; spikelets numerous ..................................................................... 4 Plants epiphytic on trees or on boulders, the peduncles arching or drooping at anthesis; anthers smooth; seeds disk-shaped, covered by a spongy arillate structure and coherent in linear packets; capsule with the exocarp peeling back away from the endocarp at dehiscence .................................................................................................... 4. Epidryos Plants mostly terrestrial, in meadows, thickets, or on rocks, rarely epiphytic, the peduncles erect at anthesis, never hanging; anthers transversely rugose; seeds prismatic or pyramidal, not arillate or coherent to each other; the endocarp of the capsule not separating from the exocarp at dehiscence .................................................................. 14. Stegolepis Involucral bracts 3 or more (or if apparently 2 as in Monotrema, then of unequal sizes) ........................................................................................ 5

R A PAT E A C E A E 415

4.

Involucral bracts 2 and equal in size and shape, or sometimes fused into a single, saccate bract that envelops the whole inflorescence ................ 7 5(4). Leaf sheaths 4–7 cm wide, the blades 3–5 cm wide; involucral bracts orbicular or suborbicular, rounded or obtuse; bracteoles obtuse ............................................................................................ 7. Marahuacaea 5. Leaf sheaths 0.7–2 cm wide, the blades 0.4–1.5 cm wide; involucral bracts broadly lanceolate, acuminate-attenuate; bracteoles finely acuminate ................................................................................................................ 6 6(5). Peduncle conspicuously ribbed, widened just below the inflorescence to 7–10 mm wide, elsewhere 3–4 mm wide; outer involucral bracts 10–32 mm wide; leaf blades prominently thick-ribbed, 60–100 cm long ......................................................................................... 1. Amphiphyllum 6. Peduncle terete to slightly ribbed, mostly 1–2 mm wide, not conspicuously broadened below the inflorescence; outer involucral bracts 2– 7 mm wide; leaf blades not thick-ribbed, mainly 20–45 cm long (60– 80 cm in M. aemulans) ......................................................... 8. Monotrema 7(4). Bracteoles of spikelets and sepals mainly red, rose, or salmon at anthesis, occasionally pale yellow or cream; corolla at anthesis either with exserted, bright red, downcurving petals, or else white to dull yellow and mostly enclosed within the sepals with just the tips of the petals slightly spreading at anthesis (in dried material the corolla sometimes is not evident) ........................................................................................ 8 7. Bracteoles of the spikelets and sepals greenish, yellow, or whitish at anthesis; corolla yellow or rarely white, exserted well beyond the sepals at anthesis, the blades of the petals usually spreading ..................... 10 8(7). Leaf blades 3–6.5 cm wide; corolla red and conspicuously exserted from the sepals .............................................................................. 6. Kunhardtia 8. Leaf blades 0.2–3 cm wide; corolla white to yellow, enclosed within the sepals except for the tips opening slightly and very briefly at anthesis ................................................................................................................ 9 9(8). Inflorescence bracts reflexed at maturity; junction of leaf blade and leaf sheath marked by an obvious contraction or else the blades very narrow and terete; peduncle 3–7 mm wide below the inflorescence; heads 2–3.5 cm diameter, ± isodiametric .......................... 12. Schoenocephalium 9. Inflorescence bracts not becoming reflexed, saccate and enveloping the head, then ruptured by the expanding spikelets into several to many persisting segments; junction of leaf sheath and leaf blade inconspicuous, the blade flattened, never terete; peduncle conspicuously enlarged to 10–17 mm wide below the inflorescence; heads 4–5 cm wide at maturity, usually widest or truncate at the base ......................... 5. Guacamaya 10(7). Inflorescence bracts widely spreading or reflexed; spikelets not subtended by a secondary bract, seeds without fine protuberances ....... 11 10. Inflorescence bracts erect or ascending, sometimes saccate, or if widely spreading the spikelets subtended by a secondary bract and the seeds with fine protuberances ....................................................................... 12 11(10). Seeds smooth, with an apical tufted appendage; leaves arching, the blade plane, with an evident midvein on the abaxial surface; spikelet bracteoles usually with recurving, glandular-setose tips; petals generally > 5 mm wide ............................................................. 2. Cephalostemon

416

R APATEACEAE

11.

Seeds longitudinally striate, without an apical appendage; leaves erect, the blade either plane, with an evident midvein as above, or else terete or compressed-cylindrical without an evident midvein; spikelet bracteoles without glandular-setose tips; petals generally < 5 mm wide ...................................................................................................... 3. Duckea 12(10). Inflorescence with 2–4 spikelets, the scape winged and flattened most of its length; anthers 2-locular .................................................. 9. Phelpsiella 12. Inflorescence with 5–many spikelets, the scape not winged or winged only near the top; anthers 4-locular ................................................... 13 13(12). Inflorescence laterally compressed, subtended by 2 opposing bracts that open like a clam shell; anthers with a hooded apical appendage; ovary with one ovule per locule .......................................................... 10. Rapatea 13. Involucral bracts saccate, connate, pierced by the developing spikelets, often persisting as a cap-like appendage protruding from the summit of the ± globose head or as torn remnants at the base; anthers without a hooded appendage; ovary with several ovules per locule .......... 11. Saxofridericia 1. AMPHIPHYLLUM Gleason, Bull. Torrey Bot. Club 58: 332, pl. 25, figs. E, F. 1931. Perennial herbs 50–100 cm tall. Leaf sheaths linear-lanceolate, 12–20 cm long, 1–2 cm wide; blades narrowly linear, 60–100 cm long, 0.6–1.5 cm wide, thick, rigid, 8–10-veined. Peduncles 70–100 cm tall, conspicuously ribbed, mostly 3–4 mm wide but flattened and widened to 8–10 mm wide at the apex; involucral bracts several, ovate-lanceolate, covered with mucilage when fresh, the outer ones 25–40 mm long by 8–16 mm wide, the inner ones 20–25 mm long, 10–15 mm wide. Spikelets 3–10 per inflorescence, sessile, 20–22 mm long; bracteoles 15–20, oblong-lanceolate, with recurved tips. Petals yellow, the exserted blade suborbicular, ca. 12 mm long × 10 mm wide. Anthers 4-locular, opening laterally by 2 subapical pores. Ovary 3-locular, pluriovulate. Venezuela (endemic to Cerro Duida, Amazonas); 1 species. Amphiphyllum rigidum Gleason, Bull. Torrey Bot. Club 58: 333, pl. 25, figs. E, F. 1931.

Swampy, open sandy areas on tepui summit, 1200–2000 m; Amazonas (Cerro Duida). Endemic. ◆Fig. 347.

2. CEPHALOSTEMON R.H. Schomb., Rapatea 9. 1845. Terrestrial herbs, mostly of swampy places. Leaves with flattened blades, the midvein evident on the abaxial surface; sheaths marcescent. Inflorescence subglobose with numerous spikelets or with just 2 or 3 spikelets, subtended by 2 linear-lanceolate, conspicuous bracts; spikelets sessile, the bracteoles gradate, setiform, often with an enlarged glandular tip. Petals yellow, obcordate-obovate, retusely apiculate, widely spreading at anthesis, generally > 5 mm broad. Anthers 4-locular, the posterior lobes dehiscing by a longitudinal, subterminal, introrse slit extending to the apex of the anterior lobes. Ovary 3-locular, 1 ovule per locule. Seed broadly ellipsoid, smooth, sub-basally attached, with a conspicuous ventral raphe and a dense tuft of papillate appendages on the top. Southeastern Colombia, southern Venezuela, Suriname, Brazil (widespread, especially in Brazilian Shield area), Bolivia (Santa Cruz); 5 species, 2 in Venezuela, both found only in the flora area.

Amphiphyllum 417

Fig. 347. Amphiphyllum rigidum

418

R APATEACEAE

The three other species in the genus are restricted to the Brazilian Shield (Cephalostemon angustatus Malme, C. gracilis (Poepp. & Endl.) R.H. Schomb., and C. riedelianus Körn.). They are morphologically similar to C. affinis, and some of them may prove with further study to be conspecific. See notes under Duckea concerning differences between the two genera. Key to the Species of Cephalostemon 1.

1.

Spikelets 5–12 per head; involucral bracts 1.5–4 cm long; bracteoles weakly veined, ending in elongate, setose, clavate appendages ....................................................................................................... C. affinis Spikelets 2 or 3 per head; involucral bracts 1–2 cm long; bracteoles conspicuously veined, the upper ones ± appressed or ascending, the apex slender, acuminate or attenuate ....................................... C. microglochin

Fig. 348. Cephalostemon affinis

Duckea 419

Cephalostemon affinis Körn., Linnaea 37: 447. 1872. Herb 30–70 cm tall; leaf blades 4–8 mm wide; peduncle 4-ridged, bracts 1.5–4 cm long; spikelets 5–12 per head. Moist areas of white-sand savannas, 50–200 m; Amazonas (Caño Ucata southeast of Síquita, base of Cerro Yapacana, Guarinuma, La Esmeralda, Pimichín, Santa Bárbara del Orinoco, lower Río Ventuari). Suriname (Tafelberg savannas), Brazil (Pará). ◆Fig. 348.

Cephalostemon microglochin Sandwith, Kew Bull. 1929: 125. 1929. Herb 30–50 cm tall; leaf blades 2–3 mm wide; bracts 1–2 cm long, ± same length as the spikelets, these just 2 or 3 per head. Wet sandy and sometimes hilly savannas, 50–600 m; Amazonas (Río Manapiare basin). Brazil (Mato Grosso), Bolivia (Santa Cruz). Cephalostemon microglochin is a rare species with a notable disjunction between Venezuela and southern Brazil and Bolivia.

3. DUCKEA Maguire, Mem. New York Bot. Gard. 10(1): 41. 1958. Herbs of swampy places. Leaves sedge-like, erect, either linear with a flat blade or cylindrical or compressed-cylindrical; sheaths marcescent. Inflorescence subglobose to cylindrical, with 2 spreading to reflexed lanceolate bracts; head with numerous spikelets; spikelets sessile, the bracteoles gradate. Sepals coriaceous, basally connate; petals basally connate, the blade lanceolate to suborbicular, generally < 5 mm diameter, ephemeral (often only lasting a few hours before wilting or dehiscing). Stamens 4-locular, opening by an oblique terminal pore. Ovary 3-locular, 1 sub-basal ovule per locule. Seed ellipsoid-oblongoid, conspicuously striate, generally with a conspicuous ventral raphe. Colombia (Guainía), southern Venezuela, Brazil (Amazonas, Pará, Roraima); 4 species, all in Venezuela and the flora area. Duckea is most closely related to Cephalostemon. Steyermark (Ann. Missouri Bot. Gard. 75: 1565–1586. 1988) treated Duckea as a synonym of Cephalostemon, but they appear to be coherent sister groups rather than a paraphyletic assemblage. Besides the nonpapillate and striate seeds, Duckea can be distinguished from Cephalostemon by its erect (versus arching) leaves, several species with terete or compressed-cylindrical leaves, the generally smaller or more lanceolate petals, and the more numerous, denser, smaller spikelets. Most Cephalostemon species are distinguished by the setose bracteoles that are usually gland-tipped. Key to the Species of Duckea 1.

1.

2(1).

2.

3(1). 3.

Leaf blades terete or compressed-cylindrical, the midvein not elevated or easily distinguishable; spikelet bracteoles rounded, obtuse, or acute, not recurved at the tips ......................................................................... 2 Leaf blades plane, flattened, the midvein elevated and visible on the lower surface; spikelet bracteoles acuminate-aristate or acute and recurved at the tips ................................................................................... 3 Spikelet bracteoles rounded or obtuse; leaf blades compressed-cylindrical, 4–8 mm wide; peduncles mostly 1–4 m tall; heads oblong-cylindric, 10–40 mm tall ................................................................................. D. flava Spikelet bracteoles acute; leaf blades terete, 1–4 mm wide; peduncles 0.8–1.5 m high; heads subhemispheric to spherical, 8–15 mm tall ............................................................................................... D. junciformis Bracteoles ending in recurved, acute to attenuate tips; outer bracteoles strongly 5–7-veined ................................................................ D. squarrosa Bracteoles ± appressed or ascending, with slender, acuminate or attenuate apex; outer bracteoles inconspicuously veined ......... D. cyperaceoidea

420

R APATEACEAE

Fig. 349. Duckea flava

Fig. 350. Duckea junciformis

Duckea 421

basin, lower Río Guasacavi, Río Pacimoni, Yavita). Colombia (Guianía), Brazil (Amazonas, Pará, Roraima). Duckea flava (Link) Maguire, Mem. New York Bot. Gard. 10(1): 43. 1958. —Schoenus flavus Link in Spreng. et al., Jahrb. Gewächsk. 1(3): 75. 1820. —Cephalostemon flavus (Link) Steyerm., Ann. Missouri Bot. Gard. 75: 1565. 1988. —Rabo de ratón. Generally robust, narrow herb 1–3 m tall; peduncles 1.5–3.5 m tall; leaves fleshyspongy, without evident veins, 4–8 mm wide; bracts 20–50 mm long, 4–6 mm wide, the head usually cylindrical, mostly 20–40 mm tall; bracteoles obtuse; petals suborbicular, ca. 5 mm wide. In wet, often flooded areas at forest edges, shrubby or open white-sand savannas, edges of granitic outcrops, 50–200 m; Amazonas (Caño San Miguel, Carmelitas, base of Cerro Yapacana, along Río Atabapo and its tributaries, Río Autana, Río Baría, Río Casiquiare and its tributaries, Río Guainía and tributaries, upper Río Guayapo, lower Río Ventuari). Colombia (Guainía), Brazil (Amazonas). ◆Fig. 349. Duckea flava is the tallest species in the genus and grows closest to streams and rivers subject to frequent flooding.

Fig. 351. Duckea squarrosa

Duckea cyperaceoidea (Ducke) Maguire, Mem. New York Bot. Gard. 10(1): 42. 1958. —Cephalostemon cyperaceoides Ducke, Arch. Jard. Bot. Rio de Janeiro 1: 10. 1915. Herb 50–140 cm tall; leaves 3–5 mm wide; head ± round to cone-shaped, 15–22 mm wide, the bracts 30–90 mm long, 3–5 mm wide; spikelets numerous, bracteoles 3–6 mm long; sepals ca. 6 mm long, petals oblong-lanceolate, 2–3 mm long, 1.5 mm wide. Swampy white-sand savannas, rock outcrops of tepui slopes, 50–800 m; Amazonas (Cerro Aracamuni, Río Casiquiare basin, Río Guainía

Duckea junciformis Maguire, Mem. New York Bot. Gard. 10(1): 43, fig. 7A–F. 1958. —Cephalostemon junciformis (Maguire) Steyerm., Ann. Missouri Bot. Gard. 75: 1565. 1988. Herb 40–100 cm tall; leaves terete, 1.5–4 mm diameter; heads globose, 14–18 mm diameter, 8–15 mm tall, the bracts 20–40 mm long, 1.5–2.5 mm wide; bracteoles 3–6 mm long, acute; sepals 7–8 mm long, petals with the blade oblong-ovate, ca. 3 mm long. Marshy areas of white-sand savannas, 50– 200 m; Amazonas (road from Maroa to Yavita, near mouth of Río Atacavi, Río Guainía basin). Adjacent Colombia (Guainía). ◆Fig. 350. Duckea squarrosa (Willd.) Maguire, Mem. New York Bot. Gard. 10(1): 42. 1958. —Dichromena squarrosa Willd. ex Link in Spreng. et al., Jahrb. Gewächsk. 1(3): 77. 1820. —Cephalostemon squarrosus (Willd.) Körn., Linnaea 37: 450. 1872.

422

R APATEACEAE

Herb 50–110 cm tall; leaf blades flattened, 3–6 mm wide, conspicuously veined; head globose or depressed-globose, 20–25 mm diameter, bracts 6–10 cm long, 5–6 mm wide; bracteoles stiff and recurved at pointed tips, 10–12 mm long, strongly 5–7-veined; petals stiff-lanceolate, the blade ± 6 mm long.

Swampy or flooded areas, edges of whitesand savannas, 50–400 m; Bolívar (middle Río Parguaza, Río Parupa, Urimán), Amazonas (base of Cerro Yapacana, Laguna Pasiba, Río Baría, San Carlos de Río Negro, near Yavita). Colombia (likely), Brazil (Amazonas, Roraima). ◆Fig. 351.

4. EPIDRYOS Maguire, Taxon 11: 57. 1962. Epiphyton Maguire, Mem. New York Bot. Gard. 10(1): 31. 1958, non Bornem. 1886. Perennial herbs, usually fan-shaped epiphytes, with ± elongate, subfleshy rhizome. Leaf bases broadly distichous, submembranous, blades linear and arching. Peduncles numerous, filiform, arching or drooping at anthesis. Inflorescence ebracteate or bracts 2 and extremely reduced, spikelets 1–7, sessile or unequally pedicellate. Petals lanceolate, yellow. Anthers 2-locular, with 2 pores ± confluent toward the apex. Ovary 3-locular, pluriovulate. Capsule ovoid, endocarp indurate, separating from the exocarp, sometimes turning inside out to expose the seeds. Seeds depressed, oblong, disciform, or ellipsoid, apiculate in the middle; testa black, alveolate, usually enveloped in a spongy arillate cover that holds the seeds together in a regular packet even as they protrude from each locule after dehiscence. Panama, Colombia (Cauca, Valle), Venezuela, Guyana, northwestern Ecuador (Carchi, Esmeraldas); 4 species, 2 in Venezuela, both in the flora area. Epidryos is closely allied to Stegolepis and probably forms part of a paraphyletic assemblage in the tribe. Key to the Species of Epidryos 1. 1.

Spikelets 1–3, sessile .............................................................. E. guayanensis Spikelets 4–6, with unequal pedicels 1–4 cm long ............................. E. sp. A

Epidryos guayanensis Maguire, Mem. New York Bot. Gard. 12(3): 89. 1965. —Pareu-rei-yek (Arekuna). Fan-shaped epiphyte on tree trunks or upper branches, occasionally on granitic boulders; leaf sheaths 12–18 cm long, 2–3 cm wide, blade arching, to 100 cm long, 1.5–2.5 cm wide; peduncles filiform and drooping; spikelets 1–3, sessile, ca. 12 mm long, bracteoles 3–7 mm long; petals narrowly rhomboid-lanceolate, with a claw 4 mm long; seeds from the same locule adhering to each other in a longitudinal packet, each one disciform, 1.2–1.5 mm diameter, with a whitish aril. Moist montane forests, 900–1400 m; Bolívar (La Escalera near Salto El Danto, upper slopes of Sierra de Lema, headwaters of Río Cuyuní and Río Venamo). Guyana (Pakaraima Mountains). ◆Fig. 352. Epidryos sp. A Herb ca. 30 cm tall growing on open rocks

or boulders; leaf sheaths 22–28 cm long, 2 cm wide, the blade arching, to 1 m long, 1–1.5 cm wide; inflorescence with 4–6 pedicellate spikelets, the pedicels unequal, 1–4 cm long; petals lanceolate, 6–7 mm long, 3–4 mm wide at the base, then with a geniculate tip or claw ca. 3 mm long; seeds in 2 rows of 3 or 4 each per locule, lenticular, ca. 1.5 mm diameter, covered by a whitish, spongy-arillate layer. Large granitic dome with sparse vegetation, ca. 1500 m; Amazonas (Caño Piedra 115 km southeast of Puerto Ayacucho in Serranía Sipapo). Endemic. ◆Fig. 353. This interesting species is known from a single specimen (Sanoja et al. 2933, PORT). It is novel in Epidryos because of its pedicellate spikelets, but it agrees with the genus in other floral and vegetative features, such as the arching, slender scapes, the lanceolate petals, and the arillate seeds aligned in rows in the characteristically dehiscing capsule.

Epidryos 423

Fig. 352. Epidryos guayanensis

424

R APATEACEAE

Fig. 353. Epidryos sp. A

Guacamaya 425

Fig. 354. Guacamaya superba

426

R APATEACEAE

5. GUACAMAYA Maguire, Mem. New York Bot. Gard. 10(1): 35. 1958. Terrestrial herbs. Stem with leaf sheaths conduplicate and not much wider than the leaf blade, twisted to appear sigmoid in cross section; union of leaf sheath and blade abruptly transversal, blades linear or lanceolate. Peduncles axillary, inflorescence subpyramidal, solitary, subtended by 2 connate, sacciform involucral bracts. Spikelets numerous, sessile, spirally attached. Sepals free, red; petals white, lanceolate, imbricate, connate at the base, largely included within the sepals, the tips barely emergent at anthesis. Anthers basally 4-locular, 2-locular apically, terminally and transversely biporate. Ovary 3-locular, locules with ca. 8 ovules. Septal nectaries present. Seeds subprismatic, striate. Southwestern Venezuela and adjacent Colombia (Guainía); 1 species. Guacamaya superba Maguire, Mem. New York Bot. Gard. 10(1): 36, fig. 6, Ba–h. 1958. —Flor de Inírida real, Flor de Maroa. Herb 70–150 cm tall; base thick, with marcescent leaves, sheaths 15–30 cm long, 2–3 cm wide, blades 50–90 cm long, 2–3.5 cm wide; peduncle usually solitary, 90–130 cm tall, ca. 6 mm thick in the middle and widening to 12–18 mm wide at the top; head showy, with reddish bracteoles, cream-white sepals, and enclosed, white petals; bracts saccateconnate, 2.5–3 cm wide, 4–4.5 cm long, perforated by the spikelets as they expand; sepals 18–22 mm long, 5–6 mm wide; petals 12–14 mm long, tightly imbricate, connate at base

for 3–4 mm, then with a lanceolate-sagittate blade 8–9 mm long. Swampy, white-sand shrubby savannas, 50–200 m; Amazonas (upper Caño Cumare, outskirts of Maroa, uppermost Río Atabapo, Río Guainía, lower Río Guasacavi). Adjacent Colombia (Guainía). ◆Fig. 354. In Guacamaya superba, individual flowers open very briefly to expose the tips of the petals when they are visited by hummingbirds seeking the nectar produced near the base of the petals. Inflorescences of this species are collected locally in Venezuela and Colombia and used as floral bouquets, either when fresh and colorful or when dried as everlastings.

6. KUNHARDTIA Maguire, Mem. New York Bot. Gard. 10(1): 32, fig. 5, A–M. 1958. Large terrestrial herbs. Leaf blades sword-shaped, strongly equitant, 70–140 cm long, 3–6 cm wide. Inflorescence depressed-globose, large, many-flowered, with separate bivalved or else saccate involucral bracts. Spikelets sessile, reddish, porrect or slightly angled downward. Sepals red, stiff, free at the base; petals red, (broadly) lanceolate, strongly exserted, slightly zygomorphic, connate at the base. Anthers linear, 2-locular, opening by 2 annular terminal pores. Ovary 3-locular, pluriovulate. Septal nectaries present. Seeds subpyramidal, striate. Southern Venezuela; 2 species, both restricted to the flora area. Flowers of Kunhardtia remain open for about two days, whereas their counterparts in Guacamaya and Schoenocephalium may open only fleetingly for a few hours. The flowers produce considerable amounts of nectar and were observed on the Sierra de Maigualida to be regularly visited by hummingbirds (Tepui goldenthroats, Polytmus milleri). Even though the anthers are poricidally dehiscent, they are smooth-walled and have a sensitive trigger device that quickly expels the pollen onto the bird’s bill when probing for nectar. The two species treated below are very similar and may need to be combined in the future under Kunhardtia rhodantha. Kunhardtia is primarily a highland species that is unusual in growing on both quartzite and granitic substrates. It is known from just a few locations in the lowlands and intermediate elevations.

Kunhardtia 427

Less common form with fused saccate bracts

Fig. 355. Kunhardtia rhodantha

428

R APATEACEAE

Key to the Species of Kunhardtia 1. 1.

Heads nearly spherical, less compressed; petals 3–4 mm wide; occurring around 200 m .............................................................................. K. radiata Heads compressed-globose, not spherical; petals 6–8 mm wide; occurring at (600–)1500–2000 m ........................................................... K. rhodantha

Kunhardtia radiata Maguire & Steyerm., Acta Amazon. 9: 267. 1979. In clumps on seepages on granitic outcrops, 50–200 m; Amazonas (south of Puerto Ayacucho at Tobogán de la Selva and Caño Fluta). Endemic. Kunhardtia rhodantha Maguire, Mem. New York Bot. Gard. 10(1): 32, fig. 5a–m, 6Aa–Ah. 1958. Schoenocephalium sipapoanum Maguire, Acta Bot. Venez. 14(3): 17. 1984.

Sandstone and granitic outcrops, 600– 2100 m; Bolívar (Serranía Uasadi, Sierra de Maigualida), Amazonas (Cerro Autana, Cerro Cuao, Cerro Guanay, Cerro Sipapo, Cerro Yutajé). Endemic. ◆Fig. 355. There are two forms of Kunhardtia rhodantha known, the more common smallbracteate one, and a rarer from with connate, saccate bracts (both pictured in Fig. 355). Both forms occur together or closeby on Cerro Guanay and Cerro Yutajé.

Fig. 356. Marahuacea schomburgkii

Monotrema 429

7. MARAHUACAEA Maguire, Acta Bot. Venez. 14(3): 16. 1984. Perennial, thick-stemmed herbs mostly 1–1.5 m tall. Leaves distichous, sheaths conduplicate, coriaceous, ca. 15 cm long, 4–5 cm wide, auriculate and scariose-margined; blades rotated at right angle to sheaths, sword-shaped, 60–70 cm long, 3–5 cm wide. Peduncles axillary, usually 5 or 6 per leaf axil, 50–70 cm long, 2–3 mm diameter; inflorescence capitate, compressed, 3 cm wide, 2 cm tall, severalflowered, subtended by 3 or 4 orbicular, hyaline and broadly depressed bracts, the outermost ones strongly conduplicate, the inner ones successively smaller. Flowers in 3 or 4 sessile glomerules, each glomerule usually 3- or 4-flowered and subtended by a bract as in the spikelets; spikelets 25–28 mm long, bracteoles ca. 15, gradate, to 15 mm long, apically obtuse-cucullate. Sepals 20–25 mm long, connate at the base; petals yellow, broadly obovate-suborbicular, ca. 30 mm long. Anthers narrowly linear, strongly corrugated, ca. 10 mm long, 4-locular, dehiscent by a ventral, subterminal pore. Ovary 3-locular, ovules axillary, numerous; style subulate, ca. 20 mm long; stigma minute. Capsule oblong, basally membranous, apically hardened. Venezuela (endemic to Cerro Marahuaka, Amazonas); 1 species. Marahuacaea schomburgkii (Maguire) Maguire, Acta Bot. Venez. 14(3): 16. 1984. —Amphiphyllum schomburgkii Maguire, Mem. New York Bot. Gard. 10(1): 30. 1958.

Herb 1–1.5 m tall, the base mucilaginous; bracts pale. Locally dominant in exposed, rocky areas, 2400–2800 m; Amazonas (summit of Cerro Marahuaka). Endemic. ◆Fig. 356.

8. MONOTREMA Körn., Linnaea 37: 475. 1872. Terrestrial herbs. Leaves narrowly linear or gramineous. Peduncles axillary, 1 or 2 per leaf axil; inflorescence small, subglobose, bracts several, imbricate, unequal, the spikelets subtended by a secondary bract. Petals yellow or whitish, lanceolate. Anthers basally 4-locular, dehiscing by a single, circular, obliquely apical pore. Ovary pyriform, 3-lobed, 3-locular, the locules uniovulate, ovules unilaterally attached. Seeds ovoid, white, finely muriculate, with a prominent flattened apical appendage. Southeastern Colombia, southern Venezuela, Brazil (Amazonas); 4 species, 3 of these and an interspecific hybrid in Venezuela, all of these in the flora area. The fourth species in the genus, Monotrema arthrophyllum (Seub.) Maguire, is known from the western edge of the Guayana Shield in Colombia (Araracuara, Mitú, Serranía Chiribiquete, Yapobodá) and may prove to be conspecific with M. aemulans. Key to the Species of Monotrema 1. 1. 2(1).

2.

Outermost bracts of the inflorescence usually longer than the heads; floral bracts white or greenish or greenish yellow ................................... 2 Outermost bracts of the inflorescence shorter to slightly longer than the heads; floral bracts dark brown, maroon, or green .............................. 3 Outermost inflorescence bracts oval or ovate-lanceolate, 15–20 mm long, 6–10 mm wide, greenish; bracteoles acute or subacute, ovate or oblongovate, shorter than or equal to the spikelet; leaves 6–16 mm wide ..................................................................................................... M. ×affine Outermost inflorescence bracts narrowly lanceolate, aristate-acuminate to long acuminate, pale white, 20–35 mm long, 2–6 mm wide;

430

3(1).

3.

R APATEACEAE

bracteoles awl-shaped or attenuate-acuminate, narrowly lanceolate, longer than the spikelet; leaves 2–12 mm wide ................. M. bracteatum Heads ovoid, broadly conic, or subhemispherical, usually slightly to clearly longer than broad, sometimes as broad as long, (9–)10–20 mm long, (9–)10–13 mm broad; outer bracts closely imbricate, concealing the spikelets; bracteoles obtuse to rounded at apex; heads 15–25-flowered; floral bracts usually dark brown or maroon ................ M. xyridoides Heads usually broader than long, sometimes appearing to have 2 tightly appressed hemispherical segments, 8–15 mm long, 14–25 mm broad; outer bracts not closely imbricate and not concealing the spikelets; bracteoles subacute, apiculate; heads 40–75-flowered; floral bracts mainly greenish ...................................................................... M. aemulans

Monotrema aemulans Körn., Linnaea 37: 477. 1872. —Stegolepis aemulans (Körn.) Benth. & Hook., Gen. Pl. 3: 859. 1883. Herb 40–100 cm tall; leaf sheaths 10–12 mm wide, blades 60–80 cm long, 1.2–1.4 cm wide, the apex falcately acuminate; peduncles commonly 2–4 per axil, 80–100 cm tall; heads hemispheric, often subdidymous (divided into paired segments), 14–25 mm wide; petals oblanceolate, 4–5 mm long. White-sand savannas and sandy open shrubland, 50–600 m; Amazonas (Caño San Miguel, Cerro Vinilla, base of Cerro Yapacana, near Guarinuma, Río Atabapo, upper Río Autana, Río Baria, Río Casiquiare, Río Negro, upper Río Orinoco, Río Pacimoni). Colombia (Guainía, Vaupés), Guyana, Brazil (Amazonas, Mato Grosso). ◆Fig. 357. Monotrema ×affine Maguire, Mem. New York Bot. Gard. 10(1): 48, fig. 7M. 1958, pro. sp. Cespitose herb 40–70 cm tall; leaves 30– 50 cm long, 0.6–1.6 cm wide; bracteoles white, petals pale yellow to white. Whitesand savannas and shrubby savannas, 50– 200 m; Amazonas (Caño Cumare southeast of San Fernando de Atapabo, base of Cerro Yapacana). Endemic. ◆Fig. 358. Monotrema ×affine is found only in mixed populations of Monotrema bracteatum and M. xyridioides and represents hybrids between these two species. It has intermediate floral and foliar characters. Monotrema bracteatum Maguire, Mem. New York Bot. Gard. 10(1): 47, fig. 7G–L. 1958. Cespitose herb 30–60 cm tall; leaf sheaths 6–10 × 1–1.5 cm, blades 20–35 cm long, 0.2–

1.2 cm wide, old ones marcescent; outer bracts 20–35 mm long, 2–6 mm wide, spikelets 8–10 mm long; petals yellow, the basal claw ca. 3 mm long, the blade oblong-lanceolate, 4–5 mm long, 1.5–2 mm wide. Edges of flooded white-sand savannas, 50–500 m. Colombia, Venezuela; 2 varieties, both in the flora area. Key to the Subspecies of M. bracteatum 1. Leaves 2–4 mm wide; outer bracts of the inflorescence 2–5 mm wide ..................... .............................. subsp. bracteatum 1. Leaves 6–12 mm wide; outer bracts of the inflorescence 6–12 mm wide ................... ....................................... subsp. major M. bracteatum subsp. bracteatum White-sand savannas, forest edges, 50– 500 m; Bolívar (base of Cerro Guaiquinima, Río Trueno, Urimán), Amazonas (base of Cerro Yapacana, Río Puruname, lower Río Ventuari). ◆Fig. 360. M. bracteatum subsp. major Maguire in Luces & Steyerm., Fl. Venez. 11(2): 175. 1982. Seasonally flooded white-sand savannas, 50–200 m; Amazonas (Río Atabapo basin). Adjacent Colombia (Guainía). Monotrema xyridoides Gleason, Bull. Torrey Bot. Club 58: 332, pl. 25, fig. B. 1931. Perennial herb 50–70 cm tall; leaf sheaths linear-lanceolate, equitant, 8–12 cm long, blades 30–50 cm long, 0.6–0.8 cm wide, prominently veined; heads ovoid-globose, 12– 18 mm long, the outermost bract ca. 15 mm long; spikelets 8–9 mm long, petals with claw

Monotrema 431

ca. 5 mm long, the blade narrowly oblong, 3– 4 mm long. Open, moist white-sand savannas, ca. 50–800 m; Amazonas (Caño Yagua, Cerro Aracamuni, Río Atabapo and tributaries, Río Guayapo, Río Guianía, Río Manapiare basin, upper Río Orinoco, Río Sipapo, Río Ventuari). Colombia (Caquetá, Guainía, Vaupés), Brazil (Amazonas, Roraima). ◆Fig. 359.

Fig. 357. Monotrema aemulans

Fig. 358. Monotrema ×affine

432

R APATEACEAE

Fig. 359. Monotrema xyridoides

Fig. 360. Monotrema bracteatum subsp. bracteatum

Phelpsiella 433

9. PHELPSIELLA Maguire, Mem. New York Bot. Gard. 10(1): 28. 1958. Small herbs to 1 m tall, caudex fleshy. Leaves linear, equitant; sheaths 18–25 cm long, 1–1.5 cm wide, linear-lanceolate, marcescent, distichous, scariose-margined, auricles prominent, erect, ovate, 7–8 mm long and wide, oblique at the base; blade linear, subcoriaceous, 40–45 cm long, 0.8–1.6 cm wide, prominently 11–13veined. Peduncle solitary, strongly compressed, 50–70 cm long, 3–5 mm wide, broadly winged, ca. 12 mm wide below the inflorescence; involucral bracts 2, spathate, orbicular-obovate, 12–20 mm long, 12–14 mm wide, 5–7-veined, erect, appressed, margins initially connate. Spikelets 2(–4), sessile; bracteoles 13 or 14, gradate. Petals 20–25 mm long overall, the blade oblanceolate-rounded, 4–5 mm wide, with a basal claw 3–4 mm long. Anthers 2-locular, smooth, 2-porate, pores rounded, apical. Ovary 3-locular, pluriovulate, ovules attached in median part of axis. Seeds subprismatic, 1.5–3 mm wide, 1.7–2 mm long, testa scariose. Venezeula (apparently endemic to the summit of Cerro Parú, Amazonas); 1 species. Phelpsiella ptericaulis Maguire, Mem. New York Bot. Gard. 10(1): 29. 1958. Flowers with petals reported as either yellow or white; leaves bluish-tinged, apparently spreading by basal offshoots. Open

tepui summit meadows, 1300–2000 m; Amazonas (Cerro Parú). Endemic. ◆Fig. 361. Phelpsiella ptericaulis is very similar to Stegolepis breweri, except for the presence of spathate bracts.

Fig. 361. Phelpsiella ptericaulis

434

R APATEACEAE

10. RAPATEA Aubl., Hist. Pl. Guiane 305, pl. 118. 1775. Mnasium Schreb., Gen. Pl. 214. 1789. Terrestrial herbs. Leaf sheaths equitant, conduplicate, attached in a sigmoid arrangement. Inflorescence subtended by 2 large, opposing bracts, closely subtending the spheroid, elongated or compressed head; flowers sessile or short-pedicellate. Anthers introrse, 4-locular, with an apical hooded appendage, dehiscing by a single apical pore below the hood. Ovary 3-locular, each with a single basal ovule, normally 2 or sometimes just 1 aborting. Capsule hardened above the middle, dehiscence loculicidal. Seed usually solitary, oblongoid, finely and longitudinally striate. Panama, Colombia, Venezuela, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 23 species, 11 in Venezuela, all in the flora area. Key to the Species of Rapatea 1. 1. 2(1).

2. 3(2).

3.

4(3).

4.

5(4).

5.

6(1).

6.

Leaf blades 0.5–2 cm wide ......................................................................... 2 Leaf blades 2–12 cm wide .......................................................................... 6 Base of leaf blades abruptly widened at the base above the petiole; apex of leaf blade abruptly narrowed; petiole as long as the blade .................................................................................................... R. longipes Base of leaf blades gradually passing into the petiole; apex of leaf blades not abruptly narrowed; petiole shorter than the blade ....................... 3 Attenuate portion of the involucral bract 10–30 mm long, < twice as long as the expanded broader basal portion; lower surface of the leaf blade often glandular-punctate ........................................................ R. spruceana Attenuate portion of the involucral bracts 20–80 mm long, 2–4 times longer than the expanded broader basal portion; lower surface of the leaf blade not glandular-punctate ......................................................... 4 Attenuate portion of the involucral bracts 20–35 mm long, 2–3.5 times longer than the expanded broader basal portion; lower surface of the leaf blade densely and minutely papillate; bracteoles aristate .................................................................................................. R. yapacana Attenuate portion of the involucral bracts 35–80 mm long, 4–5 times longer than the expanded basal portion; lower surface of the leaf blade not minutely papillate; bracteoles acute, apiculate, or sharply acuminate ......................................................................................................... 5 Expanded basal portion of involucral bracts 7–12 mm broad; attenuate portion of involucral bracts 35–50 mm long, 4–5 mm broad; leaf blades 3–7 mm wide; peduncle 4–20 cm long, shorter than the leaves; plants growing in dense circular clumps of numerous individuals .............................................................................................. R. angustifolia Expanded basal portion of involucral bracts 20–25 mm broad; attenuate portion of involucral bracts 55–85 mm long, 9–14 mm broad; leaf blades 9–15 mm wide; peduncle 35–45 cm long, longer than or equal to the leaves; plants generally growing as solitary individuals ...... R. circasiana Leaf sheath short, 3–9(–15) cm long; leaf blades 2–3.5(–5) cm wide; involucral bracts 1.5–3 cm long, narrowed apical part 5–10 mm long; heads small, 1–2 cm broad ........................................................ R. longipes Leaf sheath 15–25 cm long; leaf blades (2–)4–11.5 cm wide; involucral bracts 5–25 cm long, apical attenuate portion 3–22 cm long; heads 3– 8 cm broad .............................................................................................. 7

Rapatea 435

7(6). 7.

Base of leaf blades gradually narrowed to the sheath ............................. 8 Base of leaf blades abruptly constricted above the summit of the sheath ................................................................................................................ 9 8(7). Bractlets of the spikelets acuminate; leaves smooth throughout; involucral bracts cordate; ranging throughout the Venezuelan Guayana ................................................................................................... R. paludosa 8. Bractlets of the spikelets obtuse; lower surface of the leaves slightly scabridulous; involucral bracts rounded or truncate at base; Cerro Sipapo, Amazonas ......................................................................... R. scabra 9(7). Peduncle 3–14 cm long; involucral bracts broadly deltoid, nearly as broad as long, 6–10 cm long ........................................................................... 10 9. Peduncle 20–50 cm long; involucral bracts lanceolate, longer than broad, 10–25 cm long ...................................................................................... 11 10(9). Lower surface of leaf sheath densely brown-pubescent with elongate, crisp-villosulous, loose, multicellular hairs; upper surface of leaf with short, divaricate, pointed hairs irregularly scattered on the veins in addition to the minutely papillate surface; spikelets 16–20 mm long (excluding the sepals); lowest bracteoles 12–13 mm long; anthers 5 mm long; filaments glabrous ..................................................... R. steyermarkii 10. Lower surface of leaf sheath brown-dotted between smaller, pale punctuations; upper leaf surface densely verruculose-papillate throughout; spikelets 23–25 mm long (excluding the sepals); lowest bracteoles 17– 18 mm long; anthers 10 mm long; filaments ciliolate ..... R. aracumuniana 11(9). Bracteoles stiff, obviously different in size, the lower ones much less than 1/2 the length of the spikelet; leaf blades 35–75 cm long, 8–11 cm wide; involucral bracts 10–20 cm long; lower leaf surface with the primary veins acutely elevated, separating the broad, flat depressions; primary veins of upper leaf surface shallowly raised, not presenting a plaited appearance, 0.5–1 mm apart .................................................. R. fanshawei 11. Bracteoles submembranous, of nearly equal length; leaf blades ca. 130 cm long, 9–12 cm wide; involucral bracts ca. 25 cm long; lower leaf surface with shallowly convex surfaces separated by shallow sulcations, lacking acutely elevated primary veins; primary veins of upper surface conspicuously elevated and presenting a plaited appearance, 1–2 mm apart ................................................................................... R. chimantensis Rapatea angustifolia Spruce ex Körn., Linnaea 37: 469. 1872. Herb 15–60 cm tall, in dense circular clumps of many individual plants; peduncle 4–20 cm tall. Depressions in riparian granitic outcrops and on berms of small black-water rivers, 50–200 m; Amazonas (Caño San Miguel, streams along road from Maroa to Yavita, Río Casiquiare, Río Guainía, Río Pacimoni, Río Temi). Endemic. ◆Fig. 366. Rapatea aracamuniana Steyerm., Ann. Missouri Bot. Gard. 76: 964. 1989. Herb with leaves 35–65 cm long; flowering head 4.5–6.5 cm wide, 8–12.5 cm long. Along

rapids over granitic outcrop, ca. 600 m; Amazonas (Cerro Aracamuni). Endemic. Rapatea chimantensis Steyerm., Ann. Missouri Bot. Gard. 75: 313, fig. 2. 1988. Herb 1–2 m tall. Along stream, ca. 1000 m; Bolívar (Macizo del Chimantá [southwestern slopes of Torono-tepui]). Endemic. ◆Fig. 364. Rapatea circasiana García-Barr. & L.E. Mora, Mutisia 22: 9. 1954. Herb 30–60 cm tall; bracts lanceolate, (3–) 5–10 cm long. Wet sandy savannas and open forests, 50–200 m; Amazonas (Caño Cumare, Río Atabapo, Río Guainía, Río Sipapo). Co-

436

R APATEACEAE

lombia (Vaupés), Brazil (northern Amazonas). ◆Fig. 367. Rapatea fanshawei Maguire, Mem. New York Bot. Gard. 12(3): 96, fig. 14H–M. 1965. Herb ca. 1 m tall. Forest edges on sandy substrates, 400–1000 m; Bolívar (Cordillera Epicara, near El Paují, Macizo del Chimantá, Sierra Pakaraima). Guyana. ◆Fig. 363. Rapatea longipes Spruce ex Körn., Linnaea 37: 472. 1872. Rapatea modesta Maguire, Bot. Mus. Leafl. 14: 112. 1950. —Rapatea longipes var. modesta (Maguire) Maguire, Mem. New York Bot. Gard. 12(3): 100. 1965. Herb 30–80 cm tall; petiole long, terete, narrowly oblong leaves, bracts wider than long. Swampy, sandy savannas and low open forests, 50–200 m; Amazonas (Maroa, Río

Atacavi, San Carlos de Río Negro). Southeastern Colombia, Brazil (Amazonas). ◆Fig. 365. Rapatea paludosa Aubl., Hist. Pl. Guiane 305, pl. 118. 1775. —Mnasium paludosum (Aubl.) Willd., Sp. Pl. 2(1): 22. 1798 [1799]. Rapatea schultesiana García-Barr. & L.E. Mora, Mutisia 22: 13. 1954. Herb 0.8–1.5 m tall; petals pale yellow. Panama, Colombia, Venezuela, Trinidad, Guyana, Suriname, French Guiana, Peru, Brazil; 2 varieties, 1 in Venezuela. Variety sessiliflora is known only from Suriname.

Fig. 362. Rapatea steyermarkii

Rapatea 437

Fig. 363. Rapatea fanshawei

Fig. 364. Rapatea chimantensis

438

R APATEACEAE

Fig. 365. Rapatea longipes

Fig. 366. Rapatea angustifolia

Rapatea 439

Fig. 368. Rapatea spruceana Fig. 367. Rapatea circasiana

440

R APATEACEAE

Fig. 369. Rapatea paludosa var. paludosa

Saxofridericia 441

R. paludosa var. paludosa Swampy ground in forests and along streams, 50–500(–1500) m; widespread in Delta Amacuro, Bolívar, and Amazonas. Panama, Colombia (Valle and southeast), Trinidad, Guyana, Suriname, French Guiana, Peru, Brazil (Amazon basin, Bahia). ◆Fig. 369. This is the most common and widespread species of Rapatea.

Moriche, La Esmeralda, Río Atabapo and tributaries, Río Autana, Río Casiquiare, Río Guianía, Río Guayapo, Río Negro, Río Sipapo, Río Yatúa basin). Adjacent Colombia (Guianía), Brazil (Amazonas). ◆Fig. 368. Rapatea spruceana is similar to R. longipes but has longer bracts and the petiole less abruptly transitioning with the blade.

Rapatea scabra Maguire, Mem. New York Bot. Gard. 12(3): 93. 1965. Herb with scabrous, linear leaves to 1.5 m long; bracteoles obtuse and hooded, yellow. Along streams on granitic outcrops, edges of streams, ca. 600 m; Amazonas (Cerro Sipapo). Endemic.

Rapatea steyermarkii Maguire, Fieldiana, Bot. 28: 130. 1951. Herb 20–90 cm tall; peduncle short, 3–5 cm long. In forest understory in moist or swampy areas, 100–1000(–1700) m; Bolívar (slopes of Carrao-tepui and Sororopán-tepui, Gran Sabana, lower slopes of Ilú-tepui, Sierra de Lema). Adjacent Guyana. ◆Fig. 362.

Rapatea spruceana Körn., Linnaea 37: 470. 1872. Herb 30–80 cm tall. Sandy, flooded whitesand savannas and forests, 50–600 m; Amazonas (Cerro Aracamuni, base of Cerro

Rapatea yapacana Maguire, Mem. New York Bot. Gard. 12(3):101. 1965. Herb 25–40 cm tall. In dense hummocks in open forests and savannas, 50–200 m; Amazonas (Yapacana savannas). Endemic.

11. SAXOFRIDERICIA R.H. Schomb., Rapatea 13. 1845. Robust perennial herbs. Leaves distichous, petiolate or not. Peduncles solitary, axillary; inflorescence globose or depressed-globose to hemispheric at maturity, completely enclosed by saccate involucral bracts (in some cases evidently formed by 2 connate, foliaceous bracts, in other cases appearing as a single enveloping membrane), in bud the bracts filled with copious mucilage, at anthesis perforated by the elongating sepal tips, sometimes disintegrating as the head matures, or with a persistent acuminate tip extending beyond the head. Spikelets 35–ca. 100 per head, sessile or shortly pedicellate, with numerous bracteoles, these either gradate and scale-like or else subequal in length and ± spathulate-hooded with a shortly acute tip. Sepals lanceolate, imbricate, free to the base, ± inrolled with a firm-hardened keel, scarious margins, and a sharp, stiff tip. Petals basally connate into a narrow tube, then with separate claws leading into the yellow, obovate or suborbicular blades, which are strongly spreading at anthesis and remaining fresh for about one day. Filaments adnate to the basally connate portion of the corolla and shortly free above; anthers lanceolate, usually transversely wrinkled, tapered distally, inconspicuously 4-lobed, the 2 anterior locules ending in a bilobed terminal pore, the posterior locules extending shortly beyond the pore as a slightly hooded appendage. Ovary 3-locular, with 2–4 axile ovules per locule; style slender, with a minute terminal stigma. Capsule oblongoid, membranous below, hardened above, the valves spreading open loculicidally at dehiscence. Seeds oblongoid-reniform, with a centrally depressed hilum, transversely striate and furrowed, usually only one but occasionally 2 or 3 per capsule, (no more than one seed developing per locule, and many fruits without any seed developing). Southeastern Colombia, southern Venezuela, Guyana, Suriname, French Guiana, Amazonian Brazil; 11 species, 9 in Venezuela, all in the flora area.

442

R APATEACEAE

Key to the Species of Saxofridericia 1. 1. 2(1).

2.

3(2). 3. 4(2). 4. 5(4).

5.

6(4). 6. 7(6).

7.

8(6).

8.

Margins of the lower part of the leaf blade, petiole (when present), and the upper keel of the leaf sheath with spines or prickles ........ S. aculeata Margins of the leaf blade, petiole, and leaf sheath entire, never spinose ................................................................................................................ 2 Mature inflorescence 1.5–3.5 cm wide; leaf blades 1–3.5 cm wide; lowland plants growing in seasonally flooded banks of black-water rivers and streams ................................................................................................... 3 Mature inflorescence 4–8 cm wide; leaf blades mostly > 3.5 cm wide; montane or rarely lowland plants growing in bogs, forests, or on rocks, typically not riparian plants ........................................................................ 4 Petiole shorter than the blade or essentially lacking; leaf blades 1.4– 3.5 cm wide ................................................................................. S. inermis Petiole 1–2 times the length of the blade; leaf blades 1–2 cm wide ................................................................................................... S. petiolata Petiole present, 5–70 cm long; known only from Cerro Parú .................. 5 Petiole absent or occasionally short-petiolate, but then the leaves < 4 cm wide; more widespread .......................................................................... 6 Leaf sheaths 1.5–3 cm wide; base of the blades strongly unequal, one side offset up to 2.5 cm above the other side; peduncle 10–15 mm wide at the apex; inflorescence spherical, with hyaline bracts tightly appressed to the head, even at the apex, < 7 cm long; spikelets ca. 100, the bracteoles all of the same length .................................................... S. sp. A Leaf sheaths 5–7 cm wide; base of the blades with subequal sides; peduncle widened to 20–25 mm wide at the apex; inflorescence hemispherical, wider at the bottom, with firm, membranous bracts 7–10 cm long extending into an acuminate tip beyond the head; spikelets 50– 80, the bracteoles gradate, of unequal lengths ......................... S. grandis Peduncles abruptly enlarged and compressed below the inflorescence, 25–40 mm wide at the top ..................................................................... 7 Peduncles gradually enlarged below the inflorescence, 15–25 mm wide at the top .................................................................................................... 8 Top of peduncles strongly and sharply compressed, the margins usually narrowly winged; leaf sheaths not thick-spongy, 4–5 cm wide; leaf blades 4–5 cm wide; spikelets 20–24 mm long ...................... S. compressa Top of peduncles moderately compressed, without winged margins; leaf sheaths thick-spongy, 5–11 cm wide; leaf blades 5–10 cm wide; spikelets 26–32 mm long .................................................................. S. spongiosa Secondary bracts usually present inside the outermost spathaceous bracts, the outer ones lanceolate and to 4–5 cm long, decreasing in size inwardly; sepals 17–20 mm long; involucral bracts 8–14 cm long; Bolívar ........................................................................................... S. regalis Secondary bracts lacking or inconspicuous within the outer spathaceous bracts; sepals 15–16 mm long; involucral bracts 3–8(–12) cm long; Amazonas ...................................................................................... S. duidae

Saxofridericia 443

Saxofridericia aculeata Körn., Linnaea 37: 457. 1872. Saxofridericia subcordata Körn., Linnaea 37: 459. 1872. Saxofridericia pandanoides Linden & André, Ill. Hort. 20: 213, t. 153, 154. 1873. Saxofridericia australis Gleason, Bull. Torrey Bot. Club 60: 350. 1933. Herb 1–1.5 m tall; leaf sheaths 10–40 × 4– 10 cm, petiole usually present and 5–15(–50) cm long, blades 1–1.5 m long, 4–10 cm wide, usually abruptly widened above the petiole, spines or prickles present along the petiole, upper keel of the leaf sheath, and lower margins of the blade; peduncle 15–60 cm tall, widened to 6–10 mm wide at the apex; inflorescence globose, 2.5–4.5 diameter, the bracts hyaline and tightly enveloping the head, 4–6 cm long; spikelets 75–100, 10–15 mm long, with 28–30 subequal bracteoles; petals with an emergent, orbicular blade at anthesis. Not known yet from the flora area, but expected there, at least in southern lowland Amazonas, because it is known from the upper Rio Negro of Brazil. Colombia (Vaupés), Guyana, French Guiana, Suriname, Brazil (Amazonas, Pará). Saxofridericia compressa Maguire, Mem. New York Bot. Gard. 10(1): 25. 1958. Herb 1.5–3 m tall; leaf sheaths 30–40 × 4– 5 cm, blades 1–2 m long, 4–5 cm wide; peduncle expanded and strongly flattened and ± winged at the apex; involucral bracts 6–10 cm long, head 6–7.5 cm wide, spikelets 20–24 mm long, bracteoles 17–20, gradate; petals with claw 8–10 mm long, the blade broadly obcordate, 25–30 mm wide, ca. 20 mm long. Swampy scrub, along steep creeks, bogs, open Bonnetia and Tyleria woodlands, (500–) 1500–2300; Amazonas (Cerro Aracamuni, Sierra de la Neblina, Sierra Parima). Adjacent Brazil (Amazonas). ◆Fig. 371. Saxofridericia duidae Maguire, Mem. New York Bot. Gard. 10(1): 24. 1958. Saxofridericia duidae subsp. marahuacensis Maguire, Mem. New York Bot. Gard. 10(1): 25. 1958, syn. nov. Herb 1.5–3 m tall; leaf sheaths 20–30 cm long, 3 cm wide, blades 80–180 long, 2–3.5

cm wide; peduncle dilated near the apex, where 15–18 mm wide; inflorescence globose, 4.5–5.5 cm diameter, involucral bracts 3–8 (–12) cm long; spikelets 18–23 mm long; petals with obovate, apiculate blade. Tepui slope forests and summit meadows, 600–2000 m; Amazonas (Cerro Duida, Cerro Huachamacari, Cerro Marahuaka). Endemic. ◆Fig. 372. Saxofridericia grandis Maguire, Mem. New York Bot. Gard. 10(1): 26, fig. 4A–D. 1958. Large herb 1–2 m tall; leaf sheaths 15–20 cm long, 5–7 cm wide, petiole present, blade linear, 70–100 cm long, 4–5.5 cm wide; peduncle gradually expanded to the apex; involucral bracts 7–10 cm long, inflorescence hemispherical, spikelets 25–30 mm long; sepals 22–26 mm long, petals with orbicular blade. Tepui meadows and scrub forests, 1100–2000 m; Amazonas (Cerro Parú). Endemic. ◆Fig. 374. Saxofridericia inermis Ducke, Arq. Inst. Biol. Veg. 4: 1. 1938. Herb 0.8–1.5 m tall; leaf sheaths 13–19 cm long, 1.5–3 cm wide, blades 50–100 cm long, 1–3.5 cm wide, subsucculent, with little or no petiole, the midrib thickened along with adjacent part of the blade, no veins evident; peduncles smooth and ± flattened, (20–) 60–100(–145) cm long, 7–11 mm wide below the head; inflorescence depressed-globose, 3– 3.5 cm wide, involucral bracts 3.5–5 cm long, the basalmost 5–8 mm of the involucre hardened and persistent; spikelets 40–70, sepals 9–11 mm long, petals yellow, ca. 20 mm long, the blade elliptic-obovate. Seasonally flooded forests, stream banks, palm-bamboo thickets on white sand, savannas and scrub on granitic outcrops, 50–300 m; Amazonas (junction of Río Casiquiare with Río Negro, Río Guianía, Río Pacimoni, Río Siapa). Colombia (Guainía), Brazil (Amazonas). ◆Fig. 373. There are a number of collections from Piedra Catipán, on the Río Yatúa in Amazonas, that are related to Saxofridericia inermis and may represent a new taxon or some kind of hybrid with another species. They appear to be more robust plants, with

444

R APATEACEAE

the leaf bases thick-spongy (drying rugulose). Also, the inflorescences are quite unusual; when the outer involucral bracts are removed, there appear to be several elongate axes from which the spikelets emerge. Saxofridericia petiolata Maguire, Mem. New York Bot. Gard. 10(1): 28, fig. 4M. 1958. Herb (0.7–)0.8–1.5 m tall; leaf sheaths 15– 20(–25) cm long, ca. 3 cm wide, petiole commonly 1–2 times longer than the blades, (25–) 35–100 cm long, spongy, gradually broadening into the blades, the blades (19–)23–60 cm long, 1–2 cm wide, subsucculent; peduncles 0.7–1.0 m long, 10–12 mm thick at the apex, ± flattened, spongy; inflorescence hemispheric to subglobose, 2.5–3.0 cm wide; involucral bracts with the lower 10 mm fused to the receptacle, above that becoming pierced by the spikelets, then extending to 15 mm beyond the head; spikelets 40–55 per head, 15–16 mm long; bracteoles 20–22; se-

pals 13–15 mm long; petals with a basal tube 10–12 mm long, the blade 9–11 mm long, 9 mm wide, apiculate at the apex for 1–2.5 mm. Sandy, flooded margins of black-water rivers, thickets on low granitic outcrops, 50– 200 m; Amazonas (Río Atacavi, Río Guasacavi, Río Temi). Endemic (almost certainly occurring in nearby Colombia).

Fig. 370. Saxofridericia regalis

Saxofridericia 445

Fig. 372. Saxofridericia duidae Fig. 371. Saxofridericia compressa

446

R APATEACEAE

Fig. 374. Saxofridericia grandis

Fig. 373. Saxofridericia inermis

Schoenocephalium 447

Saxofridericia regalis R.H. Schomb., Rapatea 14. 1845. Rapatea viscosa Gleason, Bull. Torrey Club 56: 18. 1929. Large herb 1–3 m tall; leaf blades 20–40 cm long, 3–4 cm wide, the blades 80–150 cm long, 3–5 cm wide; peduncle compressed and ca. 25 mm wide at the apex; inflorescence hemispheric, 4–5 cm broad, outer spathaceous bracts lanceolate-acuminate, 8–12 cm long, spikelets 80–100, the outer ones sometimes with secondary bracts to 5 cm long, spikelets 20–25 mm long; petals with basal narrow tube, the blade obovate-orbicular, ca. 12 mm long × 16 mm wide, apiculate. Whitesand meadows, tepui shrub thickets, 700– 1700 m; Bolívar (Cerro Guaiquinima, Cerro Jaua, Cerro Marutani, widespread in the Gran Sabana and on adjacent tepui slopes, Río Carrao at Isla Orquídea, Sierra Pakaraima). Guyana, Suriname (Tafelberg), Brazil (Roraima). ◆Fig. 370. Saxofridericia spongiosa Maguire, Mem. New York Bot. Gard. 10(1): 26, fig. 4E–L. 1958. Large herb 1–3 m tall; leaf sheaths 30–35 cm long, 5–11 cm wide and thick-spongy, becoming wrinkly when dried, blades 150–350 cm long, 5–10 cm wide; peduncle somewhat expanded toward the apex, 25–30 mm wide, inflorescence hemispheric, 5–8 cm wide,

bracts 15–30 cm long; spikelets 60–100, 26– 32 mm long; sepals 25–28 mm long, petals with claw 20–22 mm long, the blade rhomboid-orbicular, 18–20 mm wide. Granitic and sandstone outcrops, 100–1700 m; Amazonas (Cerro Aracamuni, Cerro Aratitiyope, Cerro Vinilla, Piedra Arauicaua, upper Río Siapa, Sierra Parima). Brazil (Amazonas, Roraima). Saxofridericia spongiosa prefers rocky habitats and has the widest altitudinal tolerance of any species in the genus. Saxofridericia sp. A Large herb 1.5–2 m tall; leaf sheaths 25– 35 cm long. 1.5–3 cm wide, petioles 6–30(–70) cm long, blades with strongly unequal base, 75–140 cm long, 3.5–8 cm wide; peduncle 1– 1.3 m tall, 10–15 mm wide below the head; inflorescence globose, 4–5 cm diameter, tightly sheathed by the fused hyaline bracts, spikelets ca. 100, bracteoles 46–50, subequal in length, 12–15 mm long; sepals 11–13 mm long, petals connate basally into a tube for 9– 10 mm, the blade orbicular-rhombical, 8–11 mm diameter. Open woodlands on tepui summits, 1000–1100 m; Amazonas (Cerro Parú). Endemic. This species grows in close proximity to Saxofridericia grandis, which is found in more open areas at the interface of savannas and tepui woodlands. It is based on Berry et al. 4982 (MO).

12. SCHOENOCEPHALIUM Seub. in Mart., Fl. Bras. 3(1): 130. 1847; emend. Maguire, Mem. New York Bot. Gard. 10(1): 37. 1958. Herbs with a short, subfleshy caudex. Leaf sheaths ± equitant, conduplicate, blades narrowly linear. Heads globose, bracts 2, eventually reflexed; spikelets radiate, bracteoles gradate. Sepals free to the base; petals lanceolate, not extending beyond the sepals, connate at the base. Filaments adnate to corolla tube; anthers 4locular, apex cruciform, 4-porate, sometimes the lateral pores confluent. Ovary 3locular, each locule 2-ovulate (4-ovulate in S. schultesii); styles terete, truncate. Septal nectaries present. Seeds subprismatic-subpyramidal, longitudinally striate. Endemic to the Guayana Shield in southeastern Colombia, southern Venezuela, northern Amazonian Brazil; 4 species, 2 in Venezuela, both in the flora area. The other two species in the genus are Schoenocephalium martianum Seub. from the Araracuara area in Colombia, and S. schultesii Maguire from Isibukuri area of Colombia. The Venezuelan species, and most likely all species in the genus, produce nectar, are visited by hummingbirds, and harbor hummingbird mites of the genus Proctolaelaps. Maguire [Mem. New York Bot. Gard. 10(1): 37. 1958] described the flowers of Schoenocephalium as cleistogamous, but this is incorrect. The sepals relax their grip on the petals for a brief period of time such that the petals spread open slightly and allow hummingbirds to reach the nectar surrounding the ovary.

448

R APATEACEAE

Fig. 376. Schoenocephalium teretifolium Fig. 375. Schoenocephalium cucullatum

Spathanthus 449

Since this only happens on 1 or 2 spikelets per head and for a few hours at a time, most of the time the inflorescences appear to have entirely closed flowers. The attractive floral heads of this genus are gathered locally for bouquets and are dried and used as everlastings. The common names refer to the areas where they occur, and in neighboring Colombia, they are well known around the regional capital of Puerto Inírida. In areas of close sympatry, such as near Santa Rosa and along Caño Cumare in the Río Atabapo basin, Schoenocephalium cucullatum and S. teretifolium hybridize and produce individuals with intermediate floral and foliar characters. Key to the Species of Schoenocephalium 1.

1.

Leaf blades plane, linear, 6–20 mm wide; spikelets mostly 30–50 per head, with sepals projecting 1–6 mm beyond the tips of the upper bracteoles .............................................................................. S. cucullatum Leaf blades terete, 1–3 mm diameter; spikelets mostly 15–25 per head, the sepals projecting 6–10 mm beyond the tips of the upper bracteoles ............................................................................................... S. teretifolium

Schoenocephalium cucullatum Maguire, Mem. New York Bot. Gard. 10(1): 39, fig. 6Ca–Ch. 1958. —Flor de Canaripó, Flor de Inírida, Flor del Atabapo. Schoenocephalium coriaceum Maguire, Mem. New York Bot. Gard. 10(1): 38. 1958. Herb 1–1.5 m tall; leaf sheaths marcescent, the peduncles flattened near the apex; spikelets either all reddish or reddish at the base and whitish apically, or sometimes yellow-white throughout; petals yellow (sometimes whitish at the apex), tightly imbricate, the blades lanceolate and 8–11 mm long. Seasonally flooded white-sand savannas, 50–200 m; Amazonas (Caño San Miguel, savannas surrounding Cerro Yapacana, near Maroa, Río Atabapo basin, Río Casiquiare tributaries, Río Guayapo, lower and middle Río Ventuari). Adjacent Colombia (Guianía), Brazil (scattered in Amazonas). ◆Fig. 375.

Schoenocephalium teretifolium Maguire, Mem. New York Bot. Gard. 10(1): 38, fig. 6Da–Dh. 1958. Herb ca. 1 m tall; leaves erect, the blades terete, 1–3 mm diameter; peduncles 70–120 cm tall, solitary or few; head spherical, 2–3 cm diameter, with numerous radiate spikelets, these basally red, apically cream and turning red with age; petals 8–10 mm long, the blade yellowish, tightly imbricate, 6 mm long, 2.5 mm wide. Wet sandy savannas, rock outcrops on tepui slopes, 50–200(–800) m; Amazonas (Caño Cumare, Caño Pimichín, Cerro Aracamuni, near mouth of Río Atacavi, savannas near Río Autana, lower Río Guasacavi, Río Pacimoni, Santa Cruz del Atabapo). Adjacent Colombia (Guainía). ◆Fig. 376. Hybrids of Schoenocephalium teretifolium with S. cucullatum occur in the savannas of Santa Cruz, near the mouth of the Río Atacavi, and along Caño Cumare, in the lower Río Atabapo basin.

13. SPATHANTHUS Desv., Ann. Sci. Nat. (Paris) 13: 45. 1828. Terrestrial herbs 0.5–3 m tall. Leaf sheaths distichous, blades linear. Peduncles 1–several, solitary in leaf axils. Involucral bract solitary, spathate, lanceolate; inflorescence spicate, adnate to the spathe. Spikelets sessile, numerous; bracteoles numerous, distichous. Petals yellow, lanceolate, ephemeral. Anthers 4locular, opening by a single, apical, unequally dentate pore. Ovary 3-locular, 2 carpels abortive, the other with a single ovule when mature; style centrally sub-basal. Capsule 1-locular, 2-valved, 1-seeded. Seed oblong-elliptic, striate, sub-basally attached.

450

R APATEACEAE

Fig. 377. Spathanthus bicolor

Fig. 378. Spathanthus unilateralis var. unilateralis

Stegolepis 451

Southeastern Colombia, southern Venezuela, Guyana, French Guyana, Amazonian Brazil; 2 species, both in the flora area. Key to the Species of Spathanthus 1.

1.

Involucral bract green or white basally; leaves 1.5–3.5 cm wide; bracteoles obtuse and apiculate at apex; mainly in Amazonas, rare in Bolívar ........................................................................................... S. bicolor Involucral bract lavender-pink with dark purple; leaves 5–10 cm wide; bracteoles acute to acuminate, shortly pointed; Bolívar .... S. unilateralis

Spathanthus bicolor Ducke, Arq. Inst. Biol. Veg. 2: 28. 1935. Herb 0.5–2 m tall; leaf blades linear, often firm-succulent, 50–100 cm long, 1.5–3.5 cm wide, the apex often abruptly and obliquely acuminate; petiole sometimes evident, to 30 cm long; spathe 8–18 cm long. Seasonally flooded stream or river banks, forest understories, usually on white-sand soils, 50–500 m; Bolívar (Canaima, Río Acanán), widespread in southern and western Amazonas. Apure; southeastern Colombia, Brazil (Amazonas). ◆Fig. 377. Spathanthus unilateralis (Rudge) Desv., Ann. Sci. Nat. (Paris) 13: 45, pl. 4, fig. 1.

1828. —Mnasium unilaterale Rudge, Pl. Guian. 12, pl. 11. 1805. —Rapatea unilateralis (Rudge) Schult. & Schult. f., Syst. Veg. 7(2): 1149. 1830. Herb 1.5–3 m tall; leaf blade 150–250 cm long, 6–10 cm wide; spathe to 25 cm long. Southern Venezuela, Guyana, Suriname, French Guiana, Amazonian Brazil; 3 varieties, 1 in Venezuela. S. unilateralis var. unilateralis Lowland swampy forests, 50–600 m; Bolívar (Canaima, base of La Escalera below Piedra de la Virgen, Reserva Forestal Imataca). Guyana, Suriname, French Guiana, Brazil. ◆Fig. 378.

14. STEGOLEPIS Klotzsch ex Körn., Linnaea 37: 480. 1872. Herbs, terrestrial or rarely epiphytic. Caudex short to elongate. Leaf sheaths equitant, conduplicate, often with a broad flattened aspect, blades linear-lanceolate, sword-shaped, or grass-like. Inflorescence lacking subtending spathaceous bracts (except in S. gleasoniana and S. perligulata, which are represented by vestigial narrow membranous scales or flaps); peduncles several, axillary; inflorescence 1–many-flowered, the spikelets in a fan-shaped or globose arrangement; bracteoles gradate. Sepals membranous at base, most often connate into a short tube, the limb hardened, exserted, sometimes becoming reflexed at anthesis; petals differentiated into claw and limb, the claws connate into a short tube, the limb lanceolate to orbicular or subcordate. Filaments free or usually connate at the base, the tube discrete or adnate to the corolla tube; anthers 4-locular, dehiscing subterminally by a single or double pore, caudate at the base, the posterior lobes prolonged, ± hooded, transversely corrugated at least in lower 1/2–2/3. Ovary 3-locular, loculicidal; locules pluriovulate; placentation axile. Seed prismatic or ± pyramidal; testa alveolate or ± striate, usually pale. Endemic to the Guayana Shield in southern Venezuela, Guyana, and northern Brazil; 34 species, 31 in Venezuela, all in the flora area. The three species not found in Venezuela are Stegolepis piresii Maguire, which is endemic to Serra Aracá (Amazonas, Brazil), S. ferruginea J.G. Baker, from the Kaieteur plateau of Guyana, and an undescribed species from the summit of Mount Ayanganna in Guyana.

452

R APATEACEAE

Key to the Species of Stegolepis 1.

1. 2(1). 2. 3(2).

3.

4(3). 4. 5(4). 5. 6(5).

6. 7(6).

7.

8(7).

8.

9(7).

Peduncle 8–10 mm wide throughout its length, flattened, winged, ribbonlike; Bolívar (in and around sink-holes on Cerro Sarisariñama) ...................................................................................................... S. breweri Peduncle not as above, the surface angled or costate, but not winged; elsewhere in the Venezuelan Guayana ....................................................... 2 Inflorescence ± globose, many-sided, the spikelets oriented in various angles or directions, generally numerous (15–100) ............................. 3 Inflorescence compressed, 2-sided, and radiating fanwise (few to 50 flowers per inflorescence), or else composed of just 1–8 spikelets ........... 13 Stems with an enlarged, bulbous, onion-like base; leaves neither equitant nor fan-like; spikelets obliquely attached to the apex of the peduncle, somewhat inclined ............................................................. S. microcephala Stems not as above; leaves in an equitant or fan-like plane, the leaf sheaths conduplicate with a flattened, broad aspect; spikelets not obliquely attached at the apex of the peduncle, nor inclined .................. 4 Petals white, 5.5–7 mm long; peduncle slender, 0.8–1.5 mm diameter .................................................................................................... S. albiflora Petals yellow or orange, 8–40 mm long; peduncle usually stouter (except S. choripetala) ........................................................................................ 5 Inflorescence with 15–50 spikelets ........................................................... 6 Inflorescence densely flowered with > 50 spikelets ............................... 10 Bracteoles small, suborbicular to ovate, 3–7 mm long; inflorescences sometimes viviparous, with bulbils emerging from the spikelets .................................................................................................... S. vivipara At least some bracteoles longer or more acute than above; inflorescences never viviparous .................................................................................... 7 Inflorescence 1.5–3 cm diameter; spikelets 2–5 mm diameter, bracteoles 1.5–3 mm wide; peduncle slightly or scarcely dilated at apex to 2– 4 mm wide .............................................................................................. 8 Inflorescence 3–4.5(–5) cm diameter; mature spikelets 4–9 mm diameter, bracteoles 3–5 mm wide; peduncles usually conspicuously dilated at apex to 5–10 mm wide ........................................................................... 9 Leaf blades 3.5–5 cm wide; midvein not sulcate on adaxial surface; main veins on abaxial surface 15–25 on each side of midvein, 0.5–1 mm apart; leaf sheath veinless; bracteoles 12–14, 3 mm wide; inflorescence with 12–30 spikelets ............................................................. S. choripetala Leaf blades 2–3.5 cm wide; midvein prominently sulcate on adaxial surface; main veins on abaxial surface 8–10 on each side of midvein, 1–1.5 mm apart; leaf sheath veined, at least at base and along margins; inflorescence with 30–40 spikelets .................................... S. steyermarkii Petals yellow, the blade suborbicular or rhombic, 15–18 mm long; inflorescence with 14–25(–35) spikelets, these 15–20 mm long, ± saliently spaced or distant from one another, the bracteoles pale or golden brown when dry; leaf sheaths conspicuously veined, submembranous when young, with age marcescent, becoming frayed and fibrous, the lower caudex appearing nearly cylindrical as opposed to the equitant green part of the foliage ......................................................... S. angustata

Stegolepis 453

9.

10(5).

10.

11(10).

11.

12(11).

12.

13(2). 13. 14(13).

14.

15(14).

Petals orange-yellow, the blade lanceolate to narrowly ovate, 6–9 mm long; inflorescence occasionally with 40–50 spikelets, but usually many more, these 12–15 mm long, densely crowded, not saliently spaced or distant from one another, the bracteoles darker brown when dry; leaf sheaths veinless, firm-hardened, markedly equitant on the caudex ................................................................................... S. parvipetala Leaf blades glaucous on both sides, ± brittle when fresh; spikelets (including the sepals) 8–10 mm long, the bracteolate part only ± 1/2 as long as the spikelet; bracteoles tan, rounded to obtuse; peduncle much longer than the leaves ......................................................... S. maguireana Leaf blades not glaucous nor brittle when fresh (sometimes glaucous in S. grandis); spikelets (8–)10–25 mm long, the bracteolate part > 1/2 the length of the spikelet; bracteoles tan to generally dark brown, some or most acute to acuminate; peduncle usually not much longer than the leaves (except S. grandis) .................................................................... 11 Peduncle usually compressed ancipitally (like a 2-edged blade) at apex, where dilated to (12–)17–25 mm wide, elsewhere 4–14 mm wide (sometimes 3-edged instead); spikelets with 25–40 bracteoles; blade of petals > 11 mm long; midvein prominent on abaxial leaf surface, but primary lateral veins inconspicuous; tepuis of Amazonas and southwestern Bolívar ........................................................................... S. grandis Peduncle compressed and noticeably 3-edged (triquetrous) at apex, where dilated to 6–12 mm wide, elsewhere 3–5 mm wide; spikelets with 12– 25 bracteoles; blade of petals < 11 mm long; midvein on abaxial leaf surface prominent or not, the primary lateral veins conspicuous; eastern Bolívar (Gran Sabana and surrounding tepuis) .......................... 12 Inflorescence 2.5–3.5(–4) cm diameter; spikelets (8–)10–15 mm long, bracteoles 12–16(–20); petals orange-yellow, the blade lanceolate to narrowly ovate, 6–9 mm long, ca. 5 mm wide; leaves green on ventral surface, silvery green on dorsal surface with contrasting green lines from lateral veins; on the northwestern tepuis of Bolívar .......... S. parvipetala Inflorescence (3.5–)4–4.5 cm diameter; spikelets (12–)15–20 mm long, bracteoles 20–25; petals yellow, the blades ovate to suborbicular and 8–11 mm long and wide; leaves without the above color combination; restricted to the eastern tepui chain along the Guyana border with Bolívar .................................................................................... S. guianensis Leaf blades 0.3–1 cm wide ....................................................................... 14 Leaf blades 1–9 cm wide .......................................................................... 16 Spikelets (2)3–8 per inflorescence; peduncle 3–4 mm diameter, strongly ribbed, dilated to 5 mm wide at apex; leaves 3–5 mm wide, arising from below ground level in deep peat; Bolívar (Cerro Jaua) .......... S. jauaensis Spikelets 1 or 2 per inflorescence; peduncle slender, 1–2 mm diameter, faintly or inconspicuously ribbed, slightly dilated to 2–4 mm wide at apex; leaves 6–10 mm wide, arising above ground level; not known from Cerro Jaua ................................................................................... 15 Leaf sheaths stiff, with a prominently prolonged ligule at the top; leaf blades with a metallic blue iridescence when fresh; spikelets (including the sepals) 27–30 mm long; Bolívar (Macizo del Chimantá) ..................................................................................................... S. ligulata

454

R APATEACEAE

15.

16(13). 16. 17(16).

17.

18(17). 18. 19(18). 19. 20(19).

20.

21(20).

21.

22(19).

22.

23(16). 23. 24(23). 24. 25(23).

25.

Leaf sheaths membranous, gradually narrowed to the base of the leaf blade, without a ligule; leaf blades not markedly iridescent when fresh; spikelets 10–12 mm long; Amazonas (Cerros Duida, Aracamuni, and Avispa) ......................................................................................... S. linearis Spikelets 7 or more per inflorescence ..................................................... 17 Spikelets 1–6 per inflorescence ............................................................... 23 Bracteoles, at least the lower and middle ones, obtuse to rounded at apex; spikelets 1–7 per inflorescence; Amazonas (Cerro Marahuaka) ........................................................................................... S. terramarensis Bracteoles subacute to conspicuously acuminate or aristate at apex; spikelets mainly (7)8–50 per inflorescence; not known from Cerro Marahuaka ........................................................................................... 18 Leaf blades 2–4.5 cm wide .................................................. S. ptaritepuiensis Leaf blades 4–9 cm wide .......................................................................... 19 Spikelets (including the sepals) 11–16 mm long, 3–5 mm wide ............ 20 Spikelets (including the sepals) 18–38 mm long, 6–15 mm wide .......... 22 Peduncle 3–6(–8) mm wide, except at dilated apex where 6–10 mm wide; inflorescence 35–40 mm wide, 20–35 mm high; spikelets 20–50; Amazonas (Sierra de la Neblina) ................................................... S. celiae Peduncle 1.5–3 mm wide, except at slightly dilated apex where 2.5–4 mm wide; inflorescence 25–30 mm wide, 15–25 mm high; spikelets < 20; not known from Sierra de la Neblina ................................................. 21 Inflorescence strongly compressed; peduncles generally solitary in the leaf sheath; spikelets < 15 per inflorescence, 11–12 mm long, the upper bracteoles linear-lanceolate; mature leaf blades mostly > 6 cm wide; Bolívar (Sierra de Lema) ............................................................... S. minor Inflorescence laxly compressed; peduncles several per axil; spikelets > 15 per inflorescence, ca. 15 mm long, the upper bracteoles broadly ellipticlanceolate; mature leaf blades 4–6 cm wide; Amazonas ..... S. choripetala Inflorescence with 2 or 3 rows of spikelets, these 18–23 mm long, 6–8 mm wide, bracteoles ending in recurved, acute to attenuate tips; Bolívar (Cerro Guaiquinima) ...............................................................S. squarrosa Inflorescence with 1 or 2 rows of spikelets, these 30–38 mm long, 8–15 mm wide, bracteoles appressed or ascending; Amazonas (Cerro Duida) .................................................................................................... S. pungens Inflorescence of 1 or 2 spikelets, with outermost elongate bracts 14– 28 mm long, much longer than the inner bracteoles ......................... 24 Inflorescence of 1 or more spikelets, the outermost bracteoles not elongate, if > 14 mm long then not markedly longer than inner ones ..... 25 Auricles of leaf sheath poorly developed; outer bracts 14–18 mm long; Amazonas (Cerro Duida) ..................................................... S. gleasoniana Auricles of leaf sheath well developed, 3–4(–6) cm long; outer bracts 26– 30 mm long; Bolívar (La Escalera) ....................................... S. perligulata Leaf sheath thin-papery when dry, lanceolate, the widest part only slightly broader than or to 1.5 times as wide as the widest part of the leaf blade, without auricles at the apex or these poorly developed ................................................................................................... S. cardonae Leaf sheath thin-papery to firm-hardened, varied in shape but generally not lanceolate, the widest part markedly broader than the widest part of leaf blade, auricles usually well developed at the apex ................. 26

Stegolepis 455

26(25). Leaf blades 1.5–2.5 cm wide; bracteoles of spikelets dark brown ......... 27 26. Leaf blades > 2.5 cm wide, or if narrower, bracteoles of the spikelets pale brown or tan ......................................................................................... 29 27(26). Inflorescence with 4–6 spikelets; leaf blades 15–20 cm long, not prominently veined on lower surface; leaf sheath thin-membranous, the auricles 10–15 mm tall, 17–22 mm wide; summit of Kukenán-tepui and Roraima-tepui ............................................................................... S. huberi 27. Inflorescence with 1–3(4) spikelets; leaf blades 25–90 cm long, generally ± prominently veined on lower surface; leaf sheath membranous to firm-hardened, the auricles smaller than above; not known from Kukenán-tepui or Roraima-tepui ........................................................ 28 28(27). Leaf sheath ≥ 2 times wider than the base of the leaf blade, prominently auriculate at apex, the auricles 5–7 mm tall; leaf blades mostly < 35 cm long; longest spikelets of the inflorescence usually < 18 mm long; Bolívar ......................................................................................... S. humilis 28. Leaf sheath < 2 times wider than the base of the leaf blade, slightly auriculate at apex, the auricles 3–5 mm tall; leaf blades mostly > 50 cm long; longest spikelets of the inflorescence usually > 22 mm long; Amazonas .............................................................................. S. neblinensis 29(26). Inflorescence with 1–7 spikelets, but most with 4 or more spikelets .... 30 29. Inflorescence with 1 or 2 (occasionally 3) spikelets ................................ 31 30(29). Leaf sheath not conspicuously scariose-margined, the auricles 15–18 mm tall, 25–35 mm wide; apex of leaf blade rounded to falcately obtuse; bracteoles of spikelet obtuse at the apex ......................... S. terramarensis 30. Leaf sheath conspicuously scariose-margined, the auricles generally smaller than above; apex of leaf blade subacute to rounded; bracteoles of the spikelets acute at the apex ......................................... S. hitchcockii 31(29). Leaf blade (3.5–)4–5 cm wide at broadest point, coriaceous ..... S. pauciflora 31. Leaf blade 2–3.5 cm wide at broadest point, firmly chartaceous .............................................................................................................. 32 32(31). Leaf sheaths firm-hardened, margins and auricles with a slight, brownish, membranous margin, the auricles rotund, 5–8 mm long; peduncles subfiliform, 0.5–0.9 mm thick ................................................ S. wurdackii 32. Leaf sheaths membranous or slightly hardened, margins and auricles broadly white-membranous or -scariose, the auricles rotund or not, but larger than above; peduncles 1–3 mm wide ....................................... 33 33(32). Leaf sheaths membranous, thin-papery when dry, evidently veined; peduncles slender, angular or compressed, 1–2.5 mm wide; spikelets, including sepals, 16–18(–22) mm long ................................ S. membranacea 33. Leaf sheaths not membranous, ± firm-hardened when dry except for the scariose margins, not evidently veined; peduncles 1–4 mm wide; spikelets, including sepals, (18–)20–30 mm long............................. S. pulchella Stegolepis albiflora Steyerm., Ann. Missouri Bot. Gard. 74: 609. 1987. Herb 0.7–1.5 m tall; leaf sheaths 20–25 cm long, 4–6 cm wide, ligulate at the apex, the blades 58–70 cm long, 3.5–5 cm wide; peduncles numerous, 45–70 cm long, 0.8–1.5 mm diameter, sulcate; inflorescence globose, 1.8–2.5 cm diameter, spikelets radiate, lan-

ceolate, 7–10 mm long; sepals 6–7 mm long; petals white, oblanceolate or lanceolate, acute, recurved, the blade 6–7 mm long, 2.5– 3 mm wide; anthers yellow. Tepui meadows and bogs, 1700–2300 m; Bolívar (Cerro Jaua, Cerro Sarisariñama), Amazonas (Sierra de Maigualida). Endemic. ◆Fig. 381.

456

R APATEACEAE

Stegolepis albiflora is very similar to S. choripetala, differing mainly in the somewhat smaller flowers with white petals. Stegolepis angustata Gleason, Bull. Torrey Bot. Club 56: 18. 1929. Generally robust herb 1–2.5 m tall; the lower stem to 50 cm tall, ± round in cross section, on older plants with persistent, frayed, fibrous leaf sheaths; leaf sheaths 15–20 cm long, 4.5–7 cm wide, the blades 100–150 cm long, 3–4 cm wide; peduncles 1–3 per axil, to 1.5 m tall, ca. 5 mm diameter, ± expanded at apex; inflorescence globose, 3.5–4.5 cm diameter, with 14–25(–40) loosely aggregated spikelets, these 16–20 mm long; bracteoles 24–30, acute, the larger interior ones to 15 mm long and 6 mm wide, light tan-brown; sepals elliptic-oblong, 16–18 mm long, the distal part often conspicuously reflexed at anthesis; petals yellow, with a basal connate portion 6–8 mm long, the blade suborbicular and 15–18 mm diameter. Moist areas in sandy savannas and forest edges, 800– 1500(–1800) m; Bolívar (Cerro Chirikayén, 10 km east of Cerro Venado, upper slopes of Cerro Venamo, 17 km east of El Paují, widespread in the greater Gran Sabana, shoulder of Kamarkabarai-tepui, middle Río Antebare, top of Sierra de Lema, plateau south of Tereké-yurén-tepui). Guyana. ◆Fig. 383. Stegolepis angustata has one of the widest altitudinal ranges in the genus and occurs at the lowest elevation of any species of Stegolepis. It has characteristic loosely spaced spikelets with light tan bracteoles, large yellow petals, and reflexed sepals. Its persistent fibrous leaf sheaths sometimes show signs of charring from previous savanna fires. In Guyana it grows as low as 500 m elevation near Kaieteur and Imbaibadai, whereas in Venezuela the lowest elevation recorded is around 850 m, east of El Paují and in the Río Antebare region northeast of Canaima. It does not occur on the summit of any of the major tepuis, and the highest elevation recorded is 1800 m on the slopes (not summit) of a mountain in Los Testigos range. Stegolepis breweri Maguire, Bol. Soc. Venez. Ci. Nat. 33: 279. 1976. Herb to 0.5 m tall; old leaf sheaths marcescent and ± tubular, young leaf sheaths ca. 15 cm long with ligulate auricles, the blades linear-lanceolate and 35–45 cm long, 1.4–1.6

cm wide; peduncles apparently 1 or 2 per plant, ca. 50 cm long, strongly compressedwinged, 8–10 mm wide, 5–16-veined; inflorescence with 2–5 spikelets; bracteoles narrowly lanceolate, the inner ones 10–12 mm long; sepals ca. 20 mm long. In large clumps on steep cliffs (inside large sinkholes), 700– 1100 m; Bolívar (Cerro Sarisariñama). Endemic. ◆Fig. 379. Stegolepis breweri is very similar to Phelpsiella ptericaulis Maguire, except for the absence of the spathaceous bracts. Stegolepis cardonae Maguire, Mem. New York Bot. Gard. 12(3): 87. 1965. Herb 0.3–1.0 m tall; leaf sheaths papyraceous, to 30 cm long, 2.5–3 cm wide, without auricles; leaf blades linear-lanceolate, to 80 cm long, 1.8–2.8 cm wide; peduncles 4 or 5, sulcate, 1.8–2.5 mm thick, to 60 cm tall, inflorescence with (1–)3–5 spikelets; sepals 17–20 mm long, consisting of a lower tube 3 mm long, then a stiff blade 12–13 mm long and 4– 5 mm wide that is reflexed at anthesis; petals with basally connate claws for 3 mm, the blades orbicular, bright yellow, 19–23 mm long. Shrub islands and low forests on tepui summits and base of cliffs, streamsides, 1500– 2250 m; Bolívar (Angasima-tepui, Macizo del Chimantá [Acopán-tepui, Amurí-tepui, Churitepui, Murey-tepui]). Endemic. ◆Fig. 382. Stegolepis celiae Maguire, Mem. New York Bot. Gard. 12(3): 80, fig. 11A–I. 1965. Herb 0.5–1 m tall, stems sometimes leaning; leaf sheaths 18–25 cm long, 7–10 cm wide, auricles rounded, 6–10 mm tall; blades broadly linear-lanceolate, 50–80 cm long 5–9 cm wide; peduncles 3 or 4 per leaf axil, 60–80 cm tall; inflorescence with (20–)30–50 spikelets, these 12–16 mm long; sepals 11–12 mm long, 3 mm wide; petals golden yellow, with claws basally connate, the blades obcordate and 15–18 mm wide. Open areas and scrub on tepui summits and slopes, 1300–2000 m; Bolívar (Cerro Aracamuni, Cerro Avispa, Sierra de la Neblina). Adjacent Brazil (Amazonas: Serra Pirapicú). Stegolepis choripetala Maguire, Mem. New York Bot. Gard. 12(3): 75. 1965. Herb 2–3 m tall, the stem thick and to 15 cm tall; leaf sheaths 20–30 cm long, 5–6 cm wide, without a ligule, blades 70–250 cm long, 3.5–5 cm wide; peduncles 5–7 per axil,

Stegolepis 457

80–180 cm tall, 2–3 mm diameter, compressed and dilated below the inflorescence where 3–4 mm wide; inflorescence ± compressed, with 15–20(–30) spikelets in 2 or 3 series; spikelets ca. 15 mm long, sepals 9–11 mm long; petals yellow, with fleshy claws 4–5 mm long, the blade lanceolate, 5–6 mm long, 3 mm wide, acute-tipped. Marshy open areas and shrub thickets, 1500–2100; Bolívar (Sierra de Maigualida), Amazonas (Cerro Sipapo). Endemic. Stegolepis gleasoniana Steyerm., Fieldiana, Bot. 28: 130. 1951. Herb ca. 1 m tall; leaf blades linear-lanceolate, rigid-coriaceous, ca. 90 cm long, 1– 2.5(–5.5) cm wide, veins prominent on adaxial side, more veins on one side of the midvein than on the other; peduncles slender, 4–7 per axil, inflorescence with 1–3 spikelets, these 14–18 mm long; outer bracts as long as the spikelet, interior ones narrowly lanceolate, to 9 mm long; sepals lanceolate, not reflexed at anthesis; petals dark yellow, blades suborbicular, apiculate, ca. 16 mm long. Sandstone outcrops, streamside scrub, 1500–2100 m; Amazonas (southeast portion of Cerro Duida). Endemic. Stegolepis grandis Maguire, Mem. New York Bot. Gard. 12(3): 72. 1965. Stegolepis grandis subsp. phelpsiae Maguire, Mem. New York Bot. Gard. 12(3): 73. 1965, syn. nov. Stegolepis grandis subsp. jauensis Maguire, Mem. New York Bot. Gard. 23(9): 848. 1972, syn. nov. Giant herb to 3 m tall; leaf sheaths 30–50 cm long, 5–9 cm wide, the blades 100–250 cm long, 4–8 cm wide; inflorescence globose, 4.5– 5.5 cm diameter, spikelets ca. 100, ca. 20 mm long, bracteoles 25–40; petals yellow, with claws basally connate for 10–12 mm, ca. 25 mm long. Wet tepui summit meadows and shrub thickets, 1000–2400 m; Bolívar (Cerro Guanacoco, Cerro Jaua, Cerro Sarisariñama), Amazonas (Cerro Aracamuni, Cerro Avispa, Cerro Duida, Cerro Huachamacari, Cerro Marahuaka, Cerro Parú, upper Río Ventuari). Endemic. ◆Fig. 385. Stegolepis guianensis Klotzsch ex Körn., Linnaea 37: 481. 1872. Herb 1–2 m tall, the old leaf sheaths often marcescent; leaf sheaths 20–25 cm long, 6–9

cm wide, blades 60–100 cm long, 3.5–5.5 cm wide; peduncles 1–various, 1–1.5 m tall, 4–5 mm thick, widened and triquetrous (obliquely triangular in cross section) at the apex; inflorescence globose, (3.5–)4–4.5 cm diameter, with 50–90 spikelets, these 16–18 mm long; bracteoles 20–25, obtuse, inner ones to 15 mm long, scariose; sepals 13–15 mm long; petals golden yellow, with connate claws 4–5 mm long, the blades ovate to suborbicular, 8–11 mm long and 10–12 mm wide. Sandy, moist, open areas, often on rocks or cliffsides, (1800–)2000–2810 m; Bolívar (Karaurín-tepui, Kukenán-tepui, Roraimatepui, Tramen-tepui, Uei-tepui, Yuruanitepui). Guyana (Mount Roraima). ◆Fig. 384. Stegolepis guianensis is a high-elevation species restricted to the summits and upper slopes of the eastern tepui chain. Stegolepis hitchcockii Maguire, Mem. New York Bot. Gard. 12(3): 82. 1965. Herb 0.8–2 m tall; leaves strongly equitant-sigmoid, often bluish-iridescent on one side; leaf sheaths 10–20 cm long, 5–8.5 cm wide, auricles ca. 1 cm long and 2–3 cm wide, the blades 70–90 cm long, 4–7 cm wide; peduncles 4–6 per axil, 1.5–3 mm diameter, 50–70 cm tall, compressed near apex; inflorescence with 1–6 spikelets in 1 or 2 rows, spikelets 15–18 mm long; sepals reflexed at anthesis, the stiff distal portion ca. 14 mm long; petals bright yellow, broadly obovateobcordate, apiculate, the claws 6 mm long, the blades 12–14 mm long. Moist open savannas on tepui summits, 800–2000 m; Amazonas; 2 subspecies, both endemic to the flora area. Key to the Subspecies of S. hitchcockii 1. Leaf sheaths oblong, blades usually 2.5–4 cm wide; spikelets including sepals 15– 16 mm long; bracteoles somewhat obtuse, ± hooded, not marginate ................ ............................... subsp. hitchcockii 1. Leaf sheaths more inflated than above, blades usually 4–5 cm wide; spikelets including sepals 17–20 mm long; bracteoles somewhat acute, not hooded, narrowly pale-margined ..... subsp. morichensis S. hitchcockii subsp. hitchcockii 1200–2000 m; Amazonas (Cerro Parú). Endemic.

458

R APATEACEAE

A collection from Cerro Parú at 1100 m (Huber 4290, NY) may be a robust form of this subspecies. S. hitchcockii subsp. morichensis Maguire, Mem. New York Bot. Gard. 12(3):83. 1965. 700–1300 m; Amazonas (Cerro Moriche, Cerro Yapacana). Endemic. ◆Fig. 387. Stegolepis huberi Steyerm., Ann. Missouri Bot. Gard. 74: 610. 1987. Dwarf herb 0.2–0.4 m tall; leaf sheaths 7– 10 cm long, 3–3.5 cm wide, the auricles rounded at apex, 10–15 mm tall and 17–22 mm wide, blades 15–20 cm long 1.8–2.5 cm wide; peduncles compressed, 3–6-costate, 17–25 cm long; inflorescence compressed, 2– 3.5 cm wide, 1–1.7 cm tall, with 4–6 spikelets, these 15–18 mm long; sepals ca. 13 mm long; petals rhomboid, 10 mm long, 7–8 mm wide. In colonies in open rocky rock areas on high-tepui summits, 2500–2700 m; Bolívar (Kukenán-tepui). Guyana (Mount Roraima). Stegolepis humilis Steyerm., Ann. Missouri Bot. Gard. 74: 611, fig. 1. 1987. Herb 0.3–0.8 m tall; leaf sheaths 11–12 cm long, 7–9 cm wide, the apex with rounded auricles 5–7 mm tall and 12–13 mm wide; blades ± striped on adaxial surface, 27–45 cm long, 1.3–2.5 cm wide; peduncles 2 or 3, 5ribbed, 27–52 cm tall, 1–1.5 mm thick except at the top where 4–5 mm wide; inflorescence with 1 or 2 spikelets, these 15–18 mm long, bracteoles dark brown, stiff; sepals 15 mm long; petals with a long claw ca. 10 mm long, the blade broadly rhombic, 12–13 mm long, ca. 18 mm wide. Open, rocky areas on tepui slopes and summits, 1700–2500 m; Bolívar (Auyán-tepui, Los Testigos tepui chain). Endemic. ◆Fig. 389. Plants from Auyán-tepui are generally larger than those from Los Testigos and were previously treated under Stegolepis neblinensis. Stegolepis jauaensis Maguire, Mem. New York Bot. Gard. 23: 848. 1972. Juncus-like herb 0.7–1.5 m tall, in dense clumps; marcescent leaf bases in a tubular structure sunk into the ground litter or peat; leaf sheaths 10–20 cm long, 1–3 cm wide, the blades grass-like, erect, 80–120 cm long, 0.3– 0.5 cm wide, eligulate; peduncles to 1.2 m tall,

ribbed; inflorescence compressed, usually with 1–3 spikelets in 1 row, occasionally with 5–10 spikelets in 2 rows, the spikelets 15–20 mm long; petals bright yellow, fan-shaped, the blade ca. 10 mm long. Bogs on tepui summits, 1900–2100 m; Bolívar (Cerro Jaua, Cerro Sarisariñama). Endemic. ◆Fig. 388. Stegolepis ligulata Maguire, Mem. New York Bot. Gard. 12(3): 87, fig. 12B–D, H– L. 1965. Herb 0.3–0.8 m tall; leaf sheaths lanceolate, 10–20 cm long, 1.5–2.5 cm wide, with ligules to 4 cm tall; blades grass-like, 30–60 cm long, 0.5–0.8 cm wide, metallic-blue iridescent when fresh; peduncles 1–3 per plant, 50–70 cm tall, ca. 1.5 mm diameter, slightly trigonous at apex; inflorescence with a solitary spikelet 27–30 mm long; sepals 24–26 mm long, the upper portion reflexed at anthesis; petals yellow-orange, the blades 18– 20 mm long, 15–16 mm wide. Locally abundant and forming large colonies in swampy meadows, on rocks, or in thickets, 1800–2500 m; Bolívar (Macizo del Chimantá). Endemic. ◆Fig. 380. Stegolepis linearis Gleason, Bull. Torrey Bot. Club 58: 335, pl. 25A. 1931. Herb 0.2–0.8 m tall; leaf sheaths 10–12 cm long, ca. 5 cm wide, blades rigid, narrowly linear, 50–60 cm long, 0.6–0.8 mm wide, 9ribbed; peduncles 40–80 cm tall, flattened near the apex; inflorescence with 1–3 spikelets, these 10–12 mm long; petals with orbicular yellow blade 20–23 mm long. Swampy open areas on tepuis, 1300–2100 m; Amazonas (Cerro Aracamuni, Cerro Avispa, Cerro Duida). Endemic. Stegolepis maguireana Steyerm., Ann. Missouri Bot. Gard. 71: 300. 1984. Herb 1.5–3 m tall; leaf sheaths 15–30 cm long, 3–6 cm wide, blades glaucous, 150–200 cm long, 2–3 cm wide; peduncles 1.5–4 m tall, 5–6 mm diameter; inflorescence globose, 3.5– 4 cm diameter, with 50–80 spikelets, these 8– 9 mm long, sepals blunt-tipped, pale, 6–8 mm long, not reflexed at anthesis; petals with yellow, ovate blade 7–10 mm long. Wet sandy tepui summit meadows and sandstone outcrops, 1900–2200 m; Bolívar (Macizo del Chimantá [Acopán-tepui, Amurí-tepui]). Endemic.

Stegolepis 459

Stegolepis membranacea Maguire, Mem. New York Bot. Gard. 12(3): 84. 1965. Herb 0.5–1.5 cm tall; leaf sheath 12–16 cm long, 3–4 cm wide, blades 70–100 cm long, 2.5–4 cm wide, prominently veined; peduncles mostly 5–8, sometimes angled or compressed, to 50 cm long; inflorescence with 1 or 2 spikelets, these 16–18 mm long, bracteoles 18–20, narrowly triangular-lanceolate, 4–7 mm long; sepals 17–20 mm long, petals with obcordate blade 12–15 mm long. On rocks along streams, on rocky tepui bluffs, 1000–1800 m; Amazonas (Cerro Duida, Cerro Huachamacari, Cerro Marahuaka). Endemic. Stegolepis microcephala Maguire, Bol. Soc. Venez. Ci. Nat. 33: 280. 1976. Herb ca. 0.5 m tall, with a bulbous, whitish, onion-like base; leaf sheaths narrowly lanceolate, ca. 15 cm long, 1–1.5 cm wide; leaf blades ca. 40 cm long, 10–12 mm wide, prominently veined and ± striped on lower surface; peduncles several, striate and ± flattened; inflorescence angular-subglobose, obliquely affixed to the peduncle, with (15–)30– 50 spikelets, these 6–10 mm long; bracteoles gradate, 3–5 mm long; petals bright yellow, blade obovate, ca. 15 mm diameter. Growing in peaty bogs in dense tepui meadows and edges of tepui gallery forests, 1700–1800 m; Bolívar (Cerro Jaua). Endemic. ◆Fig. 391. Stegolepis minor Steyerm., Ann. Missouri Bot. Gard. 75: 315. 1988. Terrestrial or epiphytic herb 0.5–1 m tall; leaf sheaths 14–25 cm long, ca. 7 cm wide, blades ca. 60 cm long, 6.5 cm wide; inflorescence compressed, with 7–20 spikelets in 1 or 2 rows, the spikelets 11–12 mm long; sepals erect, 9–10 mm long; petals not known. Low montane forests, bases of sandstone bluffs near waterfalls, 500–1000 m; Bolívar (Río Chicanán in Sierra de Lema). Endemic. Stegolepis neblinensis Maguire, Mem. New York Bot. Gard. 12(3): 78, fig. 10E– J. 1965. Herb 0.3–0.9 m tall; leaf sheaths stiff, 10– 18 cm long, 3.5–5 cm wide, with a rounded auricle 3–5 mm tall; blades 50–70 cm long, 1.5–2.5 cm wide; peduncles 2–4, 30–50 cm long; inflorescence compressed, with 1–6 spikelets, these 25–35 mm long, bracteoles

24–28; petals yellow, with broadly obovate or obcordate blades 20–25 mm long and 15– 25 mm wide. Moist tepui meadows and bogs; 1600–2200 m; Amazonas (Sierra de la Neblina). Endemic (most likely present as well on Brazilian side of border). ◆ Fig. 394. Specimens of Stegolepis humilis from Auyán-tepui were previously included in S. neblinensis. Stegolepis parvipetala Steyerm., Fieldiana, Bot. 28: 133. 1951. Stegolepis parvipetala subsp. chimantensis Maguire, Mem. New York Bot. Gard. 12(3): 75. 1965, syn. nov. Herb 1–2 m tall; leaf sheaths firmspongiose, tan-brown, smooth-surfaced, 15– 25 cm long, 5–8 cm wide, blades to 130 cm long and 6 cm wide; peduncles 50–80 cm tall, 8–15 mm wide; inflorescence globose, with (40–)50–100 spikelets, bracteoles obtuse, 3–9 mm long; petals with a connate base ca. 6 mm long, the blade orange-yellow, lanceolate to narrowly ovate, 5–9 mm long, ca. 5 mm wide, tubular-spreading and sometimes barely emergent from the spikelets at anthesis; anthers orange. On tepui summits and slopes on open rocks, in meadows, and in Bonnetia shrublands, 1000–2500 m; Bolívar (Amaruay-tepui, Auyán-tepui, Kamarkaibarai-tepui, throughout the Macizo del Chimantá, Murisipán-tepui, Ptari-tepui, Sororopán-tepui, Tereké-yurén-tepui). Endemic. ◆Fig. 395. Stegolepis pauciflora Gleason, Bull. Torrey Club 58: 336, pl. 250. 1931. Herb 0.6–1 m tall; leaf sheaths 18–25 cm long, 4–6 cm wide, blades (3.5–)4–5 cm wide; inflorescence with 1 or 2 spikelets 15–25 mm long, the outer bracteoles 3 mm long, the inner ones to 13 mm long, obtuse; sepals ca. 25 mm long, with a membranous base, the distal 17–18 mm long, stiff, exserted, and reflexed at anthesis; petals with claws ca. 10 mm long, the blade orbicular-obovate, 20–23 mm long. Moist rock outcrops on tepui summit, 1800–2100 m; Amazonas (upper elevations of Cerro Duida). Endemic. Stegolepis perligulata Maguire, Mem. New York Bot. Gard. 12(3): 88, fig. 12M– Y. 1965.

460

R APATEACEAE

Herb to 1.5 m tall; leaf sheaths ca. 25 cm long, 3 cm wide, auricles ligulate and 30– 40(–60) mm long, ca. 15 mm wide, blades linear-lanceolate, to 120 cm long, 1.4–1.8 cm wide; peduncles 6–8, sulcate, 40–60 cm long; inflorescence with a single spikelet 26–30 mm long, with 2 narrowly lanceolate outer bracts, the outer one 26–30 mm long and 3 mm wide, the inner one 20–25 mm long and 4 mm wide, bracteoles 6–8, narrowly lanceolate, 10–21 mm long; petals yellow, broadly obovate, blade ca. 15 mm wide. Locally frequent on escarpments, 800–900 m; Bolívar (Río Uiri-yuk in the headwaters of the Río Cuyuní in the Sierra de Lema). Endemic. ◆Fig. 393. Stegolepis ptaritepuiensis Steyerm., Fieldiana, Bot. 20: 133. 1951. Herb 0.5–2 m tall, sometimes with a trailing caudex; base strongly compressed with nonfibrous, equitant, marcescent leaves; leaf sheaths 10–18 cm long, 4–5 cm wide, with auricles 8–10 mm long, blades 40–80 cm long, 2–4.5 cm wide; peduncles usually 3–5 per axil, to 1.5 m tall, 2–3 mm thick, striate, compressed and widened to 10 mm wide at the apex; inflorescence fan-shaped, with 5– 20 spikelets usually in one row, the spikelets 18–22 mm long, bracteoles 22–25, pungent, the exterior ones 3–5 mm long; sepals 13–15 mm long petals with a basally connate tube 2–4 mm long, the blades yellow, obcordate, apiculate, 8–12 mm long. Swampy whitesand savannas, 900–1400(–2400) m; Bolívar (lower slopes of Amaruay-tepui, slopes of Auyán-tepui, widespread in the Gran Sabana, middle slopes of Ilú-tepui, summit and slopes of Ptari-tepui). Guyana. ◆Fig. 386. This is mainly a mid-elevation species, most common between 900 and 1400 m elevation. At the eastern end of its range in Guyana, it extends down to 550 m in the Pakaraima region and the Merume Mountains. It is present on the slopes of a number of tepuis, but apparently only rarely reaches the summit (perhaps only on Ptari-tepui, where it is infrequent on the summit at 2400 m). Stegolepis pulchella Maguire, Mem. New York Bot. Gard. 12(3): 83. 1965.

Herb 0.5–1.5 m tall; leaf sheaths 10–20 cm long, 3–5 cm wide, with rounded auricles 20–30 mm wide, blades 70–100 cm long, 2.5– 3.5 cm wide; inflorescence with 1–3(4) spikelets 8–12 mm long (without the sepals); sepals 20–22 mm long, the claw ca. 4 mm long, the blade stiff, ca. 16 mm long, reflexed at anthesis; petals with a basal claw ca. 6 mm long, the blade obcordate, 22–25 mm long, 27–28 mm wide. Moist open meadows, on rocks, streamsides, 1000–1800 m; Amazonas (Cerro Autana, Cerro Camani, Cerro Guanay, Cerro Sipapo, Cerro Yutajé). Endemic. Stegolepis pungens Gleason, Bull. Torrey Bot. Club 58: 337, pl. 25D. 1931. Generally robust herb 0.8–2 m tall, with large flattened caudex; leaf sheaths 18–25 cm long, 7–10 cm wide, with rounded auricles at the apex to 10 mm tall; blades 50–80 cm long, 5–7 cm wide; peduncles 3 or 4 per axil, to 1 m tall, robust, striate, flattened towards the apex where 10–20 mm wide; inflorescence compressed, fan-like, with 8–20 spikelets in two rows, the spikelets 30–38 mm long, bracteoles 20–30, stiff, the outer ones triangular, acute, 6–8 mm long, the inner ones to 22 mm long, sometimes recurved; sepals 25–28 mm long, the lower part membranous and ca. 12 mm long, the upper part stiff and acuminate, 12–16 mm long; petals with claw ca. 10 mm long, the blades yellow, fleshy, orbicular-obcordate, ca. 25 mm wide. Swampy meadows and thickets on tepui summit, 1100–2200 m; Amazonas (Cerro Duida). Endemic. ◆Fig. 396. Stegolepis squarrosa Maguire, Mem. New York Bot. Gard. 12(3): 78, pl. 10A–D. 1965. Large herb 1–2 m tall; leaf blades 70–90 cm long, 5–7 cm wide, stiffly robust; peduncles 80–130 cm tall, 4–5 mm thick, triangularly widened and 10–15 wide at the apex; inflorescence compressed, fan-shaped, with 15–30 spikelets in 2 or 3 rows, the spikelets 18–23 mm long, 6–8 mm wide; bracteoles 22– 24, squarrose (abruptly recurved) when dry, the inner ones lanceolate and 16–17 mm long; sepals 20–22 mm long, petals yellow, the blades obcordate, apiculate, ca. 30 mm wide and 20 mm wide. Forming dense colo-

Stegolepis 461

nies in swampy tepui meadows, 600–1700 m; Bolívar (Cerro Guaiquinima). Endemic. ◆Fig. 398. Stegolepis steyermarkii Maguire, Mem. New York Bot. Gard. 12(3): 76. 1965. Terrestrial or epiphytic herb 0.7–1 m tall; leaf sheaths lanceolate, 15–20 cm long, 4–5 cm wide, blades linear-lanceolate, 40–60 cm long, 2.5–3 cm wide; peduncles often 4–6 per leaf axil, 40–70 cm tall, 1.5–2.5 mm diameter; inflorescence globose, 2–3 cm diameter, with 30–40 spikelets, these 12–14 mm long; bracteoles ca. 16, gradate, the apex acute, slightly hooded; sepals ca. 10 mm long, 2.5 mm wide; petals orangish yellow, connate for 2–3 mm at the base, the blades oblong-lanceolate, 10–12 mm long, recurved at anthesis. Montane forests, 800–1400 m; Bolívar (Cerro Venamo, La Escalera, north-facing slopes of Sierra de Lema). Guyana (Merume Mountains). ◆Fig. 397. Stegolepis steyermarkii is one of the few species of Stegolepis that grows occasionally as an epiphyte in tall forests. The small, narrow petals are characteristic of the group to which this species belongs (with S. ferruginea from Guyana and S. parvipetala from Venezuela). Stegolepis terramarensis Steyerm., Ann. Missouri Bot. Gard. 71: 301. 1984. Herb 0.5–1 m tall; leaf sheaths 14–15 cm long, 2.5–4 cm wide, topped by a suborbicular auricle 15–18 mm tall, 25–35 mm wide; leaf blades 45–55 cm long, 3–3.5 cm wide; peduncles 6–8, strongly ribbed, 25–55 cm long, 2–3 mm diameter except at apex where 4–6 mm wide; inflorescence with (1–)3–7 spikelets, these 10–15 mm long; sepals 14–18 mm long, petals yellow, the blade orbicular, 15–20 mm long. Subdominant in open rocky tepui summit areas, 2500–2600 m; Amazonas (Cerro Marahuaka). Endemic. ◆ Fig. 399. Stegolepis vivipara Maguire, Mem. New York Bot. Gard. 12(3): 74. 1965. Herb 1–1.5 m tall; leaf sheaths 12–18 cm long, 2.5–3.5 cm wide, exauriculate, leaf blades 30–70 cm long 1.5–2.5 cm wide; peduncles 1 or 2 per axil, 50–130 cm long; inflorescence globose, 2.5–3 cm diameter, with

25–35 spikelets, these ca. 15 mm long, many developing viviparous bulbils to 10 cm long; bracteoles ca. 15, suborbicular-ovate, 3–7 mm long; sepals reflexed at anthesis; petals yellow, obovate, 22–24 mm long, the blade 14–15 mm wide. Forming small colonies along river banks, forest edges, and in thickets on tepui summits, 1900–2300 m; Bolívar (Macizo del Chimantá [Amurí-tepui, Chimantá-tepui, Churí-tepui]). Endemic. ◆Fig. 392. Besides the type specimen, which has viviparous bulbils on the inflorescence, there are other specimens from the Macizo del Chimantá that do not show signs of vivipary, but otherwise agree with Stegolepis vivipara in floral and vegetative characters. Stegolepis wurdackii Maguire, Mem. New York Bot. Gard. 12(3): 85. 1965. Herb 0.7–1 m tall; leaf sheaths oblonglanceolate, 14–18 cm long, 3–4 cm wide, auricles rotund, 5–8 mm tall; blades 60–70 cm long, 1.5–3.5 cm wide; peduncles 5–15, subfiliform, 0.5–0.8 mm diameter, to 60 cm long; inflorescence usually with a solitary spikelet, rarely 2, each 23–25 mm long; bracteoles 18–20; sepals 22–24 mm long; petals with clawed base 4–5 mm long, the blade suborbicular, ca. 24 mm long and wide. Tepui meadows and scrub, 1600–2000 m. Venezuela; 2 subspecies, both endemic to the flora area. Key to the Subspecies of S. wurdackii 1. Peduncles 10–15; leaf blade 2.5–3.5 cm wide ................... subsp. chimantensis 1. Peduncles 3–5; leaf blade 1.5–2.5 cm wide ................................ subsp. wurdackii S. wurdackii subsp. chimantensis Maguire, Mem. New York Bot. Gard. 12(3): 86. 1965. 1700–2000 m; Bolívar (Macizo del Chimantá [Toronó-tepui]). Endemic. ◆Fig. 390. S. wurdackii subsp. wurdackii 1600–1800 m; Amazonas (escarpments of Sierra de la Neblina in headwaters of Río Yatúa). Likely in adjacent Brazil (Amazonas).

462

R APATEACEAE

Fig. 379. Stegolepis breweri

Fig. 381. Stegolepis albiflora

Fig. 380. Stegolepis ligulata

Stegolepis 463

Fig. 382. Stegolepis cardonae Fig. 383. Stegolepis angustata

464

R APATEACEAE

Fig. 386. Stegolepis ptaritepuiensis

Fig. 384. Stegolepis guianensis

Fig. 385. Stegolepis grandis

Stegolepis 465

Fig. 387. Stegolepis hitchcockii subsp. morichensis

Fig. 388. Stegolepis jauaensis

466

R APATEACEAE

Fig. 389. Stegolepis humilis Fig. 390. Stegolepis wurdackii subsp. chimantensis

Stegolepis 467

Fig. 391. Stegolepis microcephala

Fig. 392. Stegolepis vivipara

468

R APATEACEAE

Fig. 393. Stegolepis perligulata

Fig. 394. Stegolepis neblinensis

Stegolepis 469

Fig. 395. Stegolepis parvipetala

470

R APATEACEAE

Fig. 396. Stegolepis pungens

Stegolepis 471

Fig. 397. Stegolepis steyermarkii

Fig. 398. Stegolepis squarrosa

472

R APATEACEAE

Fig. 399. Stegolepis terramarensis

R H A M N A C E A E 473

RHAMNACEAE by Julian A. Steyermark and Paul E. Berry Trees, shrubs, or woody vines, rarely herbaceous, deciduous or evergreen, frequently armed with spines, or unarmed, sometimes with coiled tendrils. Leaves alternate, infrequently opposite, simple, pinnately veined or 3-veined from the base, entire or serrate; stipules usually present, but minute and deciduous (sometimes broad and persistent in Gouania). Inflorescence usually axillary and terminal, in corymbose or umbellate cymes, panicles, racemose or spicate thyrses, or reduced to a single flower. Flowers small, green or yellowish, bisexual or rarely polygamo-dioecious, 5-merous or sometimes 4-merous, perigynous to epigynous. Calyx tube obconic, turbinate, urceolate, or cylindric; calyx lobes 4 or 5, valvate; petals 4 or 5, free or absent, often concave or hooded, sessile or clawed. Stamens 4 or 5, opposite the petals and often enclosed by them; anthers 2-locular, longitudinally dehiscent. Disk intrastaminal, annular or lobed, free from the ovary or adnate around the base or to the complete length of the ovary. Ovary sessile, perigynous or epigynous, 2 or 3(–5) carpellate, imperfectly or completely many-locular; style usually short; stigma capitate or 3-lobed; ovules solitary or rarely 2–4 in each locule, anatropous. Fruit a drupe with 2–4 separate stones (pyrenes) or the stones plurilocular, or a coriaceous winged or unwinged capsule separating into pericarps. Seed solitary in the locules, often arillate, sometimes 1-grooved dorsally, often arillate at base; endosperm fleshy, scanty, or absent; embryo large, oily. Widely distributed in temperate and tropical regions of the world; ca. 55 genera and 900 species; 5 genera and 20 species in the flora area. Key to the Genera of Rhamnaceae 1. 1. 2(1).

2.

3(1). 3. 4(3).

4.

Vining shrubs or climbing plants .............................................................. 2 Trees or non-vining shrubs ........................................................................ 3 Leaves with 1 or 2 main pairs of lateral veins at the base; plants lacking tendrils; fruit a 3-locular drupe; inflorescence an elongated racemosely branched panicle, the flowers separated in small, widely spreading cymules ........................................................................... 1. Ampelozizyphus Leaves with 5–9 main pairs of lateral veins pinnately distributed; plants with tendrils; fruit a longitudinally narrowly or broadly 3-winged dry mericarp; inflorescence a narrow spicate thyrse composed of glomerules ............................................................................................. 3. Gouania Leaves with only 3 main veins, these all from the base; branches often with spines ................................................................................ 5. Zizyphus Leaves with > 3 main veins, some or all of them above the base; branches nonspiny ................................................................................................. 4 Base of leaf blade biglandular; lowest pair of leaf veins arising at the base; lateral pairs of veins 3; fruit dry, capsular-like with dehiscent locules; leaves and inflorescences frequently opposite or subopposite .................................................................................................. 2. Colubrina Base of leaf blade not glandular; lowest pair of leaf veins arising above the base; principal lateral pairs of veins 5–9; fruit fleshy; leaves and inflorescences always alternate .............................................. 4. Rhamnus

474

R HAMNACEAE

Fig. 400. Ampelozizyphus amazonicus

Colubrina 475

1. AMPELOZIZYPHUS Ducke, Arq. Inst. Biol. Veg. 2: 157. 1935. Vining shrubs or high-climbing lianas. Leaves distichous, alternate, entire, 5veined, the 2 pairs of lateral veins arising at and just above the base, or only 3veined with the lowest lateral pair arising at the base; stipules small, setaceous, caducous. Inflorescence mainly axillary, panicle racemosely elongated, bearing many alternately branched, interrupted, many-flowered cymes; cymes 1–3-dichotomous. Flowers bisexual. Calyx tube shortly turbinate, 5-lobed; petals 5, inserted on the margin of the disk, cucullate at apex, unguiculate below. Disk strongly adnate to the calyx tube, entire, orbicular, flat above. Ovary enclosed in the calyx tube, connate with it and with the disk, 3-locular. Ovules solitary, erect from base of the locule; style shortly 3-fid at the apex. Drupe stipitate above the fleshy torus base, the base of the stipe surrounded by the persistent calyx lobes; exocarp fleshy; endocarp hard, 3-locular. Seeds 3, shining, exalbuminous. Colombia, Venezuela, Guyana, Suriname, French Guiana, Peru, Brazil; 1 species. Ampelozizyphus amazonicus Ducke, Arq. Inst. Biol. Veg. 2: 158, t. 1, 2. 1935. —Bejuco pujajui, Hoja de mono, Palo de culebra. Vining shrub 2–3 m tall or high-climbing liana; young branches rufous-puberulent, older ones glabrous; leaves coriaceous, oblong-elliptic, lance-oblong, or oblong-ovate, shortly acuminate to subobtuse at apex, rounded or obtuse at base, 12–30 × 4.5–12.5 cm, quickly glabrous; inflorescence much elongated, 15–45 × 3–10 cm; flowers green to

greenish yellow; fruit subglobose at maturity, to 20 mm wide. Evergreen lowland and gallery forests, 50–200 m; Bolívar (Río Caura near Caño Guaya), Amazonas (Caño Yagua, Chipital, Río Mawarinuma, Río Siapa, San Carlos de Río Negro). Apure; Amazonian Colombia, Guyana, Suriname, French Guiana, Amazonian Peru and Brazil. ◆Fig. 400. The inner bark of Ampelozizyphus amazonicus has an odor of methyl salicylate (oil of wintergreen), and is used as a soap substitute.

2. COLUBRINA Rich. ex Brongn., Mém. Fam. Rhamnés 61. 1826, nom. cons. Shrubs or small trees, deciduous or evergreen, (sometimes with spines outside the flora area). Leaves alternate or opposite, pinnately veined or 3-veined from the base, entire or finely dentate, with or without small glands on the lower surface and/or 1 or 2 glands at the base of the blade; stipules minute. Inflorescence axillary, cymose or thyrsoid, sessile or shortly pedunculate. Flowers bisexual, small, disk fleshy, filling the floral tube, adnate to the lower half of the ovary. Calyx 5-lobed, lobes spreading, triangular-ovate, deciduous; petals 5. Stamens 5. Ovary semi-inferior, 3-locular; ovules 1 in each locule; style short, 3-lobed. Fruit a slightly 3-lobed capsule, dehiscent. Seeds 3, brown or black; testa smooth and shining. Widely distributed in the American tropics and subtropics, 1 species in Hawaii, 1 in Asia; 31 species, 2 in Venezuela, 1 of these in the flora area. Colubrina glandulosa Perkins, Bot. Jahrb. Syst. 45: 465. 1911. —Bernardino, Cartancillo, Guácimo cimarrón. Tree to 35 m tall; buds and young stems rufous brown, pubescent; inflorescence with rufous-tomentose, many-flowered cymes, equaling or shorter than the petioles; flowers cream-colored, green, pale yellow, or greenish yellow, shortly pedicellate; capsules sub-

globose, 6–8 mm diameter. Forested slopes along streams, seasonally dry evergreen forests, borders or Mauritia palm swamps in upland savannas, deciduous forests, 50–500 m; Bolívar (Altiplanicie de Nuria, La Vergareña, near Río Botanamo and Río Supamo, north of Upata). Panama, Peru, Brazil. ◆Fig. 401. The wood of Colubrina glandulosa is strong and hard and is used in construction.

476

R HAMNACEAE

Fig. 402. Gouania polygama

Fig. 401. Colubrina glandulosa

Gouania 477

3. GOUANIA Jacq., Select. Stirp. Amer. Hist. 263. 1763. Arching shrubs or lianas climbing by tendrils. Leaves alternate, pinnately veined or 3-plinerved at the base with additional veins above, dentate or entire, the teeth often inconspicuous or glandular; stipules usually narrow and deciduous or sometimes broad and persistent. Inflorescence in axillary or terminal spicate thyrses or racemes composed of glomerules, the rachis often ending in a tendril. Flowers polygamous or bisexual, small, whitish; disk 5-lobed, 5-angulate, or with 5 horn-like appendages, glabrous or pilose, epigynous, filling the calyx tube. Calyx tube obconic or subcampanulate, adherent to the ovary, 5-lobed, the lobes persistent; petals 5, cucullate, inserted below the margin of the disk. Stamens 5, concealed by the petals. Ovary inferior, immersed in the disk, 3-locular; style 3-parted with minute stigmas. Fruit coriaceous, a dry schizocarp, crowned by the persistent calyx, usually narrowly or broadly winged, with 3 indehiscent subligneous cocci, splitting longitudinally along the margin of each wing into 3 2-winged mericarps. Seeds 3, obovate; testa lustrous; endosperm scant. Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina, Uruguay, a few species in Asia, Africa, Madagascar, and islands in the Pacific Ocean; ca. 30 species, 7 in Venezuela, all in the flora area. The stems of some species, because of their saponin content, are chewed for their lather-producing effect, and when dried are exported to other countries for use in the preparation of dentrifices. The genus is greatly in need of revision. Key to the Species of Gouania 1. 1.

2(1).

2.

3(1). 3. 4(3).

4.

Disk pubescent; wings of fruit ± semilunate, longer than broad ............. 2 Disk glabrous; wings of fruit semilunate and longer than broad (horizontally extended, suborbicular or usually broader than long in G. polygama) ............................................................................................... 3 Disk pubescent throughout; marginal teeth of leaf blades salient, 15– 20 each side; leaf blades strongly discolored, with the lower surface cream- or buff-colored; lower surface of leaves densely soft-tomentose; fruit with densely pubescent wings, the trichomes spreading or lax ................................................................................................. G. wurdackii Disk pubescent in the lower (proximal) portion surrounding the stylar base, but glabrous in the upper (distal) portion; marginal teeth inconspicuous, few, mainly 3–5 in the lower 1/3–1/2 (rarely throughout); leaf blades ± concolored; lower surface of leaves mainly glabrous except for the appressed-pubescent midrib and veins; fruit with sparse or appressed pubescence or glabrous ......................................... G. frangulifolia Annulus (raised portion around style) glabrous ....................................... 4 Annulus sparsely or densely setose or pilose ........................................... 5 Leaf margins entire or with few repand callous shallow projections; lower surface of leaves glabrous, only the midrib and veins appressed-pubescent; leaf blades oblong or ovate-elliptic, 2–5 cm wide; fruit glabrous or sparsely appressed-pubescent ................................................. G. cornifolia Leaf margins generally prominently dentate with broad rounded dentations; lower surface of leaves usually densely villous; leaf blades chiefly broadly ovate, the larger ones 5–7.5 cm wide; fruit densely velutinous or villosulous with spreading or loose trichomes .................... G. colurnifolia

478

5(3).

5. 6(5).

6.

R HAMNACEAE

Leaf blades strongly discolored, the lower surface silvery- or gray-white or gray, with a minute dense indument or surface included within a fine tertiary network connecting conspicuous transverse veins ..................................................................................................... G. discolor Leaf blades not as above, the lower surface sparsely or densely covered with longer, appressed or spreading trichomes .................................... 6 Axis of fruit 3–4 mm long, the wings 3–7 × 3–7 mm, suborbicular, as wide as or wider than the axis, extending horizontally from the axis; axis much shorter than the extended wings; leaf blades usually with 8– 15 teeth on each margin, the teeth usually regularly distributed from near base to apex, rarely reduced or inconspicuous; lower surface of leaves ± pubescent, or at least appressed-pubescent on lower midrib and veins ................................................................................. G. polygama Axis of fruit 6–8 mm long, the wings 2–3(–6) × 7(–12) mm, semilunate, longer than wide; axis about equaling or slightly shorter than the length of the wings; leaf blades with either entire margins or at most inconspicuously repand-undulate or the crenations barely projecting and mainly < 8 to a margin, at most 1.5 mm long; lower surface of leaves glabrous, axillary trichomes mostly present, the midrib and lateral veins usually sparsely appressed-pubescent ............ G. blanchetiana

Gouania blanchetiana Miq., Linnaea 22: 797. 1849. —Bejuco reumo, Jaboncillo, Reuma, Wayo wayo (Yanomami). Liana climbing over shrubs or high-climbing; flowers slightly fragrant. Evergreen lowland to lower montane forests, riparian forests, semideciduous forests, forests clearings, 100–700 m; Delta Amacuro (east of El Palmar near Río Grande), Bolívar (Altiplanicie de Nuria, middle Río Caura), Amazonas (upper Río Orinoco, Río Puruname). Guyana, Suriname, French Guiana, Brazil, Bolivia. The local name Jaboncillo refers to the use of Gouania blanchetiana and other species of the genus as a shampoo and soap substitute. Gouania colurnifolia Reiss. in Mart., Fl. Bras. 11(1): 107. 1861. —Bejuco reuma, Rastrojero. Climbing over plants 3 m high or more; leaves usually densely pubescent beneath, the margins with large rounded teeth; fruit densely pubescent with spreading trichomes. Deciduous and semideciduous forests, savannas, margins of streams, disturbed areas, 100–300 m; Bolívar (near El Manteco, Isla Anacoco, Tumeremo, Upata). Widespread elsewhere in Venezuela north of the Río Orinoco; Colombia, Trinidad-Tobago, Guyana, Ecuador, Brazil.

Gouania cornifolia Reiss. in Mart., Fl. Bras. 11(1): 107. 1861. Vine; flowers white; fruit with narrow wing. Banks of rivers in evergreen lowland forests, 50–200 m; Amazonas (between mouth of Río Casiquiare and Culimacare). Peru, Amazonian Brazil, Bolivia. Gouania discolor Benth., Hooker’s J. Bot. Kew Gard. Misc. 4: 12. 1852. Gouania hypochroa Reiss. in Mart., Fl. Bras. 11(1): 106. 1861. Gouania ulei Pilger, Notizbl. Königl. Bot. Gart. Berlin 6: 315. 1915. Climbing plant; flowers white or greenish white. Riparian forests, gallery forests, disturbed areas, 100–900 m; Bolívar (Gran Sabana near Santa Elena and Icabarú, near Río Parguaza), Amazonas (Venezuela-Brazil border on Sierra Tapirapecó). Apure; Brazil. Gouania discolor is sometimes misidentified as G. hypoglauca Standl. Gouania frangulifolia (Willd. ex Roem. & Schult.) Radlk., Sitzungsber. Math.Phys. Cl. Königl. Bayer. Akad. Wiss. München 8: 393. 1878. —Trisecus frangulaefolius Willd. ex Roem. & Schult., Syst. Veg. 6: 641. 1820. Large woody vine; flowers white. Riparian

Rhamnus 479

and evergreen lowland forests, semideciduous forests, ca. 50–900 m; Bolívar (Peraitepui, north of Puerto Ayacucho, middle Río Orinoco, Río Parguaza), Amazonas (Río Orinoco at Santa Bárbara and Tamatama). French Guiana, Amazonian Brazil. Gouania polygama (Jacq.) Urb., Symb. Antill. 4: 378. 1910. —Rhamnus polygamus Jacq., Enum. Syst. Pl. 17. 1760. Gouania tomentosa Jacq., Select. Stirp. Amer. Hist. 263. 1763. Gouania pubescens Poir. in Lam., Encycl. suppl. 2, 819. 1811 [1812]. Vining shrub or high-climbing liana. Gallery forests, deciduous to evergreen lowland forests, disturbed areas, near sea level to 300 m; Delta Amacuro (Isla Guara, Sierra Imataca), Bolívar (near Anacoco, Río Botanamo, Río Caura, Río Paragua, Serranía de Imataca). Common elsewhere in Venezuelan north of the Río Orinoco; Mexico, Central America, West Indies, widespread in South America. ◆Fig. 402. Gouania polygama has frequently been

misidentified as G. lupuloides (L.) Urb. It is readily distinguished in fruit by the wings extended horizontally from the body of the fruit, resulting in a fruit broader than tall, the wings broader than the body of the fruit. In G. lupuloides the fruits are narrowly winged, with the wings narrower than the body of the fruit. In the flowering stage, G. polygama has the disk glabrous with the annulus setulose, whereas G. lupuloides has both disk and annulus glabrous. Gouania wurdackii Steyerm., Ann. Missouri Bot. Gard. 75: 1065. 1988. Woody vine; flowers cream-colored or white; disk and annulus pubescent, tomentose. Igneous substrates in deciduous or seasonally dry evergreen forests, disturbed open areas, 50–300 m; Bolívar (Cerro Pijiguao of Serranía Suapure, Cerro San Borja along Río Suapure, middle Río Orinoco), Amazonas (near Puerto Ayacucho). Endemic. Gouania wurdackii has been previously confused with both G. polygama and G. mollis Reiss.

4. RHAMNUS L., Sp. Pl. 193. 1753. Trees or shrubs, usually unarmed, deciduous or evergreen. Leaves alternate, rarely opposite, pinnately veined, entire or toothed; stipules small, deciduous. Inflorescence axillary, of umbellate cymes or reduced to a single flower, sessile or pedunculate. Flowers bisexual or polygamo-dioecious, small, greenish, sessile or pedunculate, fasciculate, racemose, cymose, or umbellate; disk filling the calyx tube. Calyx 4- or 5-lobed, lobes triangular-ovate, tube urceolate; petals 4, 5, or more, inserted on the margin of the disk, shorter than the calyx lobes, flat or cucullate, often unguiculate, margin entire or bilobed. Stamens 4 or 5, inserted on the upper margin of the calyx tube. Ovary free, 2- or 3-locular; ovules 2 or 3; style simple or 2- or 3-lobed. Fruit dark red to black, a berry-like drupe, subglobose or oblong, with 2 or 3 bony or cartilaginous nutlets. Seeds lenticular, obovoid, smooth or dorsally sulcate; testa membranaceous or crustaceous; endosperm fleshy. Temperate and tropical regions, mostly in eastern Asia and southwestern North America; ca. 150 species, 9 in Venezuela, 8 of these in the flora area. Key to the Species of Rhamnus 1.

1.

Leaf blades strongly discolored with a buff, pale brown, or gray lower surface and tawny or ferruginous veins; lower surface of leaves moderately to densely pubescent throughout ................................................. 2 Leaf blades concolorous or nearly so, the veins on the lower surface not tawny or ferruginous; lower surface of leaves glabrous, only the midrib beneath sometimes sparsely pale pubescent ........................................ 4

480

2(1).

2.

3(2).

3.

4(1). 4. 5(4).

5.

6(5). 6. 7(6).

7.

R HAMNACEAE

Inflorescence with a peduncle 1–2.5 cm long; lower surface of leaf blades minutely pubescent; tertiary venation somewhat prominent below; lateral veins of upper surface of leaves shallowly sulcate and shallowly reticulate between lateral veins .................................... R. marahuacensis Inflorescence epedunculate or with a peduncle < 0.6 cm; lower surface of leaf blades densely stellate tomentellose; tertiary venation mainly concealed by the tomentum; lateral veins of upper surface of leaves conspicuously sulcate and rugulose between lateral veins ....................... 3 Ovary glabrous; leaf margin obscurely crenulate; older petioles and most of the young branchlets glabrous; outer surface of calyx with short, appressed trichomes; plants of Bolívar ................................. R. chimantensis Ovary tomentose; leaf margin entire or essentially so; petioles, stems, and outer surface of calyx densely brown-pubescent; plants of Amazonas .............................................................................. R. sipapoensis Ovary pubescent; marginal teeth of leaf blades averaging 7–9 per cm; plants of Bolívar ................................................................................R. ulei Ovary glabrous; marginal teeth of leaf blades averaging 3–5 per cm; plants of Amazonas ................................................................................ 5 Apex of leaf blades mainly rounded or broadly obtuse; leaf blades usually obovate, broadest above the middle, obovate; tertiary venation of lower surface of leaf blade prominently elevated .......................... R. neblinensis Apex of leaf blades short- to long-acuminate; leaf blades oblong- or lanceelliptic to ovate, broadest near or below the middle; tertiary venation of lower surface of leaf blade inconspicuous or at least not prominently elevated .................................................................................................. 6 Acumen of the larger leaf blades 12–15 mm long; larger leaf blades 12– 16 × 4.5–6 cm; lateral veins mainly 9–11 each side ............. R. acuminata Acumen of the larger leaf blades 3–10 mm long; larger leaf blades 3.5–9 × 1.5–4 cm; lateral veins 6–8 each side ................................................... 7 Inflorescence or infructescence conspicuously pedunculate, the peduncles 10–15 mm long and pedicels 10–15 mm long; leaf margins subrevolute, obscurely shallowly repand-crenulate; tertiary venation of lower surface of leaves subelevated with manifest reticulation ............. R. longipes Inflorescence or infructescence ± epedunculate; leaf margins finely serrulate with 3–5 subsalient acuminate teeth per cm; tertiary venation of lower surface of leaves inconspicuous .................................. R. psilocarpa

Rhamnus acuminata Maguire & Steyerm., Mem. New York Bot. Gard. 51: 121. 1989. Shrub ca. 1.5 m tall; petioles 10–25 mm long; leaves crenulate-serrulate; peduncle 1– 4 cm long. Dwarf tepui slope forests, 1000– 1200 m; Amazonas (summit of Sierra de la Neblina). Endemic. A recent collection from granitic domes at Caño Piedra on Cerro Sipapo, in the Cuao– Sipapo Massif at 1500 m, Sanoja et al. 2926 (MO, PORT), appears to belong to this species as well.

Rhamnus chimantensis Steyerm. & Maguire, Mem. New York Bot. Gard. 17(1): 451. 1967. Shrub 0.5–3 m tall; petioles 2–4 mm long; leaf blades 1.5–4 × 0.8–2.7 cm, strongly revolute; fruit pale or dull brown. Open swamps with Chimantaea mirabilis, borders of dwarf forests, sandstone grottos and labyrinths, along small caños of dwarf forests, 1800– 2500 m; Bolívar (summits of Macizo del Chimantá and Auyán-tepui). Endemic. ◆Fig. 403.

Rhamnus 481

Although treated as a synonym of Rhamnus goudotiana Triana & Planch. (Fl. Neotrop. 20: 56. 1978), this is one of several endemic species of the genus, distinguished from R. goudotiana of the Colombian and Venezuelan Andes by the glabrous ovary, sericeous-tomentose outer surface of the calyx tube and lobes, small, oblong, strongly veined leaves densely tomentose on the lower surface with entire to remotely crenulate margins. Rhamnus longipes Steyerm., Ann. Missouri Bot. Gard. 75: 1066. 1988. Shrub 2 m tall; petiole 4–8 mm long; fruit globose, 5–7 mm long. Open, rocky savannas on tepui slopes and summits, ca. 2000 m; Amazonas (Cerro Parú along tributary of Río Asisa). Endemic. Rhamnus marahuacensis Steyerm. & Maguire, Acta Bot. Venez. 14: 22, fig. 12. 1984. Tree ca. 3 m tall; petioles 4–11 mm long; leaf blades 2–6 × 1.5–3.5 cm; petals bilobed at apex; fruit purple, 6–7 mm long. Dwarf forests and swampy areas on tepuis summits, 2400–2500 m; Amazonas (Cerro Marahuaka). Endemic.

mm long; leaf blades oblong or ovate-oblong, 2.5–7.5 × 1.5–3.8 cm, the lower surface pale buff-velutinous; petals bilobate; fruit dull brownish red. Dwarf forests bordering rocky sandstone slopes, 1900–2600 m; Amazonas (Cerro Marahuaka, Cerro Sipapo). Endemic. Rhamnus ulei Pilger, Notizbl. Königl. Bot. Gart. Berlin 6: 313. 1915. Shrub or tree 2–10 m tall; petioles 5–21 mm long; leaf blades lance-elliptic, oblong-elliptic, to elliptic-ovate, 6–17 cm long, 2–6 cm wide; petals cream-colored, suffused with pale lavender or olive green; fruit red to black, globose, 7–8 mm long. Dense mossy forests, wet montane forests, bases of sandstone escarpments, dwarf thickets on tepuis summits and slopes, 1100–2300 m; Bolívar (Amaruay-tepui, Carrao-tepui near Ptaritepui, Macizo del Chimantá, Ptari-tepui, Río Cuyuni basin, Roraima-tepui, Sororopántepui). Endemic. ◆Fig. 404. The Schomburgk collection cited from Mount Roraima (Fl. Neotrop. 20: 78. 1978) was collected on the Venezuelan side. Although the stipules are recorded as being persistent (Fl. Neotrop. 20: 77. 1978), most of the specimens cited do not have persistent stipules.

Rhamnus neblinensis Maguire & Steyerm., Mem. New York Bot. Gard. 51: 119. 1989. Shrub 1–2 m tall; petioles 12–23 mm long; leaf blades crenate with numerous rounded crenations; flowers whitish, petals bilobed at apex; fruit dark purple. Dwarf thickets on ridges and escarpments, borders of dwarf forests, 1600–2000 m; Amazonas (Sierra de la Neblina). Endemic. Rhamnus psilocarpa Maguire & Steyerm., Mem. New York Bot. Gard. 51: 119. 1989. Shrub 1–2.5 m tall; petioles 3–8 mm long; leaf blades 3.5–6.5 × 1.5–2.5 cm; petals white, bilobed at apex. Tepui slope forests along streams, borders of sandstone escarpments among dwarf vegetation, 800–2000 m; Amazonas (summit of Sierra de la Neblina). Endemic. Rhamnus sipapoenis Steyerm., Ann. Missouri Bot. Gard. 75: 1066. 1988. Shrub or tree ca. 3 m tall; petioles 4–10

Fig. 403. Rhamnus chimantensis

482

R HAMNACEAE

Fig. 404. Rhamnus ulei

5. ZIZYPHUS Mill., Gard. Dict. Abr. ed. 4. 1754. Shrubs or small trees, rarely subclimbing, mainly deciduous, frequently with stipular spines. Leaves alternate, rarely subopposite, subdistichous, usually 3–5veined from the base, entire or crenate. Inflorescence usually axillary, usually cymose, few-flowered. Flowers bisexual, small, greenish, yellow-green, or cream-colored; disk surrounding the ovary. Calyx 4- or 5-lobed; lobes deciduous, tube broadly obconic; petals 4 or 5 or rarely absent. Stamens 4 or 5, concealed within the petals or longer than them. Ovary inferior to subinferior at anthesis, superior in fruit, 2- or 3locular; ovule solitary in the locule; styles 2 or 3, free or connate; stigmas 2 or 3. Fruit a fleshy drupe, globose or oblong, the stone woody or bony, 1–3-locular. Seeds 1–3, plano-convex, ellipsoid, smooth; endosperm absent or scant. Pantropics and subtropics; 86 species, 3 in Venezuela, all in the flora area. Key to the Species of Zizyphus 1. 1.

2(1).

2.

Large trees of evergreen forests; branches without spines; leaves coriaceous, narrowly elliptic, with entire margins .................. Z. cinnamomum Small to medium trees of semideciduous forests or else escaped from cultivation; branches with stipular spines; leaves chartaceous, elliptic to subrotund, margins denticulate or serrulate ....................................... 2 Leaves strongly discolored, the lower surface of leaves, young stems, petioles, and inflorescence with a dense tan or cream-colored pubescence; introduced exotic ................................................................... Z. mauritiana Leaves nearly concolorous the lower surface of leaves glabrous or slightly pubescent, not covered with a dense tan or cream-colored pubescence; young stems, petioles, and inflorescence slightly pubescent to glabrous; native .................................................................................... Z. saeri

Zizyphus 483

Fig. 405. Zizyphus mauritiana

Fig. 406. Zizyphus saeri

484

R HAMNACEAE

Zizyphus cinnamomum Triana & Planch., Ann. Sci. Nat. Bot. sér. 5, 16: 380. 1872. —Pashomacasi, Pashonema-casiji-ajuji (Yanomami). Tree to 30 m tall; branches spineless; leaves narrowly elliptic, coriaceous. Riparian forests, 100–200 m; Amazonas (Río Orinoco at mouth of Río Orinoquito). Colombia, Guyana, French Guiana, Ecuador, Peru, Bolivia.

ests, 50–300 m; Bolívar (El Tigre, Isla Anacoco, near Tumeremo), Amazonas (near Puerto Ayacucho). Introduced and cultivated throughout Venezuela; widespread in tropical America, native to southern Asia and Africa. ◆Fig. 405. The sweet-tasting fruit of Zizyphus mauritiana is edible and is also used medicinally.

Zizyphus mauritiana Lam., Encycl. 3: 319. 1789. —Ponsigué. Shrub or tree 2–6 m tall; leaves suborbicular, oval, or oblong, 4–6 × 2–4 cm, tan-pubescent on lower surface; flowers pale green, yellowish, or whitish; fruit usually orange-red, subglobose to oblong, 12–20 mm diameter. Open fields, roadsides, abandoned cultivated areas, occasionally near cut-over moist for-

Zizyphus saeri Pittier, Bol. Minist. RR. EE. no. 12, reprinted in Arb. Arbust. Venez. 4/5: 61. 1926. —Limoncillo, Mara, Nirgua. Tree 5–25 m tall; fruit green, ovoid, edible. Semideciduous forests, gallery forests, 50– 200 m; Bolívar (El Palmar, Río Oronata). Northern and central Venezuela; northeastern Colombia. ◆Fig. 406.

RHIZOPHORACEAE by Julian A. Steyermark Trees or shrubs. Leaves opposite or verticillate; stipules interpetiolar, caducous or persistent; blades simple, entire or obsoletely serrate or crenulate. Inflorescence of solitary, axillary flowers, or of axillary to subterminal cymes, fascicles, or racemes; bracts 2 per flower or absent. Flowers actinomorphic, 4–7-merous, bisexual or unisexual. Calyx campanulate or short-tubular, 4–many-parted, lobes valvate; petals free, usually as many as calyx lobes, sometimes inserted at base of a nectary disk, fimbriate, lacerate, or entire, each petal individually enclosing 1–5 stamens. Stamens 8–40. Ovary superior to inferior, 2–6-locular; style simple, bidentate, 3- or 4lobulate, or radially fimbriate; ovules 2 per locule, pendulous, anatropous to hemitropous. Fruit a capsule or berry. Seeds 1–many, pendulous, sometimes arillate or winged; endosperm fleshy and oily. Pantropics; 15 genera and ca. 125 species, 3 genera and 8 species in the flora area.

Cassipourea 485

Fig. 407. Cassipourea guianensis

486

R HIZOPHORACEAE

Key to the Genera of Rhizophoraceae 1.

1.

2(1). 2.

Plant of mangrove coastal swamps and lower reaches of rivers, in salt or brackish water; prop roots present; seeds germinating while fruit still attached to tree; ovary half-inferior; calyx deeply parted, the tube much shorter than the lobes; petals entire or nearly so, the margins villous, emarginate at apex .................................................. 2. Rhizophora Plant of solid ground, not in swamps; prop roots absent; seeds germinating on ground after dispersal; ovary superior; calyx dentate or lobed, the tube equaling or shorter than the lobes; petals fimbriate-lacerate, 3-lobed or -parted at apex ..................................................................... 2 Flowers bisexual, solitary or in fascicles; stamens (8–)15–40 ................... .............................................................................................. 1. Cassipourea Flowers unisexual, in cymes with peduncles 1–3 cm long; stamens mainly 10 .................................................................................. 3. Sterigmapetalum

1. CASSIPOUREA Aubl., Hist. Pl. Guiane 529, t. 211. 1775. Trees or shrubs. Leaves opposite; stipules caducous; blades entire or slightly dentate. Flowers bisexual, axillary, solitary or fasciculate, sessile or pedicellate, ebracteolate. Calyx campanulate, 4–7-lobed, the lobes triangular, erect; petals 4 or 5, clawed, fimbriate-lacerate, the lateral flanges with numerous appendages, inflexed, whitish. Stamens 8–40, equaling the calyx or slightly exserted, attached to or outside the disk. Ovary superior, 3- or 4-locular; style elongated, exserted; stigma 3- or 4-lobed or capitate. Fruit a fleshy, 3- or 4-locular septifragal capsule; seeds 1 or 2 per locule, arillate. Neotropics, Africa (where most diverse), Madagascar, Sri Lanka, India; ca. 60 species, 2 in Venezuela, 1 of these in the flora area. Cassipourea guianensis Aubl., Hist. Pl. Guiane 529, t. 211. 1775. Shrub or tree 3–25 m tall. Evergreen or semideciduous forests along streams, disturbed forests, near sea level to 1000 m;

widespread in Delta Amacuro, Bolívar, and Amazonas. Trinidad-Tobago, Guyana, Suriname, French Guiana, Amazonian Brazil. ◆Fig. 407.

2. RHIZOPHORA L., Sp. Pl. 443. 1753. Glabrous trees or shrubs with large prop roots. Leaves opposite; stipules large, caducous; blades thick-coriaceous, entire. Inflorescence axillary, bracteate, with solitary flowers or in few- to many-flowered, pedunculate, dichotomous cymes. Flowers coriaceous, bisexual, bibracteolate. Calyx deeply 4-parted, persistent in fruit; petals 4, lanceolate, white, caducous, the margins entire, villous. Stamens 8– 12, epipetalous. Ovary half-inferior, 2-locular, surrounded by a fleshy disk; style subulate, stigma bidentate; ovules 2 per locule. Fruit a coriaceous berry. Seed 1 by abortion, germinating on the tree; hypocotyl elongate. Pantropics, along seashores; 4, or sometimes 8 species recognized, 3 in Venezuela, all in the flora area. The bark is important in the tanning industry and is used locally for medicine. The wood is hard and durable, much used for construction and charcoal. Key to the Species of Rhizophora 1.

Inflorescence unbranched, 1–4-flowered; pedicels 7–20 mm long; hypocotyl of germinating seed 15–20(–35) cm long ........................... R. mangle

Rhizophora 487

1. 2(1). 2.

Inflorescence branched, several- to many-flowered; pedicels 3–11 mm long; hypocotyl of germinating seed 11–65 cm long ............................. 2 Hypocotyl 25–65 cm long; stigmas 2–4; pedicels 3–5 mm long; floral bud and peduncle thick ................................................................... R. racemosa Hypocotyl 15–30 cm long; stigmas 2; pedicels 3–11 mm long; floral bud and peduncle slender .............................................................. R. harrisonii

Fig. 408. Rhizophora mangle

488

R HIZOPHORACEAE

Fig. 409. Rhizophora racemosa

Rhizophora harrisonii Leechm., Kew Bull. 1918: 8. 1918. —Mangle rojo. Tree 15–35 m tall with descending aerial roots to 20 m long. Locally dominant in mangrove forests at sea level; Delta Amacuro (Caño Angosturita, southeast of Pedernales). Monagas; Trinidad-Tobago, Guyana, Suriname, French Guiana, western Africa.

Rhizophora mangle L., Sp. Pl. 443. 1753. —Cascarón, Mangle, Mangle blanco, Mangle de burro, Mangle negro, Mangle rojo. Tree 3–7 m tall. Mangrove forests at sea level, Delta Amacuro (common). Scattered along northern coast of Venezuela; U.S.A. (Florida, lower California), widespread in the

Sterigmapetalum 489

Neotropics, western Africa, Pacific Islands. ◆Fig. 408. Rhizophora racemosa G. Mey., Prim. Fl. Esseq. 185. 1818. —Rhizophora mangle var. racemosa (G. Mey.) Engl. in Mart., Fl. Bras. 12(2): 427. 1876. —Buju (Waroa), Mangle rojo.

Tree 5–15 m tall. Mangrove and riparian forests, in salt to brackish water, ranging farther inland than the other 2 species, sea level to 50 m; Delta Amacuro (common). Monagas, Sucre; Guyana, Suriname, French Guiana, Ecuador, Brazil, more common in western Africa. ◆Fig. 409.

3. STERIGMAPETALUM Kuhlm., Arch. Jard. Bot. Rio de Janeiro 4: 359, t. 32. 1925. Trees. Leaves opposite or 3- or 4-verticillate; stipules generally persistent; blade entire or crenulate-serrulate. Inflorescence axillary or subterminal, pedunculate, cymose, unbranched or dichotomously 1–4-branched. Flowers unisexual (plants dioecious), actinomorphic, few to numerous, white or creamy-white. Calyx campanulate, 4–7-dentate or -lobed; petals 4–6, inserted between base of calyx tube and base of staminal membrane, 3-lobed or 3-parted at apex, with 2 lateral flanges. Stamens 8–12, mostly unequal. Ovary superior, 5- or 6-locular; style very short or obsolete, the short branches with many stigmatic threads. Fruit a septicidal 5- or 6locular capsule. Seeds winged; endosperm oily. Colombia, Venezuela, Guyana, Suriname, Amazonian Brazil; 9 species, 6 in Venezuela, 4 of these in the flora area. Key to the Species of Sterigmapetalum 1.

1. 2(1). 2. 3(2).

3.

Leaves opposite; resinous exudate present on stem apex, peduncle, and buds; peduncle axillary at the 2nd to 6th node below stem apex ................................................................................................. S. resinosum Leaves 3- or 4-verticillate; resinous exudate absent; peduncle situated at or just below stem apex ......................................................................... 2 Leaf margin entire; leaf broadest above the middle ................. S. guianense Leaf margin crenate or denticulate-crenulate; leaf broadest at the middle ................................................................................................................ 3 Leaves completely covered by yellow indumentum on lower surface; stipules triangular-acuminate, 3–5 mm long; petals with fimbrillate lateral flanges, not externally papillate ......................... S. chrysophyllum Leaves moderately to densely appressed-sericeous on lower surface; stipules ovate-lanceolate, subobtuse, 7–10 mm long; petals externally papillate, without fimbrillate lateral flanges ........... S. exappendiculatum

Sterigmapetalum chrysophyllum Aymard & Cuello, Novon. 5: 223. 1995. —Kasemani, Tadiji (Yekwana). Tree 15–20 m tall. Montane forests on sandstone outcrops, 1200–1300 m; Amazonas (unnamed mountains in upper Río Ventuari basin west of Cerro Parú). Endemic. Sterigmapetalum exappendiculatum Steyerm. & Liesner, Ann. Missouri Bot. Gard. 70: 188. 1983.

Tree 12–20 m tall. Montane forests on sandstone, 1000–1700 m; Bolívar (Macizo del Chimantá, Sierra de Lema and adjacent Gran Sabana). Endemic. Sterigmapetalum guianense Steyerm., Fieldiana, Bot. 28: 422. 1952. Tree 4–20 m tall. 100–1600 m. Venezuela, Suriname; 2 subspecies, both in the flora area.

490

R HIZOPHORACEAE

Fig. 410. Sterigmapetalum guianense subsp. ichunense

Key to the Subspecies of S. guianense 1. Leaf blade with prominent and subreticulate tertiary venation on upper surface .......................... subsp. guianense 1. Leaf blade lacking prominent or subreticulate tertiary venation on upper surface .......................... subsp. ichunense S. guianense subsp. guianense Tree 15–20 m tall. Dwarf or talus forests on tepuis, ca. 1600 m; southeastern Bolívar (Ptari-tepui, Sororopán-tepui). Suriname (Tafelberg). S. guianense subsp. ichunense Steyerm. & Liesner, Ann. Missouri Bot. Gard. 70:

188. 1983. Tree 4–20 m tall. Upland forests on sandy soils, shrub patches bordering white-sand savannas, granitic outcrops, 100–500 m; Bolívar (Sierra Ichún), southwestern Amazonas. Endemic. ◆Fig. 410. The collections of subsp. ichunense from Amazonas may represent a different taxon than the Bolívar one. Sterigmapetalum resinosum Steyerm. & Liesner, Ann. Missouri Bot. Gard. 70: 188. 1983. Tree 20–25 m tall. Montane forests on sandstone, 1000–1300 m; Bolívar (headwaters of Río Venamo). Endemic.

ROSACEAE by Gerardo A. Aymard C. and Nidia L. Cuello A. Trees, shrubs, perennial or sometimes annual herbs, occasionally with thorns or prickles. Leaves alternate, rarely opposite, simple or compound; stipules usually

Hesperomeles 491

present and paired, sometimes adnate to the petioles. Flowers usually bisexual, actinomorphic or rarely zygomorphic, solitary or in various types of ± cymose inflorescences. Perianth biseriate or one or both series absent; calyx often perigynous, with (3–)5(–10) basally connate, imbricate sepals; petals (3–)5(–10), free, imbricate or rarely contorted, or absent. Stamens few to numerous, often in sets of 5 or 10 in 1– several whorls, free or rarely connate; filaments slender; anthers small, 2-locular, opening by longitudinal slits, rarely by terminal pores. Ovary superior or inferior, carpels 1–many, simple or compound with 2–5 locules; ovules 2–few, basal or axile, often adnate to the receptacle; styles terminal, as many as the carpels, sometimes connate. Fruits various, commonly of separate follicles or achenes, these sometimes on a fleshy receptacle, or of laterally coherent drupelets, or a pome. Seeds usually without endosperm; cotyledons often fleshy and convex, rarely folded or convolute. Cosmopolitan (mostly temperate areas); ca. 100 genera and 3000 species, 3 genera and 10 species in the flora area. Key to the Genera of Rosaceae 1. 1.

2(1). 2.

Trees; leaves with 2 glands near the base of the blade or near the midvein in the lower half of the blade; fruit a drupe ............................... 2. Prunus Small trees, shrubs, or trailing woody vines; leaves without glands; fruit a pome or an aggregate of numerous drupelets united on enlarged receptacle ................................................................................................... 2 Leaves simple, rigid-coriaceous, crenate; inflorescences corymbose cymes 2–5 cm long; fruit a small pome ........................................ 1. Hesperomeles Leaves 3- or 5-foliolate, chartaceous to subcoriaceous, dentate-serrate; inflorescences racemes or panicles > 7 cm long; fruit of numerous drupelets united on an enlarged receptacle ................................. 3. Rubus

1. HESPEROMELES Lindl., Edward’s Bot. Reg. 23: sub. pl. 1956. 1837. Evergreen shrubs or small trees, much-branched, the young branchlets often spine-tipped. Leaves simple, often coriaceous, 3–6 cm long or less, crenate or dentate. Inflorescences usually of small terminal corymbose cymes. Flowers 5-merous, hypanthium campanulate or turbinate, 5-lobed. Petals usually obovate-oblong, pink or white. Ovary inferior at anthesis, carpels 5, distinct from each other from the flowering stage on but attached parietally at maturity with bony endocarp, enlarged and often somewhat exserted from the fleshy hypanthium; ovule 1 or rarely 2 per locule. Stamens 10–20; disk present. Fruit a small pome, < 1 cm long, with persistent sepals. Cotyledons accumbent. Neotropics; 20 species, 8 in Venezuela, 1 of these in the flora area. Hesperomeles obtusifolia (Pers.) Lindl., Bot. Reg. 23, subpl. 1956. 1837. —Crataegus obtusifolia Pers., Syn Pl. 2: 37. 1806. Hesperomeles heterophylla (Ruiz & Pav.) Hook., Icon. Pl. 9: pl. 846. 1852. —Mespilus heterophylla Ruiz & Pav., Fl. Peruv. 4: pl. 425b. 1802. Neotropics; 2 varieties, 1 in Venezuela.

H. obtusifolia var. obtusifolia Small shrub to 1 m tall; flowers white; fruit red. Low forests on tepui summits, 2000–2500 m; Bolívar (Ilú-tepui, Macizo del Chimantá [Apacará-tepui]). Anzoátegui, Distrito Federal, Lara, Mérida, Táchira, Trujillo; Costa Rica, Panama, Colombia, Ecuador, Peru, Bolivia. ◆Fig. 411.

492

R OSACEAE

Fig. 411. Hesperomeles obtusifolia var. obtusifolia

2. PRUNUS L., Gen. Pl. ed. 5, 213. 1754. Deciduous or evergreen trees or shrubs; bark reddish brown, thin, smooth, easily peeled off in layers, the lenticels pale, usually elongating transversely. Leaves simple, alternate, with glands usually occurring on petioles or at base of leaf blades; stipules paired, free from the petiole, small, early deciduous; margins usually serrate, sometimes entire or spine-toothed, prominent, various sized and shaped. Inflorescences axillary, racemose, corymbose, or of solitary flowers. Flowers bisexual, actinomorphic. Calyx 5-lobed, ebracteolate, the lobes triangular, imbricate in bud, the tube perigynous, cup-like, forming a hypanthium with the receptacular disk; petals 5, borne on the margin of hypanthium, white to pink. Stamens (10–)15–20 (–80) in 2 or more whorls and in multiples of 5, perigynous at the outer edge of the floral tube; anther small, dorsifixed; filaments free, filiform or somewhat dilated at base. Ovary 1, free from the calyx tube; style usually peltate; stigma truncate; ovules 2, collateral. Fruit a globose to ellipsoid 1-seeded drupe; mesocarp fleshy and indehiscent, to dryish and dehiscent, usually edible; endocarp hard, indehiscent, nearly globose or compressed, smooth or textured. Seed filling stone. Cosmopolitan (mostly of the northern temperate regions); 250 species, 13 in Venezuela, 6 of these in the flora area. Key to the Species of Prunus 1. 1.

Leaves lanceolate to lanceolate-elliptic; base of inflorescence rachis 3– 4 mm diameter; pedicels 7–10 mm long ................................... P. lichoana Leaves oblong, ovate, ovate-elliptic, to elliptic; base of inflorescence rachis 1–1.5 mm diameter; pedicels 2–5 mm long ................................... 2

Prunus 493

2(1). 2. 3(2). 3. 4(2). 4. 5(4). 5.

Leaves ovate to ovate-elliptic .................................................................... 3 Leaves oblong, ovate-oblong, to elliptic .................................................... 4 Leaves papery, when dried, 5–10 cm long; inflorescences 3–4 cm long ............................................................................................... P. accumulans Leaves coriaceous, 10–20 cm long; inflorescences 5–9 cm long ............ ................................................................................................. P. amplifolia Leaves ovate-oblong; petioles 8–14 mm long .............................. P. wurdackii Leaves elliptic; petioles 4–6 mm long ....................................................... 5 Leaves thick-coriaceous, 9–12 × 4–10 cm, base acute .............. P. espinozana Leaves chartaceous to subcoriaceous, 3–7 × 2–3 cm, base rounded .................................................................................................. P. myrtifolia

Prunus accumulans (Koehne) C.L. Li & Aymard, BioLlania Edición Especiál 6: 449. 1997. —Prunus myrtifolia var. accumulans Koehne, Bot. Jahrb. Syst. 52: 320. 1915. —Menta-berti, Uakadame (Arekuna). Tree 10–25 m tall; fruit black. Evergreen lowland to lower montane forests, 100–1000 m; Bolívar (Altiplanicie de Nuria, Gran Sabana, La Yagua, upper Río Caroní, Río Mawela-tributary of the Río Erebato, Ser-

Fig. 412. Prunus myrtifolia var. myrtifolia

ranía de Imataca). Falcón; Guyana, Suriname, French Guiana. Prunus amplifolia Pilger, Bot. Jahrb. Syst. 37: 538. 1906. —Majada (Yekuana). Tree 8–20 m tall; fruit black. Evergreen lowland to montane forests, 100–1200 m; Amazonas (Río Casiquiare, Río Mawarinuma, middle Río Ventuari, San Carlos de Río Negro). Amazonian Peru, Brazil (Acre, Amazonas), and Bolivia.

494

R OSACEAE

Fig. 413. Prunus wurdackii

Prunus espinozana C.L. Li, BioLlania Edición Especiál 6: 450. 1997. Tree to 20 m tall with green bark. Tepui forests, 1800–2000 m; Bolívar (Cerro Jaua). Endemic. Prunus lichoana Aymard, BioLlania Edición Especiál 6: 451. 1997.

Tree 7–10 m tall; fruits black. Forested tepui slopes, 1000–1300 m; Bolívar (Cerro Marahuaka). Colombia (Antioquia). The Colombian collection is well out of altitudinal and geographical range of the type locality, but the specimens are morphologically alike.

Rubus 495

Prunus myrtifolia (L.) Urb., Symb. Antill. 5: 93. 1904. —Celastrus myrtifolius L., Sp. Pl. 196. 1753. Prunus sphaerocarpa Sw., Prodr. 80. 1788. Neotropics; 3 varieties, 1 in Venezuela. P. myrtifolia var. myrtifolia. —Guanabanillo. Tree 10–15 m tall; fruits purple. Lower montane to montane forests, 500–1300 m; Bolívar (Cerro Impacto, northern Gran Sabana, lower Río Suapure). Anzoátegui,

Aragua, Barinas, Distrito Federal, Falcón, Lara, Mérida, Miranda, Sucre; West Indies, Guyana, Suriname, French Guiana, Brazil (Acre). ◆Fig. 412. Prunus wurdackii C.L. Li, BioLlania Edición Especiál 6: 451. 1997. Tree 3–15 m tall; calyx salmon-colored. Montane forests, tepui slopes, 900–2200 m; Bolívar (La Escalera, Macizo del Chimantá [Toronó-tepui], Ptari-tepui). Endemic. ◆Fig. 413.

3. RUBUS L., Gen. Pl. ed. 5, 557. 1754. Erect, scandent, or climbing shrubs, rarely herbaceous. Stems (canes) usually with spines or thorns. Leaves petiolate, alternate, 3- or 5-foliolate, sometimes a few leaves simple; margins often toothed or entire; stipules persistent or caducous, adnate to base of petioles. Inflorescences racemiform or paniculiform, axillary and usually terminating the canes, pedicels often lengthening in fruit. Flowers actinomorphic, bisexual (or unisexual, the plants then dioecious). Calyx 5-lobed, persistent, sometimes glanduliferous; petals 5, shorter than to exceeding the calyx lobes (or absent), white or rarely pink. Androecium of many stamens inserted at the floral cup, the outer longer than and deshiscent before the inner ones; filaments slender; anthers sometimes pubescent. Carpels numerous, separate, closely packed; styles filiform, subterminal, persistent; stigmas discoid or slightly 2-lobed. Fruit a syncarp composed of small coherent fleshy drupelets, the drupelets falling individually or coalescent and either falling from the receptacle as a unit or with it, the stones hard, variously textured. Seeds filling the stones; embryo small, the radicle superior. Cosmopolitan (most diverse in north-temperate areas); ca. 700 or more species, 11 in Venezuela, 3 of these in the flora area. Key to the Species of Rubus 1. 1. 2(1). 2.

Leaflets elliptic, margins dentate-serrate, glabrate on lower surface ............................................................................................... R. guyanensis Leaflets elliptic-oblong, margins serrate, white appressed-tomentose on lower surface .......................................................................................... 2 Stems, branches, petioles, and petiolules villous-tomentose; lower surface of leaflets appressed-tomentose .................................. R. floribundus Stems, branches, petioles, and petiolules with hispid red trichomes 1–4 mm long; lower surface of leaflets densely white-tomentose ............................................................................................... R. urticifolius

Rubus floribundus Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 219. 1823 [1824]. —Karoráy (Arekuna). Rubus robustus C. Presl, Epimel. Bot. 196. 1849 [1851]. Erect shrub, much-branched; fruit black.

Open areas on tepui slopes and summits, 1200–2300 m; Bolívar (Auyán-tepui, Roraima-tepui, Sororopán-tepui), Amazonas (Sierra de la Neblina). Anzoátegui, Aragua, Distrito Federal, Falcón, Lara, Miranda,

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Fig. 414. Rubus guyanensis

Portuguesa, Sucre, Tachirá, Trujillo; Colombia, Ecuador, Peru, Bolivia, Argentina. Rubus guyanensis Focke, Abh. Naturwiss. Vereine Bremen 4: 160. 1874. Rubus schomburgkii Klotzsch in R.M. Schomb., Reis. Br.-Guiana 3: 1102. 1848 [1849], nom. nud. Sprawling shrub or climbing vine. Open areas or edges of tepui forests, 1700–2400 m; Bolívar (Auyán-tepui, Macizo del Chimantá [Sarvén-tepui], Ptari-tepui, Roraima-tepui). Aragua, Cojedes, Distrito Federal, Miranda, Mérida; Colombia, Guyana. ◆Fig. 414.

Rubus urticifolius Poir. in Lam., Encycl. 6: 246. 1804. Trailing or scandent shrub. Open areas on tepui summits, semi-disturbed areas, 800– 2700 m; Bolívar (Kavanayén, upper Río Cuyuní, Roraima-tepui, Sierra Pakaraima), Amazonas (Cerro Marahuaka). Aragua, Barinas, Distrito Federal, Mérida, Miranda, Portuguesa, Tachirá, Yaracuy; Mexico, Central America, Colombia, Ecuador, Peru, Brazil, Paraguay.

R U B I A C E A E 497

RUBIACEAE by Charlotte M. Taylor, Julian A. Steyermark, Piero G. Delprete, Alberto Vincentini, Rocio Cortés, Daniela Zappi, Claes Persson, Cristina Bestetti Costa, and Elisete Araujo da Anunciação Trees, shrubs, annual or perennial herbs, vines, or lianas, terrestrial or sometimes epiphytic, sometimes armed with straight to curved spines, sometimes fire-adapted with soft annual stems from extensive woody underground stems, sometimes with buds resinous, sometimes with swollen hollow stems or leaf bases housing ants, rarely with nectary glands at abaxial petiole bases (Henriquezia, Platycarpum). Leaves opposite or verticillate, rarely appearing alternate by reduction of one of the leaves of each pair (Didymochlamys), sessile to petiolate, entire or rarely undulate (Psychotria) or pinnatifid (Pentagonia, juvenile Simira), pinnately veined to infrequently subtripliveined (Hillia), the tertiary venation sometimes lineolate (i.e., closely parallel, visible on abaxial surface with magnification; also called clathrate), often with foveolate or tufted domatia in the abaxial axils of the secondary veins, in some African species with bacterial nodules immersed in the blade (Psychotria, Pavetta), rarely with pellucid punctations (Rustia); stipules persistent or caducous, interpetiolar and sometimes united around the stem into a sheath, occasionally completely united into a conical cap (Amaioua, Calycophyllum, Duroia, Faramea, Coussarea, Psychotria), in the interpetiolar portion variously entire, bilobed, multifid, lacerate, or setose, the setae often glandular, infrequently the intrapetiolar portion united and the interpetiolar portion splitting so the stipules become intrapetiolar (Elaeagia, Capirona), sometimes the sheath with 1 or 2 lateral appendages, glands, or groups of glands (Rudgea, Notopleura), internally with glandular trichomes (i.e., colleters) usually present at stipule base and sometimes well developed and persistent after the stipules fall (Calycophyllum, Psychotria), or the stipules occasionally expanded into 1–several foliaceous segments resulting in the leaves appearing exstipulate and verticillate (Galium, Limnosipanea, Rubia). Inflorescence thyrsiform, cymose, compound-cymose, or occasionally capitate, racemiform, spicate, corymbose, or with flowers solitary, terminal, axillary (i.e., borne in both axils at each node), or pseudoaxillary (or lateral, i.e., borne in one axil at each node), sessile to pedunculate, bracteate or bracts lacking. Flowers usually hermaphroditic, homostylous or often distylous, or sometimes unisexual on dioecious plants, actinomorphic or rarely zygomorphic, sessile to pedicellate, sometimes the hypanthia of individual flowers in a fused inflorescence (Morinda, Pagameopsis). Calyx gamosepalous and fused to inferior ovary, the limb cupular, tubular, or occasionally obsolete, truncate to 4- or 5(–9)-lobed or sometimes spathaceous (i.e., split along one side), the lobes often slightly to markedly unequal, one of them sometimes enlarged into a petaloid calycophyll; corolla gamopetalous, actinomorphic or rarely zygomorphic, funnelform, salverform, tubular, campanulate, or infrequently rotate or urceolate, small to large and showy, white, yellow, orange, red, blue, purple, or pale green or with colors in various combinations, glabrous or pubescent internally, lobes (3)4–5(–11), in bud imbricate, valvate, contorted, or rarely open in bud, sometimes furnished with rounded to horn-like thickenings or protuberances at or below

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abaxial apex. Stamens adnate to corolla, usually the same number (isomerous) as the corolla lobes, alternate with these, inserted variously in the corolla throat or tube, or infrequently at its base and appearing free (Galium); anthers included or exserted, free, introrse, 2-celled, dorsifixed to basifixed, dehiscing by lateral slits or rarely by apical pores (Rustia), with the connective sometimes shortly prolonged at apex and/or base. Pollen of various types, 3- or 4-porate, 3- or 4(5)-colpate (the usual type), 5–25-colpate with disk-shaped or ellipsoidal grains, tetrahedral tetrads of 3or 4-porate grains, cylindrical grains with 2 pores, or inaperturate. Ovary inferior or rarely superior (Pagamea, Coryphothamnus), (1)2(–10)-locular, crowned by a fleshy, pulvinate, hemispheric, or conic disk, this annular or sometimes 2-parted or lobulate; ovules solitary or paired to numerous in each locule, from basal, axile, apical, or parietal placentas; style short or elongate, simple or 2–10-lobed, rarely 2-parted to the base (Galium), the stigmatic branches filiform, linear, or spathulate, stigmatic within or at the apex. Fruit capsular with dehiscence loculicidal, septicidal, circumscissile, or fleshy and baccate or drupaceous, or schizocarpous with the mericarps indehiscent, fully inferior), partially inferior (Gleasonia, Platycarpium, Henriquezia), superior (Pagamea, Coryphothamnus), or sometimes secondarily elongating above the calyx limb insertion to form a beak (Oldenlandia); pyrenes when present papery to hard, 1–10-locular (i.e., 1 per locule and unilocular to 1 per fruit and multilocular). Seeds 1–numerous, small to large, sometimes winged, smooth to foveolate, tuberculate, or striate; endosperm oily, usually large, fleshy, or corneous, entire or ruminate, rarely absent. Cosmopolitan; ca. 637 genera and 10,800 species, 86 genera and 524 species in the flora area. This is one of the largest families of flowering plants, represented in nearly all parts of the earth, but most abundant in the tropical regions of both hemispheres. Important members of the family are coffee (Coffea), quinine (Cinchona), and various horticultural subjects such as Gardenia, Ixora, Mussaenda, and Pentas. Additional species are expected to be found with increasing exploration of the Guayana Region. Literature Cited Andersson, L. 1995. Tribes and genera of the Cinchoneae complex (Rubiaceae). Ann. Missouri Bot. Gard. 82: 409–427. Andersson, L. 1996. Circumscription of the tribe Isertieae (Rubiaceae). Opera Bot. Belg. 7: 139–164. Andersson, L. 1997. Synopsis of the genus Ladenbergia (Rubiaceae). Nordic J. Bot. 17: 255–299. Andersson, L. 1998. A revision of the genus Cinchona (Rubiaceae–Cinchoneae). Mem. New York Bot. Gard. 81: 1–75. Andersson, L. 2001. Margaritopsis (Rubiaceae, Psychotrieae) is a pantropical genus. Syst. Geogr. Pl. 71: 73–85. Andersson, L. 2002. Relationships and generic circumscriptions in the Psychotria complex. Syst. Geogr. Pl. 72: 167–202. Andersson, L., and B. Ståhl. 1999. 162. Rubiaceae (Part 3), Tribe 8, Isertieae. In: G. Harling and L. Andersson, eds., Flora of Ecuador 62: 57–129. Council for Nordic Publications in Botany, Copenhagen. Andersson, L., and C. M. Taylor 1994. 162(1–4). Rubiaceae–Cinchoneae–

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Coptosapelteae. In: G. Harling and L. Andersson, eds., Flora of Ecuador 50: 1–114. Council for Nordic Publications in Botany, Copenhagen. Bacigalupo, N. M. 1974. Rubiaceae. In: A. Burkart, Flora Ilustrada de Entre Ríos (Argentina), Parte VI: Dicotiledóneas metaclamídeas (Gamopétalas), B: Rubiales, Cucurbitales, Campanulales (Incluso Compuestas), pp. 3–50. Colección Científica del I.N.T.A., Buenos Aires. Bacigalupo, N. M., and E. L. Cabral. 1996. Infrageneric classification of Borreria (Rubiaceae–Spermacoceae) on the basis of American species. Opera Bot. Belg. 7: 297–308. Bacigalupo, N. M., and E. L. Cabral. 1999. Revisión de las especies americanas del género Diodia (Rubiaceae, Spermacoceae). Darwiniana 37: 153–165. Boom, B. M. 1984. A revision of the genus Isertia (Isertieae: Rubiaceae). Brittonia 36: 425–454. Boom, B. M., and P. G. Delprete. 2002. Rubiaceae. In: S. Mori et al., Guide to the vascular plants of central French Guiana, Part 2, Dicotyledons. Mem. New York Bot. Gard. 76: 606–649. Bremekamp, C. E. B. 1934. Notes on the Rubiaceae of Surinam. Recueil Trav. Bot. Néerl. 31: 248–305. Bridson, D. 1988. Coffea. In: D. Bridson and B. Verdcourt, Rubiaceae (Part 2). In: R. M. Polhill, ed., Flora of Tropical East Africa, 747 pp. A. A. Balkema, Rotterdam, The Netherlands. Bridson, D., and E. Robbrecht. 1985. Validation of the African genus Hyperacanthus E. Mey. (Rubiaceae tribe Gardenieae) Kew Bull. 40: 273– 286. Bruza, J. 1982. A Revision of the Diodia teres Complex (Rubiaceae). Ph.D. Dissertation. Mississippi State Univ. Burger, W. C., and C. M. Taylor. 1993. Family # 202, Rubiaceae. In: W. Burger, ed., Flora Costaricensis. Fieldiana, Bot. n.s. 33: 1–333. Clark, H., R. Liesner, P. E. Berry, A. Fernández, G. Aymard, and P. Aquirino. 2000. Catálogo Anotado de la Flora del Área de San Carlos de Río Negro, Venezuela. Scientia Guaianae 11: 101–316. Cortés, R. 2003. Systematics and Biogeography of Retiniphyllum (Rubiaceae). Ph.D. Dissertation. City University of New York. Davis, A., D. Bridson, C. Jarvis, and R. Govaerts. 2001. The typification and characterization of the genus Psychotria L. (Rubiaceae). Bot. J. Linn. Soc. 135: 35–42. Delprete, P. G. 1996. Notes on the calycophyllaceous Rubiaceae. Part I. Morphological comparisons of the genera Chimarrhis, Bathysa, and Calycophyllum, with new combinations and a new species, Chimarrhis gentryana. Brittonia 48: 35–44. Delprete, P. G. 1999a. Riodocea (Rubiaceae, Gardenieae), a new genus from the Brazilian Atlantic forest. Brittonia 51: 15–23. Delprete, P. G. 1999b. Rondeletieae (Rubiaceae)—Part I (Rustia, Tresanthera, Condaminea, Picardaea, Pogonopus, Chimarrhis, Dioicodentron, Molopanthera, Dolichodelphys, and Parachimarrhis). Flora Neotropica Monograph 77: 137–187. New York Botanical Garden Press, New York. Delprete, P. G. 2001. Notes on some South American species of Psychotria subgenus Heteropsychotria (Rubiaceae), with observations on rubiaceous taxonomic characters. Brittonia 53: 396–404.

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Dempster, L. T. 1990. The genus Galium (Rubiaceae) in South America, IV. Allertonia 5: 283–345. Dwyer, J. D. 1980. Rubiaceae. In: R. E. Woodson Jr., R. W. Schery, and Collaborators, Flora of Panama—Part IX. Ann. Missouri Bot. Gard. 67: 1–522. Elias, T. S. 1976. A monograph of the genus Hamelia (Rubiaceae). Mem. New York Bot. Gard. 26(4): 81–144. Greuter, W., J. McNeill, F. R. Barrie, H. M. Burdet, V. Demoulin, T. S. Filgueiras, D. H. Nicolson, P. C. Silva, J. E. Skog, P. Trehane, N. J. Turland, and D. L. Hawksworth. 2000. International Code of Botanical Nomenclature (Saint Louis Code). Königstein, Germany: Koeltz Scientific Books. Gustafsson, C. G. R. 1998. The neotropical Rosenbergiodendron (Rubiaceae, Gardenieae). Brittonia 50: 453–466. Gustafsson, C. G. R. 2000. Three new South American species of Randia (Rubiaceae, Gardenieae). Novon 10: 201–208. Kirkbride, J. H., Jr. 1976. A revision of the genus Declieuxia (Rubiaceae). Mem. New York Bot. Gard. 28(4): 1–87. Kirkbride, J. H., Jr. 1985. Manipulus Rubiacearum—V. A revision of the genus Capirona. Acta Amazon. 15: 47–60. Kirkbride, J. H., Jr. 1997. Manipulus Rubiacearum—VI. Brittonia 49: 354–379. Kirkbride, J. H., Jr., and E. Robbrecht. 1984. Documentation of two recent new generic names in the Rubiaceae. Taxon 33: 102–105. Liogier, A. H. 2000. Flora of Puerto Rico and adjacent islands: A systematic synopsis. Río Piedras, Puerto Rico: Editorial de la Universidad Puerto Rico. Lorence, D. H. 1991. New species and combinations in Mexican and Central American Rondeletia (Rubiaceae). Novon 1: 135–157. Lorence, D. H. 1994. New species in Mexican and Mesoamerican Rubiaceae. Novon 4: 119–136. Nepokroeff, M., B. Bremer, and K. J. Sytsma. 1999. Reorganization of the genus Psychotria and tribe Psychotrieae (Rubiaceae) inferred from ITS and rbcL sequence data. Syst. Bot. 24: 5–27. Nicolson, D. H. 1977. Typification of names vs. typification of taxa: proposals on Article 48 and reconsideration of Mitracarpus hirtus vs. M. villosus (Rubiaceae). Taxon 26: 569–574. Ochoterena Booth, H. 1994. Revisión Taxonómica del Género Coutarea Aublet (Rubiaceae). Thesis. Universidad Nacional Autónoma de México. Persson, C. 1996. Phylogeny of the Gardenieae (Rubiaceae). Bot. J. Linn. Soc. 121: 91–109. Persson, C. 2000a. Phylogeny of the neotropical Alibertia group (Rubiaceae), with emphasis on the genus Alibertia, inferred from ITS and 5S ribosomal DNA sequences. Amer. J. Bot. 87: 1018–1028. Persson, C. 2000b. Phylogeny of the Gardenieae based on chloroplast DNA sequences from the rps16 intron and trnL(UAA)–F(GAA) intergenic spacer. Nordic J. Bot. 20: 257–269. Persson, C. 2003. Agouticarpa, a new neotropical genus of Tribe Gardenieae (Rubiaceae). Brittonia 55: 176–201. Piesschaert, F. 2001. Carpology and Morphology of the Psychotrieae (Rubiaceae– Rubioideae). Dissertation. Katholieke Universiteit Leuven, Belgium. Piesschaert, F., S. Jansen, I. James, E. Robbrecht, and E. Smets. 2001. Morphology, anatomy, and taxonomic position of Pagameopsis (Rubiaceae–

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Rubioideae). Brittonia 33: 490–504. Puff, C., L. Andersson, U. Rohrhofer, and A. Igersheim. 1993. The tribe Schradereae. Bot. Jahrb. Syst. 114: 449–479. Rendle, A. B. 1936. Spermacoce remota Lam. J. Bot. 74: 10–12. Robbrecht, E. 1993. Genera Rubiacearum. Opera Bot. Belg. 6: 173–196. Rogers, G. K. 1984. Gleasonia, Henriquezia, and Platycarpum (Rubiaceae). Fl. Neotrop. Monogr. 39: 1–135. Standley, P. C. 1931. Studies of American plants—V. Publ. Field Columbian Mus., Bot. Ser. 8: 295–398. Steyermark, J. A. 1963. Rubiaceae. In: B. Maguire and J. J. Wurdack and Collaborators, The Botany of the Guayana Highland—Part V. Mem. New York Bot. Gard. 10(5): 186–278. Steyermark, J. A. 1965. Rubiaceae. In: B. Maguire and Collaborators, The Botany of the Guayana Highland—Part VI. Mem. New York Bot. Gard. 12(3): 178– 285. Steyermark, J. A. 1967. Rubiaceae. In: B. Maguire and Collaborators, The Botany of the Guayana Highland—Part VII. Mem. New York Bot. Gard. 17(1): 230– 436. Steyermark, J. A. 1972. Rubiaceae. In: B. Maguire and Collaborators, The Botany of the Guayana Highland—Part IX. Mem. New York Bot. Gard. 23: 227–832. Steyermark, J. A. 1974. Rubiaceae. In: T. Lasser, ed., Flora de Venezuela 9(1–3): 1– 2070. Caracas: Instituto Botánico. Steyermark, J. A. 1984. Flora of the Venezuelan Guayana—I. Ann. Missouri Bot. Gard. 71: 297–340. Steyermark, J. A. 1987. Flora of the Venezuelan Guayana—II. Ann. Missouri Bot. Gard. 74: 85–116. Steyermark, J. A. 1988. Notes on Oldenlandia filicaulis and Oldenlandia tenuis (Rubiaceae). Ann. Missouri Bot. Gard. 75: 736–738. Taylor, C. M. 1993. Revision of Palicourea (Rubiaceae: Psychotrieae) in the West Indies. Moscosoa 7: 201–241. Taylor, C. M. 1994. Revision of Hillia (Rubiaceae). Ann. Missouri Bot. Gard. 81: 571–609. Taylor, C. M. 1996. Overview of the Psychotrieae (Rubiaceae) in the Neotropics. Opera Bot. Belg. 7: 261–270. Taylor, C. M. 1997. Conspectus of the genus Palicourea (Rubiaceae–Psychotrieae) with the description of some new species from Ecuador and Colombia. Ann. Missouri Bot. Gard. 84: 224–262. Taylor, C. M. 1999. 162(20). Rubiaceae–Coussareae. In: G. Harling and L. Andersson, eds., Flora of Ecuador 62: 245–320. Council for Nordic Publications in Botany, Copenhagen. Taylor, C. M. 2001a. Rubiaceae. In: W. D. Stevens, C. Ulloa Ulloa, A. Pool, and O. M. Montiel, eds., Flora de Nicaragua. Monogr. Syst. Bot. Missouri Bot. Gard. 85. St. Louis: Missouri Botanical Garden Press. Taylor, C. M. 2001b. Overview of the neotropical genus Notopleura (Rubiaceae: Psychotrieae), with the description of some new species. Ann. Missouri Bot. Gard. 88: 478–515. Taylor, C. M. 2002a. Rubiacearum Americanarum Magna Hama Pars X. New species and a new subspecies of Faramea (Coussareae) from Central and South America. Novon 12: 563–570.

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Taylor, C. M. 2002b. Rubiacearum Americanarum Magna Hama Pars XI. A new species of Alseis (Calycophylleae) from Central America and notes on the morphology of this neotropical genus. Novon 12: 571–574. Taylor, C. M. 2002c. Rubiacearum Americanarum Magna Hama Pars VI: New species of and morphological notes on Psychotria subg. Psychotria (Psychotrieae) from Mesoamerica and western South America. Novon 12: 120–132. Taylor, C. M. 2004. The neotropical genus Ronabea (Rubiaceae, Lasiantheae). Syst. Geogr. Pl. 74: 35–42. Taylor, C. M., and L. Andersson. 1999. 162(18). Rubiaceae–Psychotrieae (1). In: G. Harling and L. Andersson, eds., Flora of Ecuador 62: 134–241. Council for Nordic Publications in Botany, Copenhagen. Taylor, C. M., and J. L. Clark. 2001. Rubiacearum Americanarum Magna Hama V. Amphidasya in Mesoamerica and western South America. Novon 11: 489– 497. Taylor, C. M., and D. H. Lorence. 1993. On the status of Randia armata (Sw.) DC. (Rubiaceae: Gardenieae). Taxon 42: 865–867. Taylor, C. M., and D. H. Lorence. 2001. Rubiacearum Americanarum Magna Hama Pars II. New and newly circumscribed taxa of Guettarda (Guettardeae). Novon 11: 127–134. Verdcourt, B. 1976. Rubiaceae (Part 1). In: R. M. Polhill, ed., Flora of Tropical East Africa. 414 pp. Crown Agents for Overseas Governments and Administrations, London, United Kingdom. Verdcourt, B. 1983. Notes on Mascarene Rubiaceae. Kew Bull. 37: 521–571. Zappi, D. C., J. Semir, and N. I. Pierozzi. 1995. Genipa infundibuliformis sp. nov. and notes on Genipa americana (Rubiaceae). Kew Bull. 50: 761–771. Key to the Genera of Rubiaceae by Charlotte M. Taylor and Julian A. Steyermark 1. 1. 2(1). 2. 3(2).

3.

4(3). 4.

Plants epiphytic, usually succulent .......................................................... 2 Plants terrestrial, succulent or not ........................................................... 6 Leaves at each node strongly unequal, to apparently alternate with intrapetiolar stipules ................................................... 27. Didymochlamys Leaves at each node generally equal in size, with stipules interpetiolar ................................................................................................................ 3 Inflorescences pedunculate, each peduncle with an involucrate flower or head; plants usually lianas; juvenile growth often with adventitious roots; fruit baccate ................................................................ 77. Schradera Inflorescences with flowers sessile to pedicellate or pedunculate, borne singly and not involucrate; plants shrubby to clambering, without adventitious roots; fruit drupaceous or capsular ..................................... 4 Stipules united around stem in a truncate sheath 1–3 mm long; corollas 4–10 mm long; fruit drupaceous ......................................... 59. Notopleura Stipules interpetiolar or shortly united around the stem, the intrapetiolar portion ligulate to obovate, 10–30 mm long; corollas 15– 40 mm long; fruit capsular .................................................................... 5

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5(4). 5.

6(1). 6. 7(6). 7. 8(7). 8. 9(7). 9. 10(9). 10. 11(9). 11. 12(11). 12. 13(11). 13. 14(6). 14. 15(14). 15. 16(15).

16. 17(15). 17.

18(17). 18.

Flowers 2–several; corolla lobes imbricate in bud; seeds with an erose marginal wing but otherwise glabrous ............................. 22. Cosmibuena Flowers solitary; corolla lobes convolute in bud; seeds with an entire to erose marginal wing and with a tuft of long trichomes at one end ....................................................................................................... 43. Hillia Plants twining climbers, or lianas climbing with spines ......................... 7 Plants erect, sometimes clambering but not extensively twining, or creeping ......................................................................................................... 14 Stipules truncate to broadly rounded and fused to the petiole bases, setose ......................................................................................................... 8 Stipules interpetiolar, triangular to ligulate or obovate .......................... 9 Inflorescences axillary, glomerate or capitate, with flowers sessile .......... ..................................................................................................... 28. Diodia Inflorescences axillary and terminal, paniculate, with the flowers pedicellate ................................................................................ 32. Emmeorhiza Leaves with the tertiary venation lineolate ........................................... 10 Leaves with the tertiary venation irregularly areolate ......................... 11 Corolla salverform to funnelform, internally glabrous; inflorescences capitate to paniculate with cymose axes or flowers solitary ....... 15. Chomelia Corolla rotate to funnelform, villose in throat; inflorescences spiciform to paniculate, the axes usually spiciform ................................... 53. Malanea Stipules triangular, appressed to stem ................................................... 12 Stipules ligulate to oblanceolate or obovate, spreading from stem to reflexed .................................................................................................... 13 Fruit drupaceous, fleshy, white; ovules and seeds 1 per locule ... 14. Chiococca Fruit capsular, dry, brown; ovules and seeds numerous per locule ........... ................................................................................................. 54. Manettia Plants unarmed soft vines; fruit baccate, fleshy; seeds angled ..... 76. Sabicea Plants climbing by axillary spines, lianas; fruit capsular, dry; seeds winged ....................................................................................... 85. Uncaria Plants soft herbs, usually low and slender ............................................. 15 Plants trees, shrubs, or suffrutescent robust herbs, sometimes with soft stems from a substantial woody underground stock ......................... 30 Leaves apparently verticillate and exstipulate ...................................... 16 Leaves paired to verticillate, with stipules, remnant stipules, or glandular stipules or lines connecting the leaves .......................................... 17 Inflorescences axillary, the flowers solitary or few on each peduncle; calyx and corolla lobes 4; plants of wet montane and premontane vegetation .................................................................................................... 35. Galium Inflorescences terminal, the flowers several in branched cymes; calyx and corolla lobes 5; plants of savannas ................................. 49. Limnosipanea Plants creeping, regularly rooting at nodes including nodes with leaves; stipules triangular to bilobed; fruit fleshy or capsular ...................... 18 Plants tufted or erect, the stems sometimes weak to trailing but not regularly rooting at the nodes; stipules triangular to bilobed or setose; fruit dry, capsular, schizocarpous, or indehiscent ....................................... 21 Corolla lobes convolute in bud; fruit dry, capsular ...................... 79. Sipanea Corolla lobes valvate in bud; fruit fleshy, baccate or drupaceous .......... 19

504

R UBIACEAE

19(18). Fruit red; leaves cordate or cordulate at base ........................... 37. Geophila 19. Fruit blue or black; leaves acute to obtuse, rounded or cordulate at base .............................................................................................................. 20 20(19). Fruit baccate, blue, hollow; ovules and seeds numerous per locule ........................................................................................ 18. Coccocypselum 20. Fruit drupaceous, black to blue, solid; ovules and seeds 1 per locule, 2 per fruit ....................................................................................... 67. Psychotria 21(17). Stipules triangular, bifid, erose, or triangular and shortly setose; ovules and seeds several to numerous per locule .......................................... 22 21. Stipules setose or reduced to a glandular line; ovules and seeds 1 per locule ......................................................................................................... 25 22(21). Corolla lobes convolute in bud; seeds angled and capsules not beaked .............................................................................................................. 23 22. Corolla lobes valvate or induplicate-valvate in bud; seeds angled and capsule beaked, or seeds winged at one end and capsules not beaked .............................................................................................................. 24 23(22). Stamens included ................................................................ 49. Limnosipanea 23. Stamens exserted .......................................................................... 79. Sipanea 24(22). Flowers subsessile to shortly pedicellate; capsules not beaked ..... 55. Merumea 24. Flowers pedicellate or pedunculate; capsules beaked .......... 60. Oldenlandia 25(21). Inflorescences terminal, capitate, enclosed by 2 pairs of leaf-like involucral bracts; calyx lobes 6; fruit with 3 mericarps ................. 71. Richardia 25. Inflorescences terminal and/or axillary, capitate to cymose, not enclosed by leaf-like bracts, sometimes enclosed by smaller involucral bracts or 1 pair of leaves with sheathing bases; calyx lobes 2–4; fruit with 2 locules or mericarps ................................................................................. 26 26(25). Inflorescence pedunculate, cymose or capitate ........................... 64. Perama 26. Inflorescences sessile, glomerulate or capitate ...................................... 27 27(26). Fruit a circumscissile capsule; seeds with a cruciform attachment scar ............................................................................................ 56. Mitracarpus 27. Fruit indehiscent, schizocarpous, or a septicidal capsule; seeds with a round to linear attachment scar ......................................................... 28 28(27). Fruit indehiscent; plants with foul odor ...................................... 83. Tobagoa 28. Fruit schizocarpous or capsular; plant odor not particularly foul ......... 29 29(28). Fruit capsular ................................................................................ 8. Borreria 29. Fruit schizocarpous, the mericarps indehiscent ............................ 28. Diodia 30(14). Stipules laciniate, multifid, setose, fimbriate, or densely ciliate to ciliatesetose .................................................................................................... 31 30. Stipules entire to bilobed ......................................................................... 41 31(30). Stipules laciniate to multifid ................................................................... 32 31. Stipules setose to densely ciliate, setose-ciliate, or fimbriate ............... 35 32(31). Ovules and seeds 1 per locule, 2 per fruit; fruit drupaceous, fleshy ...... 33 32. Ovules and seeds numerous in each locule and the fruit; fruit dry, capsular or perhaps indehiscent .................................................................. 34 33(32). Inflorescences sessile to pedunculate, capitate, with well developed bracts; fruit blue to black ................................................................. 67. Psychotria 33. Inflorescences sessile to pedunculate, capitate to paniculate, bracts developed or not; fruit white, orange, red, or black ..................... 75. Rudgea

R U B I A C E A E 505

34(32). Calyx lobes 10–15 mm long .....................................................4. Amphidasya 34. Calyx lobes 1–3 mm long ........................................................ 80. Sipaneopsis 35(31). Ovary and fruit superior; fruit fleshy; stipules with tubular sheath ................................................................................................. 61. Pagamea 35. Ovary and fruit partially to fully inferior; fruit dry; stipules without tubular sheath ......................................................................................... 36 36(35). Plants branched shrubs and subshrubs, with stems not particularly stout and the internodes developed; inflorescences sessile to pedunculate; plants found widely in the flora area .................................................. 37 36. Plants little-branched or unbranched, pachycaulous shrubs, with the leaves congested near the stem apices due to short development of internodes; inflorescences pedunculate; plants of tepui slopes and summits ....................................................................................................... 39 37(36). Inflorescences pedunculate, cymose ........................................... 29. Duidania 37. Inflorescences sessile, glomerate or capitate .......................................... 38 38(37). Fruit capsular, with the segments dehiscent ............................... 8. Borreria 38. Fruit schizocarpous, with the segments indehiscent .................... 28. Diodia 39(36). Ovary partially inferior; calyx and corolla lobes 4; fruit capsular, dehiscent ........................................................................ 21. Coryphothamnus 39. Ovary fully inferior; calyx and corolla lobes 5 or 6; fruit apparently indehiscent .................................................................................................. 40 40(39). Ovary and fruit 1-locular; corolla tube entire .................... 5. Aphanocarpus 40. Ovary and fruit 2-locular; corolla tube fenestrate ............... 62. Pagameopsis 41(30). Stipules fused completely into a calyptrate cap, caducous, and densely pilose, hirsute, or sericeous ................................................................. 42 41. Stipules interpetiolar, united around the stem, divided deeply into 2 or 4 segments, or fused completely into a calyptrate cap but this glabrous .............................................................................................................. 45 42(41). Ovary and fruit partially inferior; petioles with 1–several glands at their abaxial bases; inflorescences paniculate; fruit capsular ........................................................................................... 65. Platycarpum 42. Ovary and fruit inferior; petioles without glands; inflorescences capitate, fasciculate, shortly cymose to extensively paniculate, or with flowers solitary; fruit fleshy or capsular ......................................................... 43 43(42). Flowers hermaphroditic, numerous in paniculate inflorescences; fruit capsular ....................................................................................... 6. Bathysa 43. Flowers unisexual on dioecious plants, solitary or to several in capitate to fasciculate inflorescences; fruit fleshy ................................................ 44 44(43). Pistillate flowers 3–several; plants glabrous to puberulous or strigose ................................................................................................... 3. Amaioua 44. Pistillate flowers solitary or 2–5; plants glabrous to hirsute or pilose ..................................................................................................... 30. Duroia 45(41). Stipules deeply split in the intrapetiolar portion, thus appearing to comprise 2 intrapetiolar segments or these sometimes also divided, producing 4 nearly or apparently free segments ..................................... 46 45. Stipules united across the interpetiolar portion, entire to bilobed, with the interpetiolar portion equal to or longer than the intrapetiolar portion ....................................................................................................... 50

506

R UBIACEAE

46(45). Petioles with 1–several glands at their abaxial bases; ovary and fruit partially inferior ................................................................ 42. Henriquezia 46. Petioles without glands; ovary and fruit fully inferior .......................... 47 47(46). A few to all of the flowers with 1 or more calyx lobes expanded into petaloid calycophylls; fruit capsular, 12–45 × 8–35 mm ........................... 48 47. None of the calyx lobes expanded or petaloid; fruit capsular or fleshy, 2– 10 × 2–10 mm ....................................................................................... 49 48(47). Trees reaching the canopy of rain forests; petaloid calyx lobes only 1 per flower and only present on a few flowers of each inflorescence; fruit cylindrical to oblong ............................................................... 11. Capirona 48. Shrubs or small trees of subcanopy or seasonal forests; all calyx lobes of all flowers petaloid; fruit suborbicular to obconic and somewhat flattened laterally ........................................................................ 38. Gleasonia 49(47). Fruit capsular; corolla ca. 4 mm long ......................................... 31. Elaeagia 49. Fruit fleshy; corolla 9–78 mm long ................................................. 45. Isertia 50(45). Inflorescence pseudoaxillary (i.e., regularly produced in only one axil at each node along the stem; “lateral”) ................................................... 51 50. Inflorescence terminal and/or axillary (i.e., regularly produced in both axils of each node along the stem) ...................................................... 52 51(50). Stipules with a glandular and/or succulent appendage or projection borne from near the middle of the sheath .................................... 59. Notopleura 51. Stipules unappendaged or with 1 or 2 terminal lobes arising from the top of the sheath, not glandular ................................................. 67. Psychotria 52(50). Inflorescences all axillary, borne from nodes below the stem apex ....... 53 52. Inflorescences terminal and sometimes also produced in the axils of the uppermost leaves, sometimes terminal on reduced lateral shoots.... 69 53(52). Ovules numerous in each locule, seeds numerous in each fruit; fruit capsular ..................................................................................................... 54 53. Ovules solitary in each locule, seeds 1–9 per fruit; fruits fleshy or dry and indehiscent ........................................................................................... 59 54(53). Stipules quickly caducous, present only at the stem apex ..................... 55 54. Stipules generally persistent with the leaves ........................................ 56 55(54). Capsules clavellate to obovoid; flowers protogynous; corolla 3–6 mm long; deciduous trees with flowers produced in the leafless stage ........ 2. Alseis 55. Capsules cylindrical to fusiform or oblong; flowers protandrous; corolla 12–55 mm long; evergreen trees or shrubs ............................. 69. Remijia 56(54). Flowers numerous in branched cymes; calyx lobes 0.5–1 mm long; lowland trees ............................................................................ 13. Chimarrhis 56. Flowers solitary or 2–4 in fascicles or short cymes; calyx lobes 6–32 mm long; upland or highland shrubs ......................................................... 57 57(56). Leaves broadly obtuse to rounded at apex .................... 12. Chalepophyllum 57. Leaves acute to acuminate at apex ......................................................... 58 58(57). Leaves 4.5–11 × 1.5–3.5 cm; corolla tube ca. 23 mm long .... 44. Holstianthus 58. Leaves 1.5–6.5 × 0.5–2.5 cm; corolla tube 65–130 mm long ...................... .................................................................................. 52. Maguireothamnus 59(53). Ovary and fruit laterally strongly flattened, the fruit didymous; low shrubs from a large woody underground base, in savannas ............. ............................................................................................... 25. Declieuxia

R U B I A C E A E 507

59. 60(59). 60. 61(60). 61.

62(60).

62.

63(62). 63. 64(62). 64. 65(64). 65. 66(65). 66. 67(64). 67. 68(67). 68. 69(52).

69.

70(69). 70. 71(69). 71. 72(71).

Ovary and fruit rounded, the fruit not didymous; plants of various habits and habitats ......................................................................................... 60 Ovary 1-locular; ovules 1 or 2 per ovary; seeds 1 per fruit .................... 61 Ovary 2–9-locular; ovules 1 per locule; seeds 2–9 per fruit ................... 62 Fruit subglobose to ellipsoid, white, yellow, or orange; stipules obtuse to rounded or sometimes calyptrate ......................................... 23. Coussarea Fruit subglobose to oblate and often laterally compressed, blue, black, or orange; stipules obtuse to acute, acuminate, aristate, or sometimes calyptrate ................................................................................ 33. Faramea Stipules adaxially sericeous to glabrous; leaves with tertiary venation lineolate, regularly areolate, or irregularly areolate; pyrene solitary, 2–9-locular ........................................................................................... 63 Stipules adaxially glabrous or with colleters inserted at base; leaves with tertiary venation irregularly areolate or not visible; pyrenes 2 per fruit, 1-locular ...................................................................................... 64 Calyx limb lobed; calyx and corolla lobes 4, the lobes valvate, valvateinduplicate, or shortly imbricate in bud; ovary 2-locular .....15. Chomelia Calyx limb truncate to undulate or reduced; corolla lobes 4–9, imbricate in bud; ovary 2–9-locular ...................................................... 40. Guettarda Ovules and seeds attached to septum ..................................................... 65 Ovules and seeds attached to base of locule ........................................... 67 Inflorescences capitate, the flowers separate or fused, ebracteate or with reduced bracts; corolla lobes valvate in bud ........................... 57. Morinda Inflorescence subcapitate to branched, the flowers free, subtended by one or more pairs of small bracts; corolla lobes convolute in bud ............ 66 Calyx and corolla lobes 5–8; corolla funnelform ............................. 19. Coffea Calyx and corolla lobes 4; corolla slenderly funnelform .................. 46. Ixora Inflorescences pedunculate, the peduncles 0.5–5 cm long ...... 67. Psychotria Inflorescences sessile or subsessile, the peduncles < 0.5 cm long ......... 68 Calyx and corolla lobes 4; plants of tepui summits ........... 17. Coccochondra Calyx and corolla lobes 5; plants of lowlands ............................. 72. Ronabea Plants with normally developed long-shoots and congested short-shoots that have the leaves closely grouped because the internodes are not developed, the unequal internode development usually visible along main stems if lateral short-shoots are not present; plants sometimes armed with spines; flowers terminal on long- and/or short-shoots ... 70 Plants with internodes generally all developed, the leaves somewhat grouped to well separated; plants unarmed or armed with spinescent lateral branches; flowers terminal on main stems and/or short lateral branches ............................................................................................... 71 Flowers unisexual on dioecious plants; corolla tube 2.5–30 mm long; young fruits uniformly green .................................................... 68. Randia Flowers bisexual; corolla tube 30–120 mm long; young fruit green with white spots and/or lines ........................................ 74. Rosenbergiodendron Leaves verticillate at most or all nodes .................................................. 72 Leaves opposite ........................................................................................ 74 Flowers solitary or paired; leaves 1–6 × 0.5–3 cm; fruit dry, capsular; plants of middle and upper tepui slopes .................... 58. Neblinathamnus

508

R UBIACEAE

72. 73(72). 73. 74(71). 74. 75(74). 75.

76(75). 76. 77(74). 77. 78(77). 78. 79(78). 79. 80(79). 80. 81(80). 81. 82(77). 82. 83(82). 83. 84(83). 84.

85(83).

85.

Flowers in cymes; leaves 5–23 × 2–8 cm; fruit fleshy, indehiscent; plants of lowlands ........................................................................................... 73 Fruit baccate, with numerous seeds embedded in the pulp; corolla lobes imbricate in bud ....................................................................... 41. Hamelia Fruit drupaceous, with 2–5 hard pyrenes each containing one seed; corolla lobes valvate in bud ..................................................... 63. Palicourea Some or all of the flowers with 1 to all of the calyx lobes expanded, 15 mm long or longer ....................................................................................... 75 Plants with calyx lobes small to medium size, to 10 mm long ............... 77 All the calyx lobes of each flower similarly enlarged, 15–20 mm long, ovate to broadly triangular, sessile, green ...................... 44. Holstianthus Only one calyx lobe of some flowers enlarged into a petaloid calycophyll, this 20–60 mm long, elliptic to obovate, stipitate, pale green to white or red to orange or yellow ........................................................................ 76 Inflorescence axes cymose, repeatedly branched; calycophylls white to pale green .......................................................................10. Calycophyllum Inflorescence axes spiciform, mostly unbranched; calycophylls red to orange or yellow ............................................................... 86. Warszewiczia Flowers staminate, without a developed ovary ...................................... 78 Flowers pistillate or bisexual, with a developed ovary .......................... 82 Inflorescences spiciform ..................................................... 82. Stachyarrhena Inflorescences capitate to cymose or fasciculate .................................... 79 Flowers pedicellate or pedunculate; corolla tube 15–50 mm long, the lobes 6–11 ............................................................................ 47. Kutchubaea Flowers sessile to pedicellate; corolla tube 5–14 mm long, the lobes generally 5 or 6 ............................................................................................. 80 Inflorescences cymose, pedunculate .............................................. 36. Genipa Inflorescences capitate to subcapitate, sessile to pedunculate .............. 81 Corolla tube usually > 10 mm long, rather thick-textured, not constricted below the corolla lobes ............................................................... 1. Alibertia Corolla tube usually < 10 mm long, if longer than this then membranaceous in texture or constricted below the corolla lobes .......... 20. Cordiera Ovules 1 or 2 per locule; seeds 1–5 per fruit ........................................... 83 Ovules 3 to numerous per locule; seeds several to numerous per fruit (sometimes the numerous seeds enclosed in hard pyrenes) .............. 94 Ovary 1-locular; ovules 1 or 2 per locule; seeds 1 per fruit .................... 84 Ovary 2–5-locular; ovules 1 per locule; seeds 2–5 per fruit ................... 85 Fruit subglobose to ellipsoid, white, yellow, or orange; stipules obtuse to rounded or sometimes calyptrate ......................................... 23. Coussarea Fruit subglobose to oblate and often laterally flattened, blue, black, or sometimes orange; stipules obtuse to acute, aristate, or sometimes calyptrate ................................................................................ 33. Faramea Ovary and fruit strongly laterally compressed, the fruit didymous; low plants from a large woody underground base, in savannas ............. ............................................................................................... 25. Declieuxia Ovary and fruit rounded to somewhat compressed laterally, the fruit sometimes didymous but not much compressed laterally; plants of various habits and habitats ................................................................ 86

R U B I A C E A E 509

86(85). Fruit dry, schizocarpous; plants sometimes with spinescent lateral stems .............................................................................................. 50. Machaonia 86. Fruit fleshy or perhaps dry, indehiscent; plants without regular spinescent lateral stems ...................................................................... 87 87(86). Ovary and fruit 3–5-locular ..................................................................... 88 87. Ovary and fruit 2-locular ......................................................................... 90 88(87). Stipules interpetiolar to united around the stem, truncate to triangular or spathaceous; corolla lobes convolute in bud ............. 70. Retiniphyllum 88. Stipules united around the stem or with the intrapetiolar portion reduced, truncate or shallowly emarginate to deeply bilobed; corolla lobes valvate in bud ............................................................................. 89 89(88). Corollas and inflorescence axes green to yellow, orange, red, blue, or purple; corolla swollen at the base, internally glabrous except for a ring of pubescence that closes off the basal swelling .......... 63. Palicourea 89. Corollas and inflorescences green, pale green, or white sometimes flushed with pink or purple; corolla straight at the base, internally glabrous to variously pubescent but if with a pubescent ring then this not situated shortly above the base ........................................... 67. Psychotria 90(87). Ovules and seeds attached to the septum .............................................. 91 90. Ovules and seeds attached to the base of the locule .............................. 92 91(90). Flowers free, subtended by 1 or 2 pairs of small bracts; inflorescences subcapitate to branched; corolla lobes convolute in bud .............. 46. Ixora 91. Flowers free or fused, ebracteate or with reduced bracts; inflorescences capitate; corolla lobes valvate in bud ...................................... 57. Morinda 92(90). Stipules with 1 or 2 groups of glands on each interpetiolar side, these borne on sheath side or margins or on the lobes, these often small and caducous and only evident on stem apex .................................. 75. Rudgea 92. Stipules without glands ........................................................................... 93 93(92). Corollas and inflorescence axes green to yellow, orange, red, blue, or purple; corolla swollen at the base, internally glabrous except for a ring of pubescence that closes off the basal swelling .......... 63. Palicourea 93. Corollas and inflorescences green, pale green, or white sometimes flushed with pink or purple; corolla straight at the base, internally glabrous to variously pubescent but if with a pubescent ring then this not situated shortly above the base ........................................... 67. Psychotria 94(82). Flowers solitary, sessile or subsessile to shortly pedunculate; fruit globose, 20–90 mm long ........................................................................... 95 94. Flowers several in branched to spiciform inflorescences; fruit variously shaped, 2–90 mm long ......................................................................... 99 95(94). Stipules obtuse to rounded or truncate; staminate inflorescences spiciform ......................................................................... 82. Stachyarrhena 95. Stipules rounded or obtuse to acute, acuminate, or aristate; staminate or hermaphrodite inflorescences capitate to fasciculate or shortly cymose .............................................................................................................. 96 96(95). Staminate flowers pedunculate or pedicellate; corolla tubes 15–50 mm long, the lobes 6–11 ............................................................ 47. Kutchubaea 96. Staminate or bisexual flowers sessile to pedicellate; corolla tube 5– 20 mm long, the lobes generally 5 or 6 ............................................... 97

510

R UBIACEAE

97(96). Corolla externally densely sericeous, yellow to orange or white marked with purple streaks; flowers bisexual ........................... 81. Sphinctanthus 97. Corolla externally puberulous to glabrous, white to cream or pale yellow; flowers unisexual on dioecious plants ................................................ 98 98(97). Corolla tube usually > 10 mm long, rather thick-textured, not constricted below the corolla lobes; fruits larger with mesocarp fleshy or woody or if small then glabrous with a woody mesocarp ........................ 1. Alibertia 98. Corolla tube usually < 10 mm long, if longer then membranaceous in texture or constricted below the corolla lobes; fruit small with mesocarp fleshy or woody ........................................................................ 20. Cordiera 99(94). Inflorescences spiciform, unbranched or with 1 or 2 pairs of secondary axes similar in form and length to the primary axis, the flowers borne directly from the primary axis .......................................................... 100 99. Inflorescences cymose to paniculate, branched to 2 or more orders, the secondary axes branched or if rather spiciform then much shorter than the primary axis; flowers borne on secondary and higher-order axes ............................................................................................................ 102 100(99). Deciduous trees, the flowers produced in the leafless stage; flowers protogynous; fruit capsular ........................................................ 2. Alseis 100. Evergreen shrubs or trees; flowers protandrous; fruit fleshy and indehiscent ............................................................................................ 101 101(100). Corolla tube 3–5 mm long, the lobes convolute in bud; fruit 12–20 mm long ................................................................................. 9. Botryarrhena 101. Corolla tube 4.5–8 mm long, the lobes imbricate in bud; fruit 3–5 mm long .............................................................................. 39. Gonzalagunia 102(99). Stipules caducous, present usually only at the stem apex .............. 103 102. Stipules generally persisting with the leaves .................................. 108 103(102). Flowers 2–6 in a corymbiform to umbelliform cyme; plants rather succulent ............................................................................. 22. Cosmibuena 103. Flowers 10–numerous in a corymbiform to pyramidal inflorescence; plants not particularly succulent .................................................. 104 104(103). Leaves broadly obtuse to rounded at apex; plants of tepui slopes .................................................................................. 51. Maguireocharis 104. Leaves obtuse to acute or acuminate at apex; plants of lowlands to middle elevations ........................................................................... 105 105(104). Corolla lobes imbricate in bud; fruit globose to ellipsoid .......... 78. Simira 105. Corolla lobes convolute or valvate in bud; fruit ellipsoid to oblong .... 106 106(105). Corolla tube ca. 3 mm long; fruit ca. 10 × 2 mm ............ 16. Cinchonopsis 106. Corolla tube 6–25 mm long; fruit 20–120 × 4–10 mm ..................... 107 107(106). Corolla lobes convolute in bud ....................................... 34. Ferdinandusa 107. Corolla lobes valvate in bud .............................................. 48. Ladenbergia 108(102). Flowers large, the corolla tube 20–295 mm long; fruit large, 20–90 mm long, baccate or capsular and flattened ........................................ 109 108. Flowers small to medium-sized, the corolla tube 2–22 mm long; fruit small to medium-sized, 3–20 mm long, of various types, if capsular and 20 mm long then not flattened .............................................. 112

R U B I A C E A E 511

109(108). Corolla narrowly to broadly funnelform or rather stoutly salverform, the tube 20–70 mm long; fruit baccate or capsular ..................... 110 109. Corolla slenderly salverform, the tube 40–295 mm long; fruit baccate ........................................................................................................ 111 110(109). Corolla externally glabrous or puberulous; fruit capsular and flattened ............................................................................................. 24. Coutarea 110. Corolla externally densely sericeous; fruit baccate, globose to ellipsoid ................................................................................................ 36. Genipa 111(109). Corolla white, the lobes imbricate in bud ........................... 66. Posoqueria 111. Corolla white to deep yellow, the lobes convolute in bud ..... 84. Tocoyena 112(108). Plants with flowers ............................................................................ 113 112. Plants with fruits ............................................................................... 119 113(112). Corolla lobes valvate in bud ................................................... 29. Duidania 113. Corolla lobes imbricate or convolute in bud ..................................... 114 114(113). Corolla lobes convolute in bud .......................................................... 115 114. Corolla lobes imbricate in bud .......................................................... 116 115(114). Corolla tube 2–4 mm long .......................................................... 7. Bertiera 115. Corolla tube 9–20 mm long .......................................... 26. Dendrosipanea 116(114). Corolla yellow to orange or red-orange, tubular to broadly funnelform, the tube 9–20 mm long ........................................................ 41. Hamelia 116. Corolla white sometimes flushed with pink or red, salverform to shortly funnelform or tubular, the tube 4.5–22 mm long, if tubular then the tube less than 9 mm long ............................................... 117 117(116). Corolla tubular to funnelform, the tube 5–7 mm long .............. 78. Simira 117. Corolla salverform, the tube 6.5–22 mm long .................................. 118 118(117). Corolla glabrous and smooth or sometimes thickened in throat ........................................................................................... 73. Rondeletia 118. Corolla densely pubescent and sometimes appendaged in throat ......................................................................................... 80. Sipaneopsis 119(112). Fruit 13–20 mm long .................................................................. 78. Simira 119. Fruit 2–10 mm long (unknown in several Sipaneopsis species) ...... 120 120(119). Fruit fleshy, baccate .......................................................................... 121 120. Fruit capsular or apparently indehiscent, dry ................................. 122 121(120). Inflorescences pyramidal to cylindrical, longer than wide, with the primary axis well developed .................................................. 7. Bertiera 121. Inflorescences broadly pyramidal to corymbiform, as wide as long or wider, with the primary axis well developed or not ........... 41. Hamelia 122(120). Capsules loculicidal ........................................................................... 123 122. Capsules septicidal or apparently indehiscent ................................ 124 123(122). Stipules 4–7 mm long; plants of middle and upper tepui slopes ............................................................................................. 29. Duidania 123. Stipules 2–3 mm long; plants of seasonal lowland vegetation ........................................................................................... 73. Rondeletia 124(122). Capsules 5–10 mm long and septicidal ....................... 26. Dendrosipanea 124. Capsules 2–4 mm long and apparently indehiscent (unknown in several species) ............................................................... 80. Sipaneopsis

512

R UBIACEAE

1. ALIBERTIA A. Rich in DC., Prodr. 4: 443. 1830. Genipella A. Rich. in DC., Prodr. 4: 443. 1830. Melanopsidium Poit. in DC., Prodr. 4: 443. 1830, hom. illeg., non Melanopsidium Colla 1824. Borojoa Cuatrec., Revista Acad. Colomb. Ci. Exact. 7: 474. 1949 [1950]. by Piero G. Delprete and Claes Persson Shrubs or trees, glabrous or pubescent, dioecious, unarmed, terrestrial. Leaves opposite or rarely verticillate, usually papyraceous to subcoriaceous, petiolate, venation brochidodromous, tertiary venation not lineolate; stipules interpetiolar and shortly connate or rarely free intrapetiolarly, persistent, narrowly triangular or ovate to oblong-ovate, acute, acuminate, or rarely obtuse or rounded at apex, in bud valvate. Inflorescences terminal, bracteate, sessile or rarely shortly pedunculate, the staminate fasciculate or rarely thyrse-like or with flowers solitary, the pistillate with flowers solitary or rarely 2. Flowers medium-sized or rarely small or large, sessile or shortly pedicellate, unisexual. Staminate flowers: hypanthium reduced; calyx limb cup-shaped or tubular, truncate, dentate, or rarely lobate, internally with or without colleters; corolla salverform, white throughout or tube greenish, externally minutely puberulent to sericeous or rarely glabrous, internally usually sericeous in upper portion, lobes 4–6(–8), contorted in bud; stamens 4–6(–8), inserted in the upper half of the corolla tube or rarely around its middle; anthers included or with tips slightly exserted, narrowly oblong to linear, dorsifixed, sessile; pollen 3-porate; pistillode present, slender. Pistillate flowers: hypanthium globose; calyx and corolla similar to the staminate except corolla often shorter and equipped with 1 or 2 additional lobes; staminodes present, similar to anthers but shorter; ovary 3–7-locular, ovules 9–70 per locule, on axial placentas; styles with 2–7 branches, abaxially with a revolute stigmatic area. Fruit baccate, globose, pulp fleshy, pericarp woody or fleshy and firm, usually yellowish, buff, or brown. Seeds sublenticular, irregularly lenticular, or lenticular; testa firm of elongated cells with secondary thickenings in the radial walls and outer tangental walls. Mexico, Central America, West Indies, Colombia, Venezuela, Trinidad-Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay; 21 species, 5 in Venezuela, 3 of these in the flora area. This treatment places Borojoa in Alibertia, without which Alibertia is paraphyletic (Persson 2000a). Several species that were previously included in Alibertia are here treated under Cordiera and discussed under that genus. Alibertia is similar to and frequently confused with Kutchubaea; their differences are discussed under Kutchubaea. Several species are said to have edible fruits. Key to the Species of Alibertia 1.

1.

Calyx dentate to lobate with lobes narrowly triangular to linear; corolla externally glabrous or sparsely puberulent; staminate flowers 5(6)merous; pistillate flowers 6-merous; fruits with fleshy mesocarp .............................................................................................. A. bertierifolia Calyx denticulate, dentate, subulate, or with widely triangular lobes; corolla externally puberulent or sericeous; staminate flowers 4(–6)merous; pistillate flowers 4–6(7)-merous; fruits with woody mesocarp ................................................................................................................ 2

Alibertia 513

2(1).

2.

Stipules 2–17 mm long; leaf blades 3.5–32(–35) × 1.2–10 cm, drying brown or olive green, the upper surface often shiny, the lower surface with inconspicuous or poorly developed secondary veins; corolla tube externally puberulent to densely puberulent, generally with retrorse hairs, drying gray, grayish beige, or sometimes beige-white; pistillate flowers (4)5- or 6(7)-merous; fruits 1–5.5 × 1–5.5 cm .................. A. edulis Stipules 6.5–9 mm long; leaf blades 2.8–15(–19) × 1.6–8 cm, drying dull brown, bronze, or greenish, the upper surface usually matte, the lower surface with conspicuous or well-developed secondary veins; the tertiary venation inconspicuous or not manifest; corolla tube externally densely sericeous, invariably with antrorse hairs, drying yellowish or light beige-white, sometimes with a goldish tinge; pistillate flowers 4(5)-merous; fruits 2–2.7 × 2–2.8 cm ......................................... A. latifolia

Alibertia bertierifolia K. Schum. in Mart., Fl. Bras. 6(6): 384. 1889. —Cordiera bertierifolia (K. Schum.) Kuntze, Revis. Gen. Pl. 1: 279. 1891. —Flor de rana. Alibertia stenantha Standl., Publ. Field Columbian Mus., Bot. Ser. 8: 170. 1930. Slender shrub or tree 3–8 m tall; leaves 6.5–26.5 × 1.8–12 cm, acute to obtuse at base, acute to acuminate at apex, with acumen 0.5–2.5 cm long; stipules 5–8 mm long, narrowly triangular to linear; corolla tube cylindrical, 4.5–8.5 mm long; fruits globose, to 40 × 40 mm. Gallery and lowland forests, 50– 400 m; Bolívar (Río Parguaza), Amazonas (road between El Burro and Puerto Ayacucho, Puerto Ayacucho, between San Carlos del Río Negro and mouth of Río Casiquiare, Trapichote). Apure; Panama, Colombia, Guyana, Ecuador, Peru, Brazil (Amazonas), Bolivia. ◆Fig. 415. Alibertia edulis (Rich.) A. Rich. in DC., Prodr. 4: 443. Sept. 1830. —Genipa edulis Rich., Actes Soc. Hist. Nat. Paris 1: 107. 1792. —Gardenia edulis (Rich.) Poir. in Lam., Encycl., Suppl. 2: 708. 1812. —Alibertia utilis A. Rich., Mém. Soc. Hist. Nat. Paris 5: t. 11. Dec. 1830, orth. var. —Garapatica edulis (Rich.) H. Karst., Fl. Columb. 1: 57. 1858 [1860]. —Cordiera edulis (Rich.) Kuntze, Revis. Gen. Pl. 1: 279. 1891. —Sabicea edulis (Rich.) Seem. ex B.D. Jackson, Index Kew. 772. 1895. Shrub or tree 1–10 m tall; leaves acute or obtuse to rounded at base, acute to acuminate at apex, with acumen 0.5–3.3 cm long; corolla tube cylindrical to narrowly obconical, 7–27 mm long. Mexico, Central

America, West Indies, Colombia, TrinidadTobago, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia; 2 varieties, both in the flora area. Key to the Varieties of A. edulis 1. Stipules narrowly ovate or narrowly triangular, (4–)5–17 × 2.5–7 mm, acute or acuminate to long-acuminate at apex; leaf blades acute to acuminate at apex; tertiary venation of upper leaf surface concolorous or discolorous ..... var. edulis 1. Stipules deltoid, 2–3 × 2–3 mm, acute at apex; leaf blades acute to rounded at apex; tertiary venation of upper leaf surface invariably discolorous ...................... ................................ var. obtusiuscula A. edulis var. edulis. —Carutilla, Guayaba de monte, Guayabilla morada, Guayabito, Marmelada, Puruí, Trompillo. Thieleodoxa lanceolata Cham., Linnaea 9: 253. 1834. —Borojoa lanceolata (Cham.) Cuatrec., Acta Agron. 3: 95. 1953. Cordiera acuminata Benth., J. Bot. (Hooker) 3: 221. 1841. —Alibertia acuminata (Benth.) Sandwith, Kew Bull. 1931: 470. 1931. Alibertia hexagyna H. Karst., Linnaea 30: 144. 1859. —Cordiera hexagyna (H. Karst.) Kuntze, Revis. Gen. Pl. 1: 279. 1891. Alibertia longistipulata L. Riley, Kew Bull. 1927: 122. 1927. Alibertia panamensis L. Riley, Kew Bull. 1927: 123. 1927. Alibertia tutumilla Rusby, Mem. New York Bot. Gard. 7: 375. 1927.

514

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Fig. 415. Alibertia bertierifolia

Alibertia tobagensis Sprague & R.O. Williams, Fl. Trinidad 2: 20. 1928. Alibertia trinitatis Sprague & R.O. Williams, Fl. Trinidad 2: 19. 1928. Thieleodoxa nitidula Bremek., Recueil Trav. Bot. Néerl. 33: 711. 1936. —Alibertia nitidula (Bremek.) L. Andersson, Scripta Bot. Belg. 1: 75. 1992. Shrub or tree 1–10 m tall. Savannas, deciduous to evergreen forests, 50–1300 m; widespread and common in Bolívar and Amazonas. Anzoátegui, Monagas, Zulia; Mexico, Central America, West Indies, South America. A. edulis var. obtusiuscula (Steyerm.) Delprete & C. Perss., comb. nov. —Alibertia acuminata var. obtusiuscula Steyerm., Mem. New York Bot. Gard. 12(3): 223. 1965. —Anoe, Anoeyo, Canilla de venado, Carutilla, Cazabe chiquito, Guayaba rebalsera. Alibertia davidsei Steyerm., Ann. Missouri Bot. Gard. 66: 903. 1979. Shrub 1–8 m tall. Semideciduous forests on granitic slopes, gallery forests, savannas, 50– 200 m; Bolívar (near Río Orinoco), Amazonas (western part near Río Orinoco). Guyana.

Alibertia edulis var. obtusiuscula is readily distinguishable by its short, deltoid stipules only 2–3 mm long, its leaf blades that are obtuse or rounded at apex, and its small fruits, ca. 1.5 cm in diameter. Alibertia latifolia (Benth.) K. Schum. in Mart., Fl. Bras. 6(6): 386. 1889. —Cordiera latifolia Benth., J. Bot. (Hooker) 3: 221. 1841. —Carutillo rebalsero, Cazabe chiquito, Matotoma de rebalse, Palo de boya, Paraguatán, Taparita. Alibertia latifolia var. parvifolia K. Schum. in Mart., Fl. Bras. 6(6): 386. 1889. Alibertia granulosa Rusby, Descr. S. Amer. Pl. 133. 1920. Alibertia latifolia var. pargueniana Steyerm., Mem. New York Bot. Gard. 12(3): 225. 1965. Shrub 3–5 m tall or tree 3–8(–10) m tall; leaves acute to rounded at base, acute to acuminate at apex, with acumen 0.5–1.7 cm long; corolla tube cylindrical to narrowly obconical, 6.5–16 mm long. Gallery forests, seasonally flooded forests, near sea level to 500 m; Delta Amacuro (Caño de Corisal, Caño Jota-Sabuca, Sacupana, Tucupita), Bolívar (Río Paragua), Amazonas (Río Asisa, Río Ventuari). Anzoátegui, Apure, Guárico; Colombia, Guyana, Suriname, Ecuador, Peru, Brazil, Bolivia.

Alseis 515

2. ALSEIS Schott in Spreng., Syst. Veg. 4: 404. 1827. by Charlotte M. Taylor and Julian A. Steyermark Trees or shrubs, terrestrial, unarmed, usually deciduous. Leaves opposite, subsessile to petiolate, venation not lineolate; stipules interpetiolar, triangular, caducous, imbricate or twisted in bud. Inflorescences axillary and sometimes also terminal, pedunculate, generally produced in the leafless stage, many-flowered, bracteate, the axes few and long-spiciform. Flowers small, homostylous, sweetly fragrant, subsessile to pedicellate, protogynous with the pistillate stage often markedly different from the staminate stage. Hypanthium turbinate to ellipsoid. Calyx limb lobed, lobes 5, without calycophylls; corolla white to pale yellow, tubular in pistillate stage, funnelform to campanulate in staminate stage, internally pubescent in upper part of tube, lobes 5, reduced, open to narrowly imbricate or valvate in bud. Stamens 5, inserted near base of corolla tube; anthers exserted, narrowly ellipsoid, dorsifixed; filaments villous. Ovary 2-locular; ovules numerous in each locule, on axile placentas. Fruits capsular, obovoid to clavellate, septicidal from apex, chartaceous to woody, the valves often later splitting from the apex. Seeds small, fusiform to rhombic, flattened, winged at each end. Mexico, Central America, Colombia, Venezuela, Guyana, French Guiana, Ecuador, Peru, Brazil, Bolivia; ca. 20 species, 3 in Venezuela, 1 of these in the flora area.

Fig. 416. Alseis labatioides

516

R UBIACEAE

Alseis is not well understood taxonomically. As noted by Taylor (2002b), its study is complicated by its usually leafless flowering stage and the protogynous flowers that change their morphology markedly between the pistillate and staminate stages. Alseis labatioides H. Karst. ex K. Schum. in Mart., Fl. Bras. 6(6): 190. 1889. —Carutillo, Totumillo, Lengua de vaca. Alseis leiantha S.F. Blake, Contr. U.S. Natl. Herb. 20: 531. 1924. Alseis trichocarpa Standl. & Steyerm., Fieldiana, Bot. 28: 566. 1953. Tree to 20 m tall; leaves 15–45 × 4–12.5 cm; stipules 9–15 mm long; inflorescences 9– 40 cm long; calyx limb 1–2 mm long; corolla

tube 2.5–4.5 mm long, lobes 0.5–1.5 mm long; capsules 10–23 × 3–4 mm. Deciduous, semievergreen, and gallery forests, near sea level to 700 m; northern and eastern Delta Amacuro (near El Palmar, Serranía de Imataca), eastern and northeastern Bolívar (Altiplanicie de Nuria, south of El Manteco, between Upata and Altagracia). Carabobo, Distrito Federal, Falcón, Guárico, Miranda, Yaracuy; Guyana. ◆Fig. 416.

3. AMAIOUA Aubl., Hist. Pl. Guiane 13. 1775. by Charlotte M. Taylor and Julian A. Steyermark Trees or shrubs, unarmed, terrestrial, dioecious; branching often distinctive, producing first an elongated internode followed by several short shoots. Leaves opposite or ternate, usually grouped at the ends of the stems, petiolate, venation not lineolate; stipules calyptrate (i.e., fused into a conical cap), densely sericeous, caducous, often splitting along one side and appearing spathaceous shortly before falling. Inflorescence terminal, few- to many-flowered, bracteate, sessile to pedunculate, the staminate fasciculate to cymose, the pistillate capitate or similar to the staminate. Flowers medium-sized, unisexual, sessile to pedicellate. Staminate flowers: calyx limb subtruncate to dentate, lobes 5 or 6, without calycophylls; corolla salverform, white to cream or greenish, externally sericeous, internally glabrous or pubescent in throat, lobes 5 or 6, convolute in bud; stamens 5 or 6, inserted at middle of corolla tube; anthers subsessile, narrowly oblong, dorsifixed, included; pistillode present, similar to style and stigma. Pistillate flowers: hypanthium turbinate; calyx and corolla similar to the staminate or sometimes smaller; staminodes present, similar to anthers; ovary 2-locular; ovules numerous in each locule, on axile placentas. Fruit baccate, subglobose, fleshy, becoming red-purple then black, reportedly sometimes becoming 1-locular. Seeds compressed, suborbicular, smooth, medium-sized to small, embedded in pulp. Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 7 species, 3 in Venezuela, 2 of these in the flora area. Amaioua is similar to Duroia; their separation is discussed under Duroia. The treatment here of Amaioua follows the classification of Steyermark (1974). However, his separation of these two species based on the arrangement of the pistillate inflorescence has been questioned by some field botanists, who consider this character continuously variable. Key to the Species of Amaioua 1.

Pistillate flowers sessile or borne on peduncles or pedicels 0–22 mm long, with corolla tube 6–9 mm long; fruits pedicellate or pedunculate, 7– 11 mm diameter; peduncles of staminate inflorescence 0–50 mm long;

Amaioua 517

1.

secondary leaf veins 5–12 pairs, with barbate trichomes often present in their abaxial axils .............................................................. A. corymbosa Pistillate flowers sessile, with corolla tube 4–11 mm long; fruits sessile, 12–15 mm diameter; peduncles of staminate inflorescences 0–11(–20) mm long; secondary leaf veins 8–16 pairs, with barbate trichomes usually not present in their abaxial axils ................................... A. guianensis

Amaioua corymbosa Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 3: 419, pl. 294. 1818 [1820]. —Cacho de venado, Canalete, Mergo. Amaioua peruviana Desf., Mém. Mus. Hist. Nat. 6: 16, pl. 4, fig. B. 1820. Tree or shrub to 10 m tall; leaves 7.5–21 × 3.5–12 cm; stipules 5–20 mm long; staminate inflorescences with 1–3 fasciculate peduncles terminating in congested cymes of flowers, pistillate inflorescences congested-cymose or similar to staminate; calyx limb 3.5–9.5 mm long, dentate; corolla tube 6–9 mm long, lobes 5–9 mm long; fruit 7–11 mm diameter. Semideciduous to evergreen lowland or lower montane forests, 50–700 m; common and widespread in Bolívar (vicinity of Panare village of Corozal, Reserva Forestal Imataca), frequent in Amazonas (Río Casanare be-

Fig. 417. Amaioua corymbosa

tween Río Siapa and Caño Momoni). Anzoátegui, Apure, Barinas, Lara, Monagas, Zulia; Mexico, Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil. ◆Fig. 417. Amaioua guianensis Aubl., Hist. Pl. Guiane 13, t. 375. 1775. Tree or shrub to 11 m tall; leaves 10–26 × 3.5–14 cm; stipules 5–22 mm long; staminate inflorescences cymose or with 1–3 fasciculate peduncles terminating in cymes, pistillate inflorescences capitate; calyx limb 5–12.5 mm long, dentate; corolla tube 4–11 mm long, lobes 3.5–16 mm long; fruit 12–15 mm diameter. Lowland and slope forests. Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Amazonian Brazil, Bolivia; 2 varieties, both in the flora area.

518

R UBIACEAE

Key to the Varieties of A. guianensis 1. Staminate calyx tube 5–6 mm long; staminate corolla with lobes 7–12 mm long, tube 9–10 mm long .... var. guianensis 1. Staminate calyx tube 6–9 mm long; staminate corolla with lobes 13–16 mm long, tube 10–11 mm long .......... var. macrantha A. guianensis var. guianensis. —Cacho de venado, Canilla de venado, Carutillo. Rain forests, semi-evergreen and deciduous forests, 50–500 m; frequent in Delta Amacuro (near El Palmar, Río Amacuro,

Serranía de Imataca), common and widespread in Bolívar (between El Dorado and Santa Elena de Uairén, near base of Sierra de Lema, north and east of Upata), occasional in Amazonas (Isla Ratón, Río Orinoco between Sananipo and mouth of the Vichada). Guyana, Suriname, French Guiana, Colombia, Ecuador, Peru, Amazonian Brazil, Bolivia. A. guianensis var. macrantha Steyerm., Mem. New York Bot. Gard. 12(3): 216. 1965. Lowland and slope forests, ca. 700 m; Amazonas (Sierra de la Neblina). Endemic.

4. AMPHIDASYA Standl., Field Mus. Nat. Hist., Bot. Ser. 11: 180. 1936. by Charlotte M. Taylor and Julian A. Steyermark Low shrubs or subshrubs, often unbranched, terrestrial, unarmed. Leaves opposite, petiolate, venation not lineolate; stipules interpetiolar, persistent, generally erect and flatly appressed in bud, entire or usually deeply multifid to laciniate with 5–17 narrow segments. Inflorescence pseudoaxillary and/or terminal, sessile to pedunculate, few- to several-flowered, cymose to subcapitate, bracteate, the bracts entire to multifid. Flowers sessile to shortly pedicellate, small to medium-sized, homostylous, protandrous, rather showy, at least sometimes nocturnal. Hypanthium oblong to turbinate. Calyx limb deeply lobed, lobes (4)5, well developed, narrow to spatulate or obovate, sometimes unequal but without calycophylls; corolla slenderly salverform with the tube usually well developed, white, internally glabrous, lobes 5, valvate in bud, on their margins and/or adaxial surface frequently verruculose or verrucose. Stamens 5, inserted in the upper part of the corolla tube; anthers linear, dorsifixed near the base, included, sometimes with the connective prolonged into a linear apical appendage. Ovary 2-locular; ovules numerous in each locule, on axile placentas. Fruit baccate, subglobose to ellipsoid, fleshy, apparently green, with the calyx lobes persisting and frequently enlarging markedly as the fruits develop. Seeds small, angled to subglobose, reticulate. Nicaragua, Costa Rica, Panama, Colombia, Venezuela, Ecuador, Peru, northern Brazil; ca. 15 species, 2 in Venezuela, 1 of these in the flora area. Amphidasya is recognizable by its low habit, multifid stipules, and congested axillary inflorescences. This genus was recently reviewed by Taylor and J. L. Clark (2001). The other Venezuelan species, A. venezuelensis (Standl.) Steyerm., is endemic to northern Venezuela and is anomalous in this genus in its branched shrub habit, entire stipules, and relatively short corollas. Amphidasya neblinae Steyerm., Mem. New York Bot. Gard. 23: 320, fig. 59. 1972. Suffrutescent plant to 1.5 m tall; leaves 20–24 × 7.5–10 cm; petioles 15–35 mm long; stipules 6.5–7.5 mm long, deeply 3–5-lobed; peduncle 0.5–1 cm long; flowers unknown; fruits 7–8 × 5 mm, with the persistent calyx

lobes 10–15 mm long. Lower montane forests, 200–300 m; Amazonas (Sierra de la Neblina). Brazil (side of Serra da Neblina along the Río Tucano). ◆Fig. 418. Amphidasya neblinae is similar to A. colombiana (Standl.) Steyerm. of low to middle elevations in western Amazonian Colombia, Ecuador, and Amazonian Peru; A.

Aphanocarpus 519

Fig. 418. Amphidasya neblinae

colombiana is allopatric and differs in its stipule lobes 5–13 mm long, versus 4–5.5 mm long in A. neblinae. Amphidasya colombiana may be expected in the southwestern part of the flora area.

5. APHANOCARPUS Steyerm., Mem. New York Bot. Gard. 12(3): 263, fig. 36a–b. 1965. by Charlotte M. Taylor and Julian A. Steyermark Low pachycaulous shrubs, unarmed, terrestrial. Leaves 3- or 4-verticillate but commonly strongly congested near the stem apices, sessile, venation not lineolate; stipules interpetiolar and fused to leaf bases, truncate, densely setose or densely ciliate, generally erect and valvate to flatly appressed in bud, persistent with the leaves, the stipule and leaves usually falling together as a unit. Inflorescence terminal and axillary, pedunculate, capitate, multiflowered, bracteate with the external bracts often involucral. Flowers small, sessile. Hypanthium turbinate. Calyx limb lobed, lobes 5, without calycophylls; corolla shortly funnelform, blue, purple, or lavender, internally glabrous in the tube but densely hirtellous adaxially on lobes, lobes 5, valvate in bud. Stamens 5, inserted near middle of corolla tube; anthers ellipsoid, dorsifixed, included. Ovary 1-locular; ovules 1 per locule, basal. Fruit capsular, obovoid, woody, apparently indehiscent. Seeds small, subtrigonous, puncticulate. Venezuela; 1 species. Aphanocarpus is similar to and apparently sympatric with Coryphothamnus and Pagameopsis. The distinctions between Aphanocarpus and Coryphothamnus are outlined under Coryphothamnus. Pagameopsis can be distinguished from Aphanocarpus by its 2-locular ovary and fenestrate corolla tube. Steyermark (1965, 263; 1974) classified Aphanocarpus in the tribe Psychotrieae, together with several other unusual genera of the Venezuelan tepuis. However, Aphanocarpus has several features that are unusual in both the Rubiaceae and the Psychotrieae (Taylor 1996), and its taxonomic placement in this tribe has

520

R UBIACEAE

been questioned (Piesschaert 2001, 38–39). Piesschaert noted that although Steyermark described the ovary as 1-locular, his figure (Steyermark 1974, 1016, fig. 157) shows two longitudinal sections of the ovary, one of which is depicted as 1-locular while the other is shown as 2-locular. It is not clear if Steyermark’s conclusion that the ovary is unilocular was based on flowers or on fruits; in Rubiaceae, frequently the fruits do not develop seeds in all the locules of the ovary, and thus appear to have fewer locules than are present in the ovary. Aphanocarpus steyermarkii (Standl.) Steyerm., Mem. New York Bot. Gard. 12(3): 264, fig. 36A–B. 1965. —Pagamea steyermarkii Standl., Fieldiana, Bot. 28: 589, fig. 131. 1953. Aphanocarpus steyermarkii f. elongatus Steyerm., Ann. Missouri Bot. Gard. 71: 331. 1984. Aphanocarpus steyermarkii f. glabrior Steyerm., Ann. Missouri Bot. Gard. 71: 330. 1984. Shrub to 1 m tall, silvery-sericeous; leaves 20–45 × 3–7 mm, strikingly pilose on mid-

Fig. 419. Aphanocarpus steyermarkii

vein and margins at least when young; stipule sheath 5–25 mm long, cilia or setae numerous, ca. 1 mm long; peduncles 1.5–10 cm long; heads ca. 1 cm wide; calyx limb 2– 3.5 mm long; corolla with tube 1.5–2 mm long, lobes 1.5–2 mm long; fruits 1.5–2 × 1 mm. Open rocky sandstone savannas, open scrub forest formations, sandstone slopes and tepui outcrops, 1000–2500 m; Bolívar (Aprada-tepui, Auyán-tepui, Camarcaibaraitepui, Carrao-tepui, Macizo del Chimantá, Murisipán-tepui, Ptari-tepui, Tereké-yuréntepui). Endemic. ◆Fig. 419.

Bathysa 521

6. BATHYSA C. Presl, Abh. Königl. Böhm. Ges. Wiss. ser. 5, 3: 514. 1845. by Charlotte M. Taylor Shrubs or trees, unarmed, terrestrial. Leaves opposite, petiolate, venation not lineolate; stipules interpetiolar, triangular, acute, and imbricate to valvate in bud to calyptrate (i.e., fused into a conical cap), persistent or caducous. Inflorescences terminal, subsessile to pedunculate, multiflowered, paniculate and often expansive, bracteate. Flowers sessile to pedicellate, rather small, homostylous, protandrous, fragrant. Hypanthium ellipsoid to turbinate. Calyx limb truncate to lobed, lobes 4 or 5, equal or in some species with one lobe of some flowers expanded into a cream to yellow petaloid calycophyll; corolla funnelform to campanulate, white to pale green, internally with a pubescent ring in tube or throat, lobes 4 or 5, imbricate in bud. Stamens 4 or 5, inserted in upper part of corolla tube; anthers ellipsoid, dorsifixed, exserted; filaments sometimes pubescent at base. Ovary 2-locular; ovules numerous, on axile placentas. Fruits capsular, ellipsoid to obovoid, septicidal from apex, chartaceous to woody. Seeds small, angled or somewhat flattened. Costa Rica, Panama, Colombia, Venezuela, Ecuador, Peru, Brazil, Bolivia; ca. 14 species, 2 in Venezuela, 1 of these in the flora area. Bathysa is similar to and often confused with Chimarrhis; their separation is outlined under Chimarrhis. The other species of Bathysa found in Venezuela, B. pittieri (Standl.) Steyerm., is endemic to middle elevations of the Coastal Cordillera. Delprete (1996) reviewed Bathysa in Venezuela. Bathysa bathysoides (Steyerm.) Delprete, Brittonia 48: 42. 1996. —Chimarrhis bathysoides Steyerm., Mem. New York Bot. Gard. 12(6): 181, fig. 29. 1965.

Tree to 25 m tall; leaves 14–22 × 8–15 cm; stipules calyptrate, 12–15 mm long, densely sericeous; inflorescences 15–25 × 10–25 cm; flowers sessile or subsessile; calyx limb ca.

Fig. 420. Bathysa bathysoides

522

R UBIACEAE

0.5 mm long, without calycophylls; corolla tube ca. 1.5 mm long, lobes ca. 1 mm long; capsules ca. 3 × 1 mm. Evergreen lowland to lower montane forests, 100–1000 m; Bolívar (Río Botanamo, trail to San Juan de Manapiare, 20 km east of Túriba), Amazonas (base of Cerro Sipapo, Río Coro Coro, tributary of

Río Padauiri on the Venezuelan-Brazilian boundary at Río Castanho, Río Cuao, Simarawochi). Endemic. ◆Fig. 420. Bathysa bathysoides is sometimes confused with Amaioua and Duroia, which have similar sericeous calyptrate stipules.

7. BERTIERA Aubl., Hist. Pl. Guiane 180, t. 69. 1775. by Charlotte M. Taylor and Julian A. Steyermark Shrubs or small trees, unarmed, terrestrial. Leaves opposite, petiolate to subsessile, venation not lineolate; stipules interpetiolar and usually at least shortly united around the stem, persistent, generally valvate to imbricate in bud, triangular or rarely spathaceous (i.e., completely fused and split along one side). Inflorescence terminal, pedunculate, multiflowered, thyrsiform-paniculate, bracteate, the secondary axes usually dichasial at basalmost node, then monchasial and secund at subsequent nodes. Flowers small, sessile to pedicellate, homostylous, protandrous. Hypanthium turbinate. Calyx limb partially lobed, lobes 4 or 5, without calycophylls; corolla salverform to narrowly funnelform, white to pale green, internally hirsute in tube, the lobes 4 or 5(6), convolute in bud, acute to acuminate. Stamens 5, inserted in the corolla throat; anthers narrowly oblong, included, dorsifixed, with the connective prolonged apically into an apiculate appendage; filaments very short. Ovary 2-locular; ovules several to numerous in each locule, on axile placentas. Fruit fleshy, baccate, subglobose, blue-black to black. Seeds small, angled, foveolate to granular or tuberculate. Mexico, Central America, West Indies, South America, tropical Africa, Madagascar, Mascarene Islands; ca. 50 species, 4 in Venezuela, 3 of these in the flora area. In fruit Bertiera is sometimes confused with Hamelia. Hamelia can be distinguished by its narrow small stipules and corymbiform rather than pyramidal inflorescences. Also found in Venezuela is Bertiera angustifolia Benth., known from Táchira and distinguished by its spathaceous stipules. Key to the Species of Bertiera 1. 1. 2(1).

2.

Corolla with tube 2–3 mm long; inflorescences with branched portion 6– 10 × 2.5–3 cm .......................................................................... B. parviflora Corolla with tube 2.5–5 mm long; inflorescence 4–19 × 3–9 cm .............. 2 Calyx limb truncate or lobed to 1/3 its length, the lobes broadly rounded; leaves glabrous except sericeous to pilosulous on midvein and secondary veins and densely hirtellous in abaxial vein axils ..... B. diversiramea Calyx limb lobed for 1/2 or more its length, the lobes acute to acuminate; leaves glabrescent or usually pilosulous to puberulous or hirtellous on both lamina and veins at least abaxially ............................. B. guianensis

Bertiera diversiramea Steyerm., Mem. New York Bot. Gard. 17(1): 319, fig. 36. 1967. —Carutillo blanco. Shrub or small tree to 4 m tall; leaves 10– 26 × 3.5–10 cm; petioles 5–15 mm long;

stipules 4–7 mm long; inflorescences with branched portion 6–15 × 4.5–9 cm; peduncles 3–8 cm long; calyx limbs ca. 0.5 mm long; corollas with tube ca. 4 mm long, the lobes 1.5– 2 mm long; fruits 3.5–4 mm diameter. Ever-

Bertiera 523

green forests, 100–400 m; eastern and northeastern Bolívar. Guyana, Suriname (Wilhelmina Mountains), adjacent Brazil (Amazonas, Roraima). Steyermark originally distinguished this species from Bertiera guianensis based primarily on its leaves with hirtellous domatia in the abaxial vein axils and its secondary inflorescence axes that are often branched dichotomously twice, versus no domatia and once branched in B. guianensis. However, hirtellous domatia also are present in some plants of B. guianensis where they may on occasion be densely pubescent and rather large, and occasional plants of B. guianensis may have the secondary inflorescence axes branched dichotomously more than once. Thus, these particular characters are not consistently informative for separating these species. Bertiera guianensis Aubl., Hist. Pl. Guiane 180, pl. 69. 1775. Shrub or small tree to 6 m tall; leaves 9– 20 × 2–2.5 cm; petioles 3–5 mm long; stipules 5–10 mm long; inflorescences with branched

portion 4–19 × 3–8 cm; peduncles 2.5–7 cm long; calyx limbs ca. 1 mm long; corollas with tubes 3–4 mm long, the lobes 2–3 mm long; fruits 3.5–5 mm diameter. Evergreen lowland forests, 100–300 m; widely distributed in Delta Amacuro, Bolívar, and Amazonas. Apure, Aragua, Barinas, Mérida, Monagas, Portuguesa, Táchira, Yaracuy, Zulia; Mexico, Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. Bertiera guianensis is notably variable in the size of the leaves and inflorescences and in the density and form of the pubescence. Steyermark (1967) distinguished two subspecies and three varieties within Bertiera guianensis. However, his subsp. pubiflora Steyerm., described from northeastern Peru, has been shown to belong to a separate species (Andersson and Ståhl 1999). Within Steyermark’s subsp. guianensis, var. leiophylla Steyerm. was described from Panama based effectively only on its lesser pubescence and has not been recognized by subsequent authors. The plants from the flora area fall into Steyermark’s var. guianensis.

Fig. 421. Bertiera parviflora

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Steyermark’s remaining variety, var. glabriflora Steyerm. of Amazonian Peru, has not been evaluated here but similarly to var. leiophylla was distinguished only by pubescence characters. Bertiera parviflora Spruce ex K. Schum. in Mart., Fl. Bras. 6(6): 325. 1889. Shrub or small tree to 4 m tall; leaves 8– 22 × 2–6.5 cm; petioles 1–4 mm long; stipules 4–6 mm long; inflorescences with branched portion 6–10 × 2.5–3 cm; peduncles 2–6.5 cm

long; pedicels ca. 0.5 mm long; calyx limbs 0.5–0.8 mm long; corollas with tube 2–3 mm long, the lobes ca. 1 mm long; fruits 3–4 mm diameter. Rain forests, 100–200 m; Amazonas (base of Sierra de la Neblina along Río Mawarinuma north to the Brazo Casiquiare). Amazonian Colombia, Peru, and Brazil (Amazonas: Rio Negro). ◆Fig. 421. Plants from near Sierra de la Neblina have smaller inflorescences, with both the primary and secondary axes shorter, than plants from western Amazonia.

8. BORRERIA G. Mey., Prim. Fl. Esseq. 79, pl. 1. 1818. by Charlotte M. Taylor and Julian A. Steyermark Annual or perennial herbs or low shrubs, terrestrial, unarmed, often weedy. Leaves opposite, sometimes developed in fascicles from axillary buds and apparently verticillate, subsessile to petiolate, venation not lineolate; stipules interpetiolar and fused to petiole bases, persistent, generally erect and valvate to appressed in bud, sheath truncate to triangular, setose. Inflorescences axillary and/ or terminal, sessile, glomerulate or capitate, few-flowered to multiflowered, bracteate, the bracts often setose or chaffy. Flowers usually small, homostylous, protandrous, sessile or subsessile. Hypanthium turbinate to subglobose. Calyx limb deeply 2- or 4-lobed, lobes equal or sometimes unequal in pairs, without calycophylls, the sinuses between the lobes sometimes setose or with small triangular projections; corolla salverform to funnelform or rotate, white, pink, or blue, internally glabrous or variously pubescent, lobes 4, valvate in bud. Stamens 4, inserted in corolla throat; anthers narrowly oblong, dorsifixed, usually exserted. Ovary 2locular; ovules solitary in each locule, axile. Fruit capsular, ellipsoid to subglobose or obovoid, septicidal from the apex or rarely the apex and the valves then loculicidal, papery to cartilaginous. Seeds small, hemispherical or ellipsoid. Pantropics; ca. 150 species, 19 species in Venezuela, 17 of these in the flora area. As with other members of its tribe, Borreria is distinguished by the dehiscence mode of its fruit and has been circumscribed somewhat differently by different authors. The systematics of this tribe are controversial and unresolved. Therefore, the generic limits used by Steyermark (1972; 1974) are followed here provisionally. Diodia is very similar to Borreria and commonly confused with it; Mitracarpus and Tobagoa are also quite similar. Diodia is separated from Borreria by its fruits that are indehiscent (e.g., D. kuntzei) or schizocarpous, splitting along the septum into two cocci that are indehiscent (e.g., D. sarmentosa) or sometimes tardily loculicidal along the abaxial face (e.g., D. ocymifolia). Most recent North American and European authors have combined Borreria with Spermacoce under this second name. The combined genus Spermacoce s.l. comprises about 250 pantropical and warm temperate species. Spermacoce s. str. is apparently Neotropical (though widely adventive in the Old World) and comprises perhaps a dozen species recognizable by their fruits with a characteristic dehiscence mode, with one half of the fruit (containing one seed) capsular and dehiscent but the other half (with the other seed) indehiscent. Spermacoce s. str. is found in northern Venezuela but not yet known from the flora area.

Borreria 525

Neotropical authors have taken a varied view of this group, ranging from accepting a broad Spermacoce s.l. (e.g., Burger and Taylor 1993; Liogier 2000), to accepting the two “traditional” genera (e.g., Steyermark 1972; 1974), to accepting the two traditional genera but with some species transferred from Diodia to Borreria (Taylor 2001a), to transferring some species from Diodia into Borreria and also recognizing several smaller genera separated from Borreria, Diodia, and Spermacoce sensu stricto (e.g., Bacigalupo and Cabral 1996; 1999). Due to both these varying generic limits and the need for mature fruits in all the classifications, generic identifications of species of Borreria and related genera are often difficult. Thus, a combined key is presented here to Borreria and the genera usually confused with it. In any condition, magnification is necessary for accurate determination of these plants. Previous authors have noted that several species differ in annual versus perennial habit, but the perennial species will apparently sometimes flower during their first year of growth, so this distinction is not a complete one. Key to the Species of Borreria, Diodia, Mitracarpus, and Tobagoa 1. 1. 2(1). 2. 3(2).

3. 4(3). 4. 5(4). 5. 6(5). 6. 7(5). 7. 8(7). 8.

9(4).

Calyx lobes 8–12 mm long; corolla tube 15–16 mm long ........................... ................................................................................ Borreria macrocephala Calyx lobes 0.5–3.5 mm long; corolla tubes 0.5–8 mm long ..................... 2 Plants creeping and usually rooting at nodes; fruits corky, solitary in leaf axils ...................................................................................... Diodia kuntzei Plants creeping to erect, not or sometimes rooting at nodes; fruits papery to cartilaginous, borne several in nodes and at stem apex .................. 3 Leaves dimorphic, those at flowering nodes reduced, half or less as large as vegetative leaves, the flowering apex of the stem thus spiciform; vegetative leaves 17–110 × 7–35 mm ................................. Diodia spicata Vegetative and reproductive leaves similar, the flower portion of the stem similar to the vegetative; leaves 3–100 × 0.5–40 mm .......................... 4 Calyx lobes 2 .............................................................................................. 5 Calyx lobes 3 or 4 ....................................................................................... 9 Plants usually with fascicles of small leaves produced in the axils of the vegetative leaves .................................................................................... 6 Plants without fascicles of small leaves in the axils of the vegetative leaves ...................................................................................................... 7 Terminal inflorescences and infructescences hemispherical; corolla tubes 1–1.5 mm long; fruits 3–4 mm long ............................. Borreria densiflora Terminal inflorescences and infructescences globose; corolla tubes 0.5– 1 mm long; fruit 1.5–2 mm long ................................. Borreria verticillata Corolla tubes 1.5–2 mm long; fruits 3.5–4 mm long ....... Diodia sarmentosa Corolla tubes 0.5–1 mm long; fruits ca. 1 mm long .................................. 8 Capsules cartilaginous; seeds ornamented with ca. 8 longitudinal rows of circular pits .................................................................... Borreria prostrata Capsules membranaceous; seeds ornamented with longitudinal rows of short low ridges and shallow depressions, often appearing irregularly striate .................................................................................. Borreria repens Leaves linear, narrowly oblong, very narrowly elliptic, or narrowly lanceolate, widest at base or near middle, with lateral margins nearly straight ................................................................................................. 10

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9. Leaves elliptic, with lateral margins curved .......................................... 18 10(9). Plants herbaceous and very slender with stems wiry; fruits ca. 1 × 1.2 mm .......................................................................... Mitracarpus microspermus 10. Plants herbaceous and slender to woody and robust; fruits 2–4.5 × 1– 2.5 mm .................................................................................................. 11 11(10). Flowers 1 or 2 per node, enclosed by the subtending leaves and stipule sheaths ............................................................................ Borreria pygmaea 11. Flowers 1–numerous per node, usually not enclosed by the subtending leaves and stipule sheaths .................................................................. 12 12(11). Leaves 4–7 × 0.5–1 mm ...................................................... Borreria intricata 12. Leaves 8–60 × 0.5–9 mm ......................................................................... 13 13(12). Largest leaves 0.5–2 mm wide ................................................................ 14 13. Largest leaves 2–9 mm wide ................................................................... 16 14(13). Leaves 8–13 mm long and calyx lobes ca. 1 mm long ...... Borreria jangouxii 14. Leaves 12–60 mm wide, calyx lobes 0.2–2 mm long .............................. 15 15(14). Leaves 0.5–1 mm wide; calyx lobes 1.5–2 mm long; corolla tube 2.8– 3.5 mm long ............................................................... Borreria confertifolia 15. Leaves 1–8 mm wide; calyx lobes 0.2–1.5 mm long; corolla tube 1.5– 2.2 mm long .................................................................. Diodia hyssopifolia 16(13). Corolla tube 1.5–2.2 mm long; fruits ellipsoid-oblong, 1–2 mm diameter; stipule setae glabrous .................................................. Diodia hyssopifolia 16. Corolla tube 2.5–8 mm long; fruits obovoid to subglobose; stipule setae glabrous or pubescent .......................................................................... 17 17(16). Plants perennial, woody at base, the roots usually not collected with the stems; calyx lobes equal or with 2 of them much longer; each segment of fruit on dorsal (abaxial) face with 3–5 well-developed ribs; corolla tube 4–8 mm long ............................................................ Diodia apiculata 17. Plants annual, soft or sometimes woody at base, the roots usually collected with the stems; calyx lobes equal to subequal; each segment of fruit on dorsal (abaxial) face smooth or with 1 well-developed rib and sometimes two shorter ribs; corolla tube 2.5–4.5 mm long ... Diodia teres 18(9). Calyx lobes 3–3.5 mm long; corolla tube ca. 6 mm long .......... Borreria alata 18. Calyx lobes 0.5–3 mm long; corolla tube 0.5–7 mm long ....................... 19 19(18). Corollas 1–1.5 mm long and not or hardly exceeding the calyx lobes; fruit 0.8–1 × 0.8–1 mm; low soft herbs ........................................................ 20 19. Corollas 1.5–6 mm long, equaling or exceeding the calyx lobes; fruits 1– 5 × 0.8–1 mm; low soft herbs to robust low shrubs ............................ 21 20(19). Corolla tube ca. 0.5 mm long; fruits ca. 1 mm long, septicidal .................. ........................................................................................ Borreria prostrata 20. Corolla tube 0.8–1 mm long; fruits ca. 0.8 mm long, circumscissile .......... ................................................................................... Mitracarpus diffusus 21(19). Plants with fruits (easier to identify) ...................................................... 22 21. Plants with flowers .................................................................................. 35 22(21). Fruits circumscissile ................................................................................ 23 22. Fruits septicidal or indehiscent ............................................................... 25 23(22). Fruits 2–3.5 × 2–3.5 mm ................................................... Mitracarpus hirtus 23. Fruits 1–1.5 × 1–1.5 mm .......................................................................... 24 24(23). Glomerules of flowers 8–15 mm diameter; plants to 0.5 m tall ................. .................................................................................... Mitracarpus frigidus

Borreria 527

24. 25(22). 25. 26(25). 26. 27(26). 27. 28(27). 28. 29(25). 29. 30(29). 30. 31(29). 31. 32(31). 32. 33(32). 33. 34(32). 34. 35(21). 35. 36(35). 36. 37(36). 37. 38(37). 38. 39(38). 39. 40(37). 40. 41(40).

Glomerules of flowers 3–5 mm diameter; plants to 0.1 m tall ................... .................................................................................. Mitracarpus parvulus Fruits indeshicent or with the segments separating but individually indehiscent .............................................................................................. 26 Fruits with the segments dehiscent ........................................................ 29 Leaves narrowly elliptic to narrowly oblanceolate, 1–7 mm wide, with secondary veins usually plane adaxially ....................... Diodia multiflora Leaves elliptic, 5–23 mm wide, with secondary veins usually sulcate adaxially ............................................................................................... 27 Calyx lobes ca. 0.5 mm long; plant with foul odor ........... Tobagoa maleolens Calyx lobes 1.2–5 mm long; plant without foul odor .............................. 28 Calyx lobes 1.5–5 mm long, at least some of them more than 3 mm long ............................................................................................... Diodia radula Calyx lobes 1.2–2.5 mm long ............................................ Diodia sarmentosa Fruits narrowly oblong to oblanceolate .................................................. 30 Fruits subglobose to obovoid ................................................................... 31 Plants of rocky riversides at 1500–1700 m ................ Borreria cataractarum Plants widespread in various habitats, sea level to 1400 m ...................... .......................................................................................... Borreria capitata Fruits 4.5–5 mm long .................................................... Borreria bolivarensis Fruits 2–3.5 mm long ............................................................................... 32 Fruits 2.5–3.5 mm long ............................................................................ 33 Fruits 1–2 mm long .................................................................................. 34 Calyx lobes 0.8–3 mm long; plants often drying yellow-green .................. .......................................................................................... Borreria latifolia Calyx lobes 0.5–1.2 mm long; plants not drying yellow-green .................. ............................................................................................ Borreria remota Plants erect; fruits 1–1.5 mm long ....................................... Borreria hispida Plants creeping, rooting at nodes; fruits ca. 2 mm long .. Borreria wurdackii Corolla with tube 5–7 mm long, lobes 2.5–4 mm long ............ Diodia radula Corolla with tube 1–5 mm long, lobes 0.5–2.5 mm long ........................ 36 Corolla tubes 1–2 mm long ...................................................................... 37 Corolla tubes 2–5 mm long ...................................................................... 42 Robust perennials, erect or often clambering or sprawling; leaves with secondary veins plane or usually sulcate adaxially ........................... 38 Slender annuals, usually erect; leaves with secondary veins generally plane adaxially .................................................................................... 40 Plants erect; leaves usually smooth adaxially; calyx lobes 0.5–1.2 mm long ..................................................................................... Borreria remota Plants weak to clambering; leaves usually scabrous adaxially; calyx lobes 0.5–2.5 mm long ................................................................................... 39 Calyx lobes 1.2–2.5 mm long; plant odor not particularly foul .................. ....................................................................................... Diodia sarmentosa Calyx lobes ca. 0.5 mm long; plants with foul odor ......... Tobagoa maleolens Inflorescences subtended by 2 decussate pairs of leaves; plants hirsute to glabrous, often drying yellow-green ................................ Borreria hispida Inflorescences subtended by 1 pair of leaves; plants hirtellous or strigose to glabrescent, not drying yellow-green.............................................. 41 Leaves 5–20 mm wide; corolla equaling or somewhat longer than the

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41. 42(36). 42. 43(42). 43. 44(43). 44. 45(44). 45.

calyx lobes, with tube 1.5–2 mm long .......................... Mitracarpus hirtus Leaves 1–5 mm wide; corolla well exceeding the calyx lobes, with tube 1– 1.5 mm long .............................................................. Mitracarpus parvulus Plants creeping, rooting at nodes ..................................... Borreria wurdackii Plants erect to weak, not rooting at nodes.............................................. 43 Flowers usually pale blue; plants often drying yellow-green .................... .......................................................................................... Borreria latifolia Flowers white; plants not drying yellow-green ...................................... 44 Plants of rocky riversides, 1700–1800 m ................... Borreria cataractarum Plants of various habitats, sea level to 1400 m ...................................... 45 Flowers several in leaf axils ............................................... Diodia multiflora Flowers numerous in dense hemispherical to subglobose heads ............... ................ Borreria bolivarensis, Borreria capitata, Mitracarpus frigidus

Borreria alata (Aubl.) DC., Prodr. 4: 544. 1830. —Spermacoce alata Aubl., Hist. Pl. Guiane 60, pl. 22, fig. 7. 1775. Herbs to 0.5 m tall, glabrous; leaves 40–70 × 10–30 mm, acute at base; stipule sheath 2– 3 mm long, setae 6–8, 2.5–8 mm long; inflorescences terminal and sometimes in upper leaf axils; calyx lobes 4, 3–3.5 mm long; corolla white to pale blue, tube ca. 6 mm long, lobes ca. 3.5 mm long; fruit subglobose, 1–2 × 1–2 mm. Riparian forests, 50–200 m; Amazonas (at base of Piedra Arauicaua, Río Casiquiare between mouth of Río Siapa and Caño Momoni, Río Yatúa). Guyana, Suriname, French Guiana, Amazonian Brazil. ◆Fig. 424. This species is similar to Borreria latifolia, and the separation of these can be subtle if the plants do not have mature flowers or fruits. The specific name refers to the shortly to markedly winged stem angles, but these vary in their degree of development and intergrade to some extent with those of B. latifolia. Plants of B. alata are frequently weak-stemmed and found in muddy microsites. These plants frequently dry with a characteristic yellow-green color, similarly to plants of B. latifolia. Borreria bolivarensis Steyerm., Mem. New York Bot. Gard. 23: 819. 1972. Subshrub to 0.5 m tall, hirtellous to glabrescent; leaves 50—70 × 9–15 mm, acute at base; stipule sheath ca. 5 mm long, setae 8–9, 2–7 mm long; inflorescences axillary and sometimes terminal; calyx lobes 4, 0.6–1 mm long; corolla white, tube ca. 4 mm long, lobes ca. 2.5 mm long; fruit obovoid, 4.5–5 mm long. Open rocky slopes, forested hills, 100–1100 m; eastern and northeastern Bolívar (Altiplanicie

de Nuria, Cerro Venamo, Serranía de Imataca). Possibly in Sucre. ◆Fig. 429. Borreria bolivarensis is similar to and perhaps not distinct from B. capitata. No material of this species has been seen by C.M. Taylor for the preparation of this flora. Borreria capitata (Ruiz & Pav.) DC., Prodr. 4: 545. 1830. —Spermacoce capitata Ruiz & Pav., Fl. Peruv. 1: 61, pl. 91, fig. b. 1978. Perennial herb or low shrub to 1.2 m tall, hirtellous or strigillose to glabrescent; leaves 1–100 × 1–10 mm, acute at base; stipule sheath 1–4 mm long, setae 4–9, 1–10 mm long; inflorescences terminal and sometimes axillary; calyx lobes 4, 1–2.5 mm long; corolla white, tube 2–4 mm long, lobes 1–3 mm long; fruit oblanceolate, 2–3 × 0.8–1 mm. Sandy and rocky savannas, along streams, granite outcrops, roadsides, rocky open places, sea level to 600(–1400) m; throughout Bolívar and Amazonas. Mexico, Central America, Antilles, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina; 3 varieties, 2 in Venezuela, both in the flora area. Steyermark (1972) recognized three varieties and three forms of this species, and reported two varieties from the flora area. He considered var. suaveolens (G. Mey.) Steyerm. to be a rare plant confined to Guyana and Suriname, although C.D. Adams (Burger and Taylor 1993), has applied this name to Central American plants [as Spermacoce suaveolens (G. Mey.) Kuntze]. This species or group of species and the application of the names Borreria capitata and B. suaveolens G. Mey. have not been carefully studied across the range of these plants. Borreria capitata as circumscribed by

Borreria 529

Steyermark comprises a notable range of morphological variation, in particular between the most vigorous plants of var. capitata and the most slender plants of var. tenella. Steyermark considered this variation to be continuous and not completely separable into smaller taxa, although the two varieties found in the flora area seem well marked. The terminal inflorescences of this species are characteristically subtended by 2 decussate pairs of leaves, which are similar to the vegetative leaves or sometimes lanceolate rather than elliptic. Key to the Varieties of B. capitata 1. Leaves mainly linear-lanceolate or linearoblanceolate, (2–)5–10 mm wide, with usually 3 or 4 pairs of secondary veins evident, these usually prominent on the lower surface; stipule setae mainly 4– 10 mm long .................... var. capitata 1. Leaves usually narrowly linear, mainly 1– 3(–5) mm wide, with the secondary veins not visible or 1 or 2 pairs of them evident, these plane on the lower surface; stipule setae mainly 2–7 mm long .......... .......................................... var. tenella B. capitata var. capitata Spermacoce ferruginea A. St.-Hil., Pl. Usuel. Bras. pl. 13. 1824. —Borreria ferruginea (A. St.-Hil.) DC., Prodr. 4: 547. 1830. —Borreria capitata [var. capitata] f. ferruginea (A. St.-Hil.) Steyerm., Mem. New York Bot. Gard. 23: 822. 1972. Borreria kappleriana Miq., Linnaea 18: 746. 1844. Borreria tenella var. platyphylla K. Schum. in Mart., Fl. Bras. 6(6): 55. 1888. Borreria capitata [var. capitata] f. glabra Steyerm., Mem. New York Bot. Gard. 23: 823. 1972. Rocky savannas, open ground, roadsides, rocky open places, along streams, 50–1400 m; Bolívar (Canaima, Cerro Bolívar, near Santa Elena de Uairén, Sierra de Lema, base of Uaipán-tepui), Amazonas (upper Río Ventuari, Río Sipapo, Yavita). Anzoátegui, Apure, Barinas, Carabobo, Distrito Federal, Guárico, Lara, Mérida, Miranda, Portuguesa, Trujillo; Guyana, Suriname, French Guiana, Peru, Brazil, Paraguay, Argentina. B. capitata var. tenella (Kunth) Steyerm., Mem. New York Bot. Gard. 23: 823.

1972. —Spermacoce tenella Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 3: 345. 1818 [1819]. —Borreria tenella (Kunth) Cham. & Schltdl., Linnaea 3: 317. 1828. Spermacoce orinocensis Willd. ex Roem. & Schult., Syst. Veg. 3: 531. 1818. Spermacoce aturensis Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 3: 345. 1818 [1819]. Sandy and rocky savannas, along streams, granite outcrops, 50–600(–1300) m; common throughout Bolívar and Amazonas. Apure, Aragua, Carabobo, Cojedes, Distrito Federal, Lara, Mérida, Monagas, Zulia; Colombia, Guyana, Suriname, French Guiana, Amazonian Peru, Brazil. ◆Fig. 427. Borreria cataractarum Steyerm., Mem. New York Bot. Gard. 23: 813. 1972. Subshrub to 0.3 m tall, hirtellous to glabrescent; leaves 8–33 × 2–8 mm, acute at base; stipule sheath 1.5–2 mm long, setae 4– 5, 1.5–2 mm long; inflorescences terminal and sometimes axillary; calyx lobes 4, 1–2 mm long; corolla white, tube ca. 2.8 mm long, lobes ca. 1.2 mm long; fruit 1.5–2 × 1 mm. Rocky ledges around waterfalls and cascades, 1700–1800 m; Bolívar (Macizo del Chimantá [Toronó-tepui]). Guyana (southern Pakaraima Mountains). ◆Fig. 426. Borreria cataractarum is similar to and perhaps not distinct from B. capitata. No material of this species was seen by C.M. Taylor for the preparation of this flora. Borreria confertifolia Steyerm., Mem. New York Bot. Gard. 23: 828. 1972. Much-branched subshrub to 0.2 m tall, hirtellous to glabrescent; leaves 12–25 × 0.5– 1 mm, acute at base; stipule sheath 1.5–2 mm long, setae 3–4, 1.5–3 mm long; inflorescences terminal and sometimes axillary; calyx lobes 4, 1.5–2 mm long; corolla white, tube 2.8–3.5 mm long, lobes 1–2 mm long; fruits unknown. Rocky stream banks, on sandy or granitic substrate, sandy savannas, 100–500 m; Bolívar (Macizo del Chimantá near Apacará-tepui), Amazonas (Sabanita Morocoto along the Río Orinoco near the confluence of the Río Atabapo). Guyana. Borreria confertifolia has densely clumped branches with relatively long linear leaves, including those that subtend the inflorescences. The name Borreria tenella var. [delta] crispata K. Schum. [in Mart., Fl. Bras.

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6(6): 55. 1889, Gardner 4175, photo MO!] was not cited by Steyermark but likely also applies to this species. As discussed under B. pygmaea below, Steyermark’s name B. pygmaea var. robusta may also apply to this species. Borreria densiflora DC., Prodr. 4: 542. 1830. —Spermacoce densiflora (DC.) Alain, Phytologia 54: 113. 1983. —Choori yo’ (Panare). Spermacoce spinosa Sw., Obs. Bot. 45. 1791, hom. illeg., not Spermacoce spinosa L. 1762, nom. illeg., not Spermacoce spinosa Jacq. 1763, hom. illeg. Apparently annual herb, scabridulous to glabrescent; leaves 40–100 × 3–15 mm, acute at base; stipule sheath 3–5 mm long, setae 6– 8, 2–7 mm long; inflorescences terminal and axillary; calyx lobes 2, 1–2.5 mm long; corolla white, tube 1–1.5 mm long, lobes ca. 0.5 mm long; fruit oblanceolate, 3–4 × 1 mm. Widespread but infrequent in weedy areas, roadsides, 50–400 m; Bolívar (Ciudad Bolívar, Río Maniapure). Anzoátegui, Aragua, Carabobo, Distrito Federal, Guárico, Mérida, Miranda, Portuguesa, Zulia; Mexico, Central America, West Indies, Trinidad, Colombia, Guyana, and likely in Suriname, French Guiana, Ecuador, Peru, Bolivia, and Brazil. Borreria densiflora is used medicinally for sores and wounds. It has relatively large inflorescences and characteristic fascicles of leaves growing from the axillary buds of the vegetative leaves, so that the leaves appear densely verticillate. Borreria hispida Spruce ex K. Schum. in Mart., Fl. Bras. 6(6): 62. 1888. Borreria hispida var. glabrescens K. Schum. in Mart., Fl. Bras. 6(6): 62. 1888. Annual herb to 0.3 m tall, hirsute to glabrous; leaves 10–40 × 1–10 mm, acute at base; stipule sheath 1–2 mm long, setae 5–6, 1.5–2 mm long; inflorescences terminal and axillary; calyx lobes 5, 1.8–3 mm long; corolla white, tube ca. 2 mm long, lobes ca. 0.5 mm long; fruit ellipsoid, 1–1.5 × 1–1.5 mm. Sandy savannas, rocky sandstone slopes, and granite outcrops, 100–1100 m; Bolívar (Canaima, El Paují, Gran Sabana). Amazonas (base of Cerro Yaví, near Puerto Ayacucho, near Río Ventuari). Anzoátegui, Sucre, Yaracuy; Guyana, Suriname, French Guiana, Brazil. ◆Fig. 431.

Borreria hispida characteristically has inflorescences subtended by 2 decussate pairs of lanceolate to ovate leaves that are not separated by an internode and thus surround the flowers as in Richardia. Most of the dried specimens seen of this species have a distinctive yellow-green color similar to that in Borreria latifolia and to B. poaya (A. St.-Hil.) DC. and Diodia schumannii Standl. ex Bacigalupo of Brazil, Bolivia, and Paraguay. Steyermark (1972) recognized two varieties of Borreria hispida distinguished by the presence, but this is variable and the varieties are not recognized here. Borreria intricata Steyerm., Mem. New York Bot. Gard. 23: 828, fig. 92. 1972. Shrub to 1 m tall, glabrescent; leaves 4–6 × 0.5–1 mm, acute at base; stipule sheath 1– 1.5 mm long, setae 3, 0.5–1 mm long; inflorescences terminal and sometimes axillary; calyx lobes 4, 1.7–2 mm long; corolla white, tube ca. 2.5 mm long, lobes 1.5–2 mm long; fruit ellipsoid, 3.5–4 × 1–1.5 mm. Savannas, 100–200 m; Amazonas (Río Atabapo, Río Sipapo). Endemic. Borreria jangouxii Steyerm. in Lasser & Steyerm., Fl. Venez. 9: 1965. 1974. Subshrub to 0.1 m tall, hirtellous to glabrescent; leaves 8–13 × 1–2 mm long, acute at base; stipule sheath 0.8–1 mm long, setae 3–4, 0.8–1.5 mm long; inflorescences terminal; calyx lobes 4, ca. 1 mm long; corolla white, tube ca. 2.2 mm long, lobes ca. 2 mm long; fruit ellipsoid, ca. 3 × 2 mm. Savannas, 100–200 m; Amazonas (savanna 8–10 km east of Caño Parucito and south of Cerro Corobita). Endemic. ◆Fig. 423. Borreria latifolia (Aubl.) K. Schum. in Mart., Fl. Bras. 6(6): 61, pl. 80. 1888. —Spermacoce latifolia Aubl., Hist. Pl. Guiane 55, pl. 19, fig. 1. 1775. —Chiyaya. Spermacoce caerulescens Aubl., Hist. Pl. Guiane 55, pl. 19, fig. 2. 1775. Borreria fockeana Miq., Linnaea 18: 299. 1844. —Borreria latifolia var. fockeana (Miq.) Bremek., Recueil Trav. Bot. Néerl. 31: 307. 1934. —Borreria latifolia [var. latifolia] f. fockeana (Miq.) Steyerm., Mem. New York Bot. Gard. 23: 809. 1972.

Borreria 531

Borreria latifolia var. minor K. Schum. in Mart., Fl. Bras. 6(6): 61. 1888. —Borreria latifolia [var. latifolia] f. minor (K. Schum.) Steyerm., Mem. New York Bot. Gard. 23: 810. 1972. Perennial herb to 1 m tall, hirtellous to glabrescent; leaves 10–70 × 5–40 mm, acute at base; stipule sheath 1–3 mm long, setae 5– 7, 1.5–6 mm long; inflorescences axillary and sometimes terminal; calyx lobes 4, 0.8–3 mm long; corolla white to pale blue, tube 2.5–5 mm long, lobes 1–1.5 mm long; fruit subglobose, 2.5–4 × 2.5–4 mm. Stream banks, riparian forests, savannas bordering Mauritia palm swamps, open and waste ground, 50–200 m; widespread in Delta Amacuro, Bolívar, and Amazonas. Apure, Barinas, Guárico; Colombia, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, and adventive in Africa, Asia, and the Pacific Islands. ◆Fig. 430. Borreria latifolia characteristically has inflorescences produced at many nodes along the stem and relatively large subglobose capsules that are usually pubscent. The dry plants frequently have distinctive yellowgreen color, similar to that of B. hispida, B. alata, and B. wurdackii. Borreria latifolia is widespread and variable. With further careful study of this group, B. latifolia may be found to comprise the plants included in B. wurdackii. Borreria alata is also similar. Borreria macrocephala Standl. & Steyerm., Fieldiana, Bot. 28: 566, fig. 120. 1953. Subshrub to 2 m tall, glabrous; leaves 35– 70 × 2–10 mm, acute at base; stipule sheath 1–3 mm long, setae 1–5, 0.5–2.5 mm long; inflorescences terminal; calyx lobes 4, 8–12 mm long; corolla white, tube 15–16 mm long, lobes 4–4.5 mm long; fruit 4.5–5 × 2 mm. White-sand savannas, along streams, on granitic outcrops and white sands, 50–200 m; Amazonas (Caño Cumare, Río Atabapo, Río Atacavi, Río Caname, Río Pasimoni). Colombia. ◆Fig. 422. Borreria macrocephala is a distinctive white-sand species that has a cane-like habit quite distinct from the other species of Borreria and related genera. Borreria prostrata (Aubl.) Miq., Stirp. Surinam. Select. 177. 1850 [1851]. —Spermacoce prostrata Aubl., Hist. Pl.

Guiane 58, t. 20, fig. 3. 1775. Herb to 0.4 m tall, often weak-stemmed, hirtellous to glabrescent; leaves 6–35 × 1–10 mm, acute at base; stipule sheath 1–1.5 mm long, setae 3–7, 1–3 mm long; inflorescences terminal and axillary; calyx lobes (2, 3)4, equal to strongly unequal, the longer ones 1– 1.2 mm long; corolla white, 1–1.2 mm long, lobed for ca. 1/2 its length; fruit ellipsoid, ca. 1 × 1 mm. Weedy in disturbed moist sites such as path edges and riversides, 50–400 m; common throughout Delta Amacuro, Bolívar, and Amazonas. Widespread elsewhere in Venezuela; Mexico, Central America, Antilles, Colombia, Guyana, Surniname, French Guiana, Ecuador, Peru, Brazil, Bolivia. Borreria prostrata has been previously combined with several other taxa under the name B. ocimoides (Burm. f.) DC. [see comments on this situation in the discussion of B. repens]. Within its complex B. prostrata is characterized by its cartilaginous capsules and its seeds ornamented with about 8 longitudinal rows of rounded relatively large pits. Steyermark (in herbaria) apparently sometimes confused B. prostrata with B. hispida. The name Spermacoce gracilis Ruiz & Pav. is probably a synonym of B. prostrata. Borreria pygmaea Spruce ex K. Schum. in Mart., Fl. Bras. 6(6): 58. 1889. Annual herb to 0.3 m tall, hispidulous to glabrescent; leaves 3–8 × 0.5 mm, acute at base; stipule sheath 0.5–1 mm long, setae 3, ca. 0.5 mm long; inflorescences terminal and axillary; flowers 1 or 2; calyx lobes 4, 0.8–2 mm long; corolla white, tube 1.5–2 mm long, lobes ca. 0.8 mm long; fruit ellipsoid, 2.5–3 × 1.5 mm. Sandy savannas and depressions on granite outcrops, 50–200 m; Amazonas (near Puerto Ayacucho and Samariapo), Bolívar (north of Puerto Ayacucho). Endemic. ◆Fig. 433. Borreria pygmaea is distinctive due to its reduced size, its numerous closely grouped small narrow leaves, and its flowers and fruits solitary or paired and enclosed by the subtending leaves and their stipule sheaths. Steyermark (1972, 812) recognized two varieties of this species, with var. robusta Steyerm. distinguished from var. pygmaea by its significantly larger leaves, 15–27 mm long, and larger corollas, 2.6–3 mm long. Although Steyermark explicitly considered these taxa united by intermediate speci-

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mens, the plants he considered intermediates are only so in stature, and have corollas that fall within the measurements of var. pygmaea. Unfortunately, no material has yet been seen by C.M. Taylor of var. robusta, and because Steyermark considered this only a variety he did not provide a full description of it. However, its leaf and flower measurements are so different that it does not seem possible to include these plants in B. pygmaea. Therefore var. robusta is here excluded from B. pygmaea, and although its identity is not established at this time, the details provided by Steyermark are consistent with B. confertifolia. This situation and the interpretation of Steyermark’s taxonomy of Borreria in general are complicated by his strong reliance on pubescence details over other morphological characters to distinguish and key species. Borreria remota (Lam.) Bacigalupo & E.L. Cabral, Darwiniana 37: 334. 1999. —Spermacoce remota Lam., Tabl. Encycl. 1: 273. 1791 [1792]. Spermacoce assurgens Ruiz & Pav., Fl. Peruv. 1: 60, t. 92, fig. c. 1798. —Borreria assurgens (Ruiz & Pav.) Griseb., Abh. Königl. Ges. Wiss Göttingen 19: 159. 1874. Spermacoce capitellata Willd. ex Roem. & Schult., Syst. Veg. 3: 530. 1818. Spermacoce guianensis Bremek., Recueil Trav. Bot. Néerl. 33: 714. 1936. Perennial herb to 0.5(–1.5) m tall, hirtellous to glabrescent; leaves 20–60 × 5–20 mm, acute at base; stipule sheath 2.5–5 mm long, setae 4–7, 1–3.5 mm long; inflorescences terminal and axillary; calyx lobes 4, 0.5–1.2 mm long; corolla white, tube 1–2 mm long, lobes 1–2 mm long; fruit ellipsoid, 2.5–3.5 × 1.5–2 mm. Roadsides, waste ground, riparian forests, along streams, 100–200 m; Bolívar (Caño Aliviadero near Tumeremo), Amazonas (above Platanal). Widely distributed in southeastern U.S.A., Mexico, Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, and adventive in Africa, Asia, and the Pacific Islands. Borreria remota is the most commonly collected species of the genus in Venezuela, but it has been only rarely recorded in the flora area. This species is found from near sea level to as high as 2800 m in the Andes. It has

long been treated under the names B. laevis (Lam.) Griseb. and Spermacoce laevis Lam. However, Verdcourt (1983, 521–571) noted that the type of the name S. laevis actually represents a very different species and in fact is a synonym of S. tenuior L., and he concluded that the species found in the flora area is correctly called S. assurgens. More recently Bacigalupo and Cabral (1999, 333– 334) agreed with Verdcourt in general but applied the name S. laevis to a different species than Verdcourt did, albeit a related one, S. riparia Cham. & Schltdl. Bacigalupo and Cabral did not cite Verdcourt’s work or address any of his conclusions. Bacigalupo and Cabral also came to a different conclusion about the correct name for the species found in the flora area, concurring with Rendle (1936, 10–12) that the oldest name for this species is another Lamarck name, S. remota, which is based on a specimen from Hispaniola (J. Martin s.n., P-LA, microfiche!) and has priority over S. assurgens. Borreria repens DC., Prodr. 4: 544. 1830. —Spermacoce mauritiana Gideon, Kew Bull. 37: 547. 1983, not Spermacoce repens Willd. ex Cham. & Schltdl. 1828. —Camaguari, Coneja sabanera, Orégano. Borreria exilis L.O. Williams, Phytologia 28: 227. 1974. —Spermacoce exilis (L.O. Williams) C.D. Adams, Fieldiana, Bot. n.s. 33: 316. 1993. Herb to 0.4 m tall, often weak-stemmed, hirtellous to glabrescent; leaves 8–40 × 3–13 mm, acute at base; stipule sheath 1–1.5 mm long, setae 7–11, 1–4 mm long; inflorescences terminal and axillary; calyx lobes 2, 0.5–1 mm long; corolla white, 0.8–1 mm long, lobed for ca. 1/2; fruit ellipsoid, ca. 1 × 1 mm. Weedy in moist disturbed sites such as along paths and creeks, in towns, and by gardens, 50–400 m; common in Delta Amacuro, Bolívar, and Amazonas. Widespread elsewhere in Venezuela; Central America, Antilles, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, and widely adventive in the Old World. ◆Fig. 432. The name Borreria ocimoides (Burm. f.) DC. has been applied by many authors to a group of low, weedy, soft Borreria species found throughout the wet tropics and characterized by their relatively short elliptic leaves, stipule setae that are often all about

Borreria 533

the same length, relatively small flowers with the corolla scarcely longer than the calyx lobes, and relatively small capsules. These plants were considered by a number of authors to belong to one morphologically very variable species (e.g., Steyermark 1972; 1974; Bremekamp 1934; Verdcourt 1976), but more recent authors have considered these plants to include several distinct species (Verdcourt 1983; Burger and Taylor 1993), and that the name B. ocimoides does not apply to any of them. Borreria repens is distinguished within this complex by its capsules that are delicately membranaceous and strongly laterally flattened, and by its seeds that are ornamented with numerous closely packed ridges oriented perpendicularly to the longitudinal axis of the seed, so that the seed usually appears irregularly striate. The other species of this complex found in the flora area is B. prostrata. These species are represented by about equal numbers of specimens from the flora area. Borreria ocimoides is based on a plant from Java, and its identity has not been entirely clarified, but workers who have reviewed the situation have not applied this name to any plants found in the Neotropics. The nomenclature of the species in this group no doubt has not been completely worked out yet. The species recognized and the names used for them here follow the work of C.D. Adams (in Burger and Taylor 1993). However, Bacigalupo (e.g., 1974) continues to use the name Borreria ocimoides for apparently native neotropical plants that have 4 calyx lobes and seeds ornamented with about 10– 15 rows of generally rounded, rather small pits. Her application of this name thus corresponds to the plants treated as Spermacoce ovalifolia (M. Martens & Galeotti) Hemsl. by Adams (Burger and Taylor 1993). Plants that correspond to S. ovalifolia have not yet been seen from the flora area, but probably should be expected. Borreria verticillata (L.) G. Mey., Prim. Fl. Esseq. 83. 1818. —Spermacoce verticillata L., Sp. Pl. 102. 1753. Borreria oligodonta Steyerm., Mem. New York Bot. Gard. 23: 826. 1972. Herb or subshrub to 1.5 m tall, hispidulous to glabrescent; leaves 14–55 × 1–15 mm, acute at base; sheath 1.5–2 mm long, setae 5–6, 1–5 mm long; inflorescences termi-

nal and sometimes axillary; calyx lobes 2, 1– 2 mm long; corolla white, tube 0.5–1.2 mm long, lobes 0.5–1.5 mm long; fruit ellipsoid, 1.5–2 × 1–1.5 mm. Sandy and rocky savannas, on sandstone and granite outcrops, open slopes, along streams, disturbed sites, and roadsides, 50–1700 m; common in Delta Amacuro, Bolívar, and Amazonas. Common elsewhere in Venezuela; usually common in southeastern U.S.A., Mexico, Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Brazil, Paraguay, Uruguay, and Argentina, and adventive in Africa and Asia. ◆Fig. 425. Borreria verticillata is distinctive due to its fascicles of small leaves produced in the axils of the vegetative leaves, its calyx lobes 2, and its relatively short corolla that is about as long as the calyx lobes. The number of calyx lobes is consistently 2 in this species in Central America and the Antilles, and in the specimens so far seen from South America. The measurements given here for this species thus differ in number of calyx lobes and several other aspects from those given by Steyermark (1974). Steyermark apparently based his description in part on misidentified specimens. Steyermark separated Borreria oligodonta based on several characters that he himself noted overlapped with those of B. verticillata, including shape of the inflorescences, length of the bracts, and leaf and corolla sizes. Borreria wurdackii Steyerm., Mem. New York Bot. Gard. 23: 812. 1972. Creeping herb to 0.3 m long, scabridulous and hispidulous to glabrescent; leaves 13–27 × 5–11 mm, acute at base; stipule sheath ca. 1 mm long, setae 4 or 5, 1.5–2 mm long; inflorescences axillary and sometimes terminal; calyx lobes 4, 1–1.2 mm long; corolla white, tube ca. 3 mm long, lobes ca. 1.5 mm long; fruit subglobose to ellipsoid, ca. 2 × 1.8 mm. Riparian black-water forests, sandy openings, 100–200 m; Amazonas (known only from the Río Temi between Yavita and Pimichín). Endemic. ◆Fig. 428. Borreria wurdackii is similar in general aspect to B. latifolia, and may eventually be shown to be comprised within the wide morphological variation of that species. However, the creeping stems regularly rooting at the nodes are not evident on specimens of B. latifolia.

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Fig. 422. Borreria macrocephala

Fig. 423. Borreria jangouxii

Fig. 424. Borreria alata

Fig. 425. Borreria verticillata

Borreria 535

Fig. 426. Borreria cataractarum

Fig. 427. Borreria capitata var. tenella

Fig. 428. Borreria wurdackii

Fig. 429. Borreria bolivarensis

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Fig. 430. Borreria latifolia

Fig. 431. Borreria hispida

Fig. 433. Borreria pygmaea

Fig. 432. Borreria repens

Botryarrhena 537

9. BOTRYARRHENA Ducke, Notizbl. Bot. Gart. Berlin-Dahlem 11: 476. 1932. by Charlotte M. Taylor and Julian A. Steyermark Trees, terrestrial, unarmed. Leaves opposite, petiolate, venation not lineolate; stipules interpetiolar or shortly united around the stem, generally imbricate in bud, triangular, persistent. Inflorescence terminal, racemose to racemiform-paniculate, pedunculate, multiflowered, pendulous to erect, bracteate with the bracts small and tuberculiform. Flowers small to medium-sized, pedicellate, homostylous, protandrous. Hypanthium turbinate. Calyx limb well developed, truncate to 5-denticulate, without calycophylls; corolla salverform to funnelform, white, tube generally rather short, internally sericeous in upper part and throat, lobes 5, convolute in bud. Stamens 5, inserted in corolla throat; anthers narrowly oblong, dorsifixed, partially exserted, subsessile. Ovary 2-locular; ovules 2 or 4 in each locule, on axile placentas. Fruit baccate, subglobose, leathery, mature color unknown. Seeds subglobose, smooth. Amazonian Colombia, Venezuela, Amazonian Peru and Brazil; 2 species, both in the flora area. Botryarrhena is infrequently collected and quite poorly known taxonomically, and the distinctions between these species probably will eventually deserve more evaluation. Botryarrhena is similar to and often confused with Stachyarrhena; their separation is discussed under Stachyarrhena.

Fig. 434. Botryarrhena venezuelensis

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Key to the Species of Botryarrhena 1. 1.

Inflorescences pendulous, racemose ............................................. B. pendula Inflorescences erect, paniculate-racemiform ........................ B. venezuelensis

Botryarrhena pendula Ducke, Notizbl. Bot. Gart. Berlin-Dahlem 11: 476. 1932. Tree to 25 m tall; leaves 14–30 × 6–15 cm; stipules 2–5 mm long; inflorescences 4–8.5 × 1.5–2 cm, to 15 × 4 cm in fruit; calyx limb 1–2 mm long; corolla tube 3–5 mm long, lobes 2–3 mm long; fruit 12–15 mm diameter. Evergreen lowland forests, 100–200 m; Amazonas (Caño Caripo, Río Atabapo, alto Río Casiquiare). Amazonian Colombia and Brazil.

Botryarrhena venezuelensis Steyerm., Ann. Missouri Bot. Gard. 70: 207. 1983. Tree to 10 m tall; leaves 17–30 × 9–13.5 cm; stipules ca. 4 mm long; inflorescences ca. 7.5 × 2 cm; calyx limb ca. 1 mm long; corolla tube ca. 4 mm long, lobes ca. 4 mm long; fruit 15–20 mm diameter. Evergreen lowland forests dominated by Leopoldinia palms, 100– 200 m; Amazonas (known only from the middle part of Caño Yagua at Cucurital de Yagua). Amazonian Peru. ◆Fig. 434.

10. CALYCOPHYLLUM DC., Prodr. 4: 367. 1830. Semaphyllanthe L. Andersson, Ann. Missouri Bot. Gard. 82: 421. 1995. by Charlotte M. Taylor and Julian A. Steyermark Trees, often reaching the canopy, unarmed, terrestrial, the bark often smooth and peeling in plates. Leaves opposite, petiolate, venation not lineolate; stipules interpetiolar, triangular and twisted in bud to calyptrate (i.e., united into a conical cap), caducous, sometimes exposing a ring of persistent white or clear trichomes. Inflorescence terminal and in axils of uppermost leaves, corymbose-cymose, pedunculate, multiflowered, bracteate with the bracts often reduced or calyptrate. Flowers small, sessile to pedicellate, homostylous, protandrous, fragrant. Hypanthium ellipsoid to turbinate. Calyx limb truncate or shallowly 4- or 5(–8)-lobed, the lobes equal or in some flowers of most species one of them expanded into a white to cream or pale green, petaloid calycophyll; corolla campanulate to shortly funnelform, white to cream, internally pubescent in upper part of tube, lobes 4 or 5(–8), imbricate or convolute in bud. Stamens 4 or 5, inserted in middle or upper part of corolla tube; anthers narrowly elliptic, dorsifixed, exserted; filaments glabrous or pubescent. Ovary 2-locular; ovules numerous in each locule, on axile placentas. Fruits capsular, ellipsoid to obloid or oblong, septicidal from apex with each valve often later partially splitting from apex, chartaceous to woody. Seeds numerous, small, generally oblong, somewhat flattened, marginally winged. Mexico, Central America, Cuba, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Bolivia, Paraguay, Argentina; ca. 10 species, 3 in Venezuela, 2 of these in the flora area. Calycophyllum is often confused with Bathysa. These genera have similar capsules, similar flowers (including several species with white petaloid calycophylls), generally similar stipules, and similar habits and habitats. These genera are traditionally separated by their seeds, which are generally flattened and clearly winged in Calycophyllum versus angled to somewhat flattened and usually not winged in Bathysa. Calycophyllum is also often confused with Chimarrhis, which can be distinguished by its protogynous flowers and imbricate to valvate corolla lobes. Steyermark apparently separated species of Calycophyllum based primarily on shape of the leaf apex and details of pubescence. Species distinctions in this genus probably deserve some review.

Calycophyllum 539

Delprete (1996, 35–44) and Kirkbride (1997, 354–379) both found intermediates between Semaphyllanthe (with interpetiolar stipules, convolute corolla lobes, and hirsute filaments) and Calycophyllum sensu stricto (calyprate stipules, imbricate corolla lobes, and glabrous filaments), so that Semaphyllanthe is here treated as a synonym of Calycophyllum. Key to the Species of Calycophyllum 1. 1.

Apex of the leaf blade rounded or broadly obtuse; corolla lobes puberulous externally ................................................................................. C. obovatum Apex of the leaf blade acute to acutish; corolla lobes glabrous externally ............................................................................................. C. venezuelense

Calycophyllum obovatum (Ducke) Ducke, Trop. Woods 49: 2. 1937. —Warszewiczia obovata Ducke, Notizbl. Bot. Gart. Berlin-Dahlem 11: 475. 1932. —Semaphyllanthe obovata (Ducke) L. Andersson, Ann. Missouri Bot. Gard. 82: 422. 1995. —Guayabillo. Tree to 15 m tall, with a coppery-red exfoliating bark; leaves 14–25 × 6–16 cm; secondary veins 7–12 pairs; petioles 1–4 cm long; stipules interpetiolar or shortly fused at base, 1.5–2 cm long; inflorescences 20–35 × 18–30 cm (not including calycophylls); calyx limb 0.3–0.6 mm long, calycophylls 4.5–7 × 1.5–3 cm; corolla tube 1.5–2.5 mm long, lobes 4 or 5; capsules 10–14 × 4–6 mm; seeds 7–13 × 2–6 mm. Evergreen lowland to lower montane forests, 100–500 m; Bolívar (Río Acanán near Amaruay-tepui), Amazonas (Cerro Cucurito along the Caño Yagua, Yavita). Colombia (basin of the Río Vaupés and upper Río Negro), Guyana. ◆Fig. 435. Liesner & Holst 20087, MO, of Bolívar, Venezuela, is provisionally included here in the distribution and description of Calyco-

phyllum obovatum. This plant has relatively large leaves, large capsules, and broad seeds. It was previously treated by Steyermark as C. venezuelense, but actually keys out to C. obovatum because of its broadly obtuse to rounded leaf apices. The specimen bears only mature fruits; future flowering collections may show it to be better treated as a separate species.

Fig. 435. Calycophyllum obovatum

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Calycophyllum venezuelense Steyerm., Mem. New York Bot. Gard. 10(5): 189. 1963. —Semaphyllanthe venezuelense (Steyerm.) L. Andersson, Ann. Missouri Bot. Gard. 82: 422. 1995. Tree to 15 m tall, with coppery-red exfoliating bark; leaves 13–20 × 6–9 cm; secondary veins 10–15 pairs; petioles 1–3 cm long; stipules 2.5–3.5 cm long; inflorescences 20– 30 × 20–30 cm; calyx limb 0.5–1.5 mm long, calycophylls 3–7 × 1–3 cm; corolla white, tube 3–4 mm long, lobes 4, 1.5–2 mm long;

capsules 10–20 × 5–6 mm; seeds 6–10 × 2 mm. Moist lowland and lower montane evergreen forests, 200–1100 m; Bolívar (La Escalera, Río Cuyuní, Sierra de Lema), Amazonas (Caño Piedra 115 km S of Puerto Ayacucho). Endemic. As Steyermark separated them, Calycophyllum obovatum is found in Amazonas and Colombia, and C. venezuelense in Bolívar. However, as these species are distinguished here, both species are known from both Venezuelan states.

11. CAPIRONA Spruce, J. Proc. Linn. Soc., Bot. 3: 200. 1859. by Charlotte M. Taylor and Julian A. Steyermark Trees, often reaching the canopy, terrestrial, unarmed, with copper- or brickcolored bark exfoliating in plates. Leaves opposite, petiolate, venation not lineolate; stipules completely fused in bud, later splitting at 90° to the leaves into 2 intrapetiolar ligulate portions, usually caducous. Inflorescences terminal and in uppermost leaf axils, pedunculate, paniculate, multiflowered, bracteate. Flowers sessile and pedicellate, homostylous, protandrous, diurnal, rather large, showy. Hypanthium turbinate. Calyx limb subtruncate to 5-lobed, in some flowers with one lobe enlarged in a pink, red, or cream petaloid calycophyll; corolla campanulate to funnelform, pink, red-purple, or white, internally glabrous except for a pubescent ring near base of tube, lobes 5, convolute in bud. Stamens 5, inserted near base of corolla tube; anthers narrowly oblong, basifixed, included. Ovary 2-locular; ovules numerous, on axile placentas. Fruit capsular, oblong to cylindrical, septicidal from the apex, woody to chartaceous. Seeds small, flattened, papery, margins winged and irregular. Colombia, Venezuela, Peru, Brazil; 1 species. This genus was long considered to include two species (Kirkbride 1985), but Andersson and Taylor (1994, 3–7) have made a convincing case for treating Capirona as comprising one wide-ranging, morphologically variable species. The unusual stipules of Capirona help distinguish it, particularly combined with its coppery-red exfoliating bark. Capirona decorticans Spruce, J. Proc. Linn. Soc., Bot. 3: 299. 1859. Capirona leiophloea Benoist, Bull. Mus. Hist. Nat. (Paris) 27: 367. 1921. Capirona wurdackii Steyerm., Mem. New York Bot. Gard. 10(5): 190, fig. 69. 1963. Tree to 40 m tall; leaves 11–44 × 6–24 cm; stipules 1.5–8 cm long; inflorescence 10–30 × 12–36 cm; calyx limb 3–6 mm long; calycophylls with stipe 1–4.5 cm long and blade 2.5–8.5 × 1–5 cm; corolla with tube 10– 35 mm long, lobes 10–15 mm long; capsules

12–45 × 8–13 mm. Evergreen lowland forests, tepui slope and summit forests, 100– 200 and 1200–1300 m; Amazonas (locally on the summit of Cerro Autana, base of Cerro Sipapo, region of Gavilán south of Samariapo, Río Casiquiare). Amazonian Colombia, Peru, and Brazil. ◆Fig. 436. Capirona decorticans appparently shows some vegetative dimorphism, with juveniles having larger stipules and obovate leaves, versus leaves generally elliptic in mature plants.

Chalepophyllum 541

Fig. 436. Capirona decorticans

12. CHALEPOPHYLLUM Hook. f. in Benth. & Hook. f., Gen. Pl. 2: 50. 1873. by Piero G. Delprete, Julian A. Steyermark, and Charlotte M. Taylor Shrubs, terrestrial, unarmed, gummy-resinous on upper portion of stems at bases of petioles, on peduncles, and on pedicels. Leaves opposite, petiolate to subsessile, on lower surface with golden yellow, finely tomentose, elevated veins, producing a rugose, checkered appearance, but venation not lineolate; stipules interpetiolar, broadly triangular, short, generally persistent with the leaves, in bud generally imbricate to valvate. Inflorescence axillary, pedunculate, with flowers solitary or 2–4 in cymes, bracteate with the bracts reduced. Flowers medium-sized, pedicellate to subsessile, homostylous, protandrous. Hypanthium turbinate to ellipsoid. Calyx limb deeply lobed, the lobes 5, equal or unequal but without calycophylls; corolla salverform or narrowly funnelform, white, internally densely barbate in the mouth of the tube with elongate trichomes, lobes 5, convolute in bud.

542

R UBIACEAE

Stamens 5, inserted in upper part of corolla tube; anthers narrowly oblong, subsagittate or sagittate to bilobed at the base, included, dorsifixed. Ovary 2-locular; ovules numerous in each locule, on apparently axile placentas; style glabrous. Fruit capsular, ellipsoid, subligneous to chartaceous, septicidal from the apex. Seeds small, angular, compressed, narrowly winged or margined, with testa spongyfoveolate. Venezuela, Guyana; 2 species, 1 in Venezuela. Chalepophyllum is similar to and sometimes sympatric with Duidania and Maguireocharis; the separation of these genera is discussed under Maguireocharis. Chalepophyllum guianense Hook. f. in Hook., Icon. Pl. 12. pl. 1148. 1873. —Aporuh-yaré (Pemón). Chalepophyllum guianense var. cuneatum Steyerm., Mem. New York Bot. Gard. 10(5): 193. 1963. Chalepophyllum longilobum Steyerm., Mem. New York Bot. Gard. 10(5): 193. 1963. Shrub 0.4–2.5 m tall; leaf blades 2–9 × 1.3–5.5 cm; stipules 2–6 mm long; calyx lobes 2–17 mm long; corolla tube 1–1.6 cm long, the lobes acute to obtuse at apex, 6–15 × 4– 5.5 mm; capsules 8–17 × 5.5–6.5 mm. Scrub formations in sandy savannas, wet savannas with Stegolepis, open dwarf Bonnetia forests, 1000–1600 m; Bolívar (near Kavanayén, from Ulí-tepui north to Cerro Venamo and west to Ptari-tepui). Guyana. ◆Fig. 437. Steyermark (1963, 193) recognized two species and two varieties in this genus, but recent collections showed that the corolla lobe length and the leaf size and shape of these taxa have overlapping ranges, and

therefore they are here treated as synonymous.

Fig. 437. Chalepophyllum guianense

13. CHIMARRHIS Jacq., Select. Stirp. Amer. Hist. 61. 1763. Pseudochimarrhis Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 255. 1922; 4: 177, pl. 23. 1925. —Chimarrhis subg. Pseudochimarrhis (Ducke) Delprete, Fl. Neotrop. 77: 169. 1999. by Charlotte M. Taylor and Julian A. Steyermark Trees with soft branches, unarmed, terrestrial, often reaching the canopy and with large buttresses, with the wood often yellowish colored. Leaves opposite, often congested at the stem apices, petiolate, venation not lineolate; stipules interpetiolar, usually twisted in bud, ovate to triangular, persistent or caducous. Inflorescences axillary, pedunculate, corymbiform to paniculate, multiflowered, bracteate. Flowers sessile to pedicellate, small, homostylous, protogynous, fragrant. Calyx limb reduced, truncate, or shortly (4)5(6)-lobed, with the lobes equal or in a few species with one lobe expanded in some flowers into a white to greenish white petaloid calycophyll; corolla shortly funnelform, pale green to white, villous in the throat, the lobes (4)5(6), narrowly imbricate but appearing valvate in bud. Stamens 5, inserted in upper part of corolla tube; anthers narrowly oblong, exserted,

Chimarrhis 543

dorsifixed; filaments exserted, villosulous in basal portion. Ovary 2-locular; ovules numerous in each locule, on axile placentas. Fruit capsular, woody, subglobose to turbinate, septicidal from the apex. Seeds small, suborbicular, compressed or turgid, sometimes marginally winged, with the testa reticulate. Central America, Antilles, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 14 species, 3 in Venezuela, 2 of these in the flora area. Chimarrhis has recently been revised by Delprete (1999b), who recognized two subgenera: subgenus Chimarrhis, with 9 species including C. microcarpa; and subgenus Pseudochimarrhis, with 5 species including C. brevipes. The citation here of the place of publication of Chimarrhis follows Delprete’s usage. Chimarrhis is similar to and often confused with Warszewiczia and Bathysa. Warszewiczia can be separated by its usually terminal inflorescences with spiciform, little-branched axes. Bathysa can be separated by its protandrous flowers; fruiting plants of Bathysa are difficult to separate from those of Chimarrhis. Bathysa and Chimarrhis have traditionally been separated by their corolla lobe aestivation, said to be valvate in Chimarrhis versus imbricate in Bathysa. However, Delprete (1999b) has shown that the corolla lobes of Chimarrhis are also imbricate in bud, if narrowly so, so this character does not actually distinguish these genera. Key to the Species of Chimarrhis 1. 1.

Stipules mostly persistent with the leaves; disk pubescent ........ C. brevipes Stipules quickly caducous; disk glabrous ................................ C. microcarpa

Chimarrhis brevipes Steyerm., Mem. New York Bot. Gard. 12(6): 179. 1965. Small tree; leaves 17–23 × 11.5–22 cm, obovate to obpyriform; calyx limb truncate, reduced, rarely with a petaloid calycophyll 5.5 × 3.5 cm on a stipe ca. 2 cm long; corollas unknown; capsules 4.5–6 mm long. Evergreen lowland forests, 100–200 m; Amazonas (base of Cerro Sipapo, near San Carlos de Río Negro). Endemic. ◆Fig. 438. Chimarrhis brevipes has been collected in-

frequently and is characterized by Delprete (1999b) as poorly known. It is very similar to C. gentryana Delprete of the central and western Amazon basin, and these species may be found to actually represent two extremes of continuous variation when more collections are available from the region. Chimarrhis gentryana was reported from the region of San Carlos de Río Negro by H. Clark et al. (2000), but no material of this species has been seen for this flora.

Fig. 438 Chimarrhis brevipes

544

R UBIACEAE

Chimarrhis microcarpa Standl., Bull. Torrey Bot. Club 53: 471. 1926. —Amarilla, Carutillo. Chimarrhis longistipulata Bremek., Recueil Trav. Bot. Néerl. 31: 260. 1934. Canopy tree to 40 m tall, often with strong buttresses; wood very hard with orange heartwood; leaves 10–22 × 5–9.5 cm, elliptic

to obovate; calyx limb truncate to undulate, reduced; corolla 4–5 mm long; capsules ca. 2 mm long. Semideciduous to evergreen lowland forests, near sea level to 600 m; Delta Amacuro (east of Upata), Bolívar (Serranía de Imataca, Sierra de Lema). Anzoátegui, Lara, Monagas, Sucre; Trinidad, Guyana, Suriname, French Guiana, northern Brazil.

14. CHIOCOCCA P. Browne, Civ. Nat. Hist. Jamaica 164. 1756. by Charlotte M. Taylor and Julian A. Steyermark Shrubs, often clambering to twining, unarmed, terrestrial. Leaves opposite, petiolate to subsessile, venation not lineolate; stipules interpetiolar or usually shortly united around the stem, persistent, triangular, acute to acuminate, generally imbricate to valvate in bud. Inflorescence axillary and sometimes also terminal, several-flowered to multiflowered, racemose, paniculate, or infrequently fasciculate or with the flowers solitary, bracteate or the bracts reduced. Flowers small to medium-sized, pedicellate and often secund on inflorescence axes, homostylous, protandrous. Hypanthium ellipsoid, usually laterally flattened. Calyx limb partially lobed, the lobes 4 or 5(6), without calycophylls; corolla campanulate to funnelform, white to yellow or purple, internally glabrous, the lobes 4 or 5, valvate in bud. Stamens 4 or 5, inserted at the base of the corolla tube; anthers narrowly oblong, dorsifixed near the base, included or exserted. Ovary 2(3)-locular; ovules solitary in each locule, pendulous from the locule apex. Fruit drupaceous, suborbicular to ellipsoid, laterally compressed, fleshy to coriaceous or spongy, white to purple; pyrenes 2, hemispherical, laterally compressed, chartaceous to bony, 1-locular. Seeds small, ellipsoid, laterally compressed, smooth. Southeastern U.S.A., Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay; ca. 20 species, 6 in Venezuela, 5 of these in the flora area. Species of Chiococca are often variable morphologically, across their ranges as well as within a local population, and the current taxonomy is provisional and probably includes more species than are warranted. Key to the Species of Chiococca 1. 1. 2(1). 2. 3(2).

Inflorescence 1- or 2-flowered; calyx limb 4–4.5 mm long, lobes 4, lanceolate ........................................................................................... C. lucens Inflorescence 1–75-flowered; calyx limb 0.5–2.5 mm long, lobes 4 or 5, suborbicular to triangular, ovate, or oblong ......................................... 2 Hypanthium and fruit as broad as long or broader than long, orbicular to suborbicular in outline .......................................................................... 3 Hypanthium and fruit generally longer than broad, ellipsoid to oblong in outline .................................................................................................... 4 Inflorescence a simple raceme or branched at the basalmost node, with flowers 1–27, crowded to well spaced, secund or not, with the axes glabrous to pubescent; leaf veins slightly impressed to obscure and glabrous to pubescent on the lower surface, slightly impressed to obscure

Chiococca 545

3.

4(2). 4.

on the upper surface; calyx limb internally glabrous; corollas 5–10 mm long ................................................................................................... C. alba Inflorescence paniculate, generally regularly branched throughout, with flowers 17–75, crowded, not appearing secund, with the axes ± pilosulous; leaf veins conspicuously elevated and ± pubescent on the lower surface, usually conspicuously sulcate on the upper surface; calyx limb internally pubescent; corolla 11–12 mm long ......... C. pubescens Pedicels arising directly from the base of the petiolar axil, epedunculate ....................................................................................... C. auyantepuiensis Pedicels arising from a developed, pedunculate rachis ................... C. nitida

Chiococca alba (L.) A.S. Hitchc., Rep. Missouri Bot. Gard. 4: 94. 1893. —Lonicera alba L., Sp. Pl. 175. 1753. —Babandi, Cruceta de la reina, Cruceto, Francisco Javier. Chiococca racemosa L., Syst. Nat. ed. 10. 917. 1759. Chiococca brachiata Ruiz & Pav., Fl. Peruv. 2: 67, t. 219. 1799. Chiococca parvifolia Wullschl. ex Griseb., Fl. Brit. W.I. 337. 1861. —Chiococca alba subsp. parvifolia (Wullschl. ex Griseb.) Steyerm., Acta Bot. Venez. 6: 138. 1971. Chiococca micrantha J.R. Johnst., Proc. Amer. Acad. Arts 40: 696. 1905. —Chiococca alba [subsp. parvifolia] var. micrantha (J.R. Johnst.) Steyerm., Acta. Bot. Venez. 6: 139. 1971. Chiococca alba [subsp. parvifolia var. micrantha] f. pilosa Steyerm., Acta. Bot. Venez. 6: 141. 1971. Shrub to 3 m tall, usually scandent; leaves 1.5–11 × 0.8–4.5 cm; stipules 0.3–2 mm long; inflorescences 0.5–8.5 cm long; calyx limb 0.7–1.8 mm long; corolla tube 3–6 mm long, lobes 1.8–4 mm long; fruit 4–6 × 4–6 mm. Deciduous to evergreen, lowland to montane forests, 50–1300 m; Bolívar (Altiplanicie de Nuria, Puerto Ordaz, eastern part west to the Río Paragua, San Felix, near Upata, southeast of Villa Lola). Widely distributed elsewhere in Venezuela; U.S.A. (Florida, southwestern Texas), Mexico, Central America, West Indies, Colombia, Guyana, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay. Chiococca alba is commonly found in secondary vegetation in a range of habitats, and is quite variable morphologically. Several authors, notably Steyermark, have recognized several infraspecific taxa based on leaf size

and texture, number of flowers, degree and arrangement of branching of the inflorescences, and presence and distribution of pubescence. However, the variation in these characters is continuous, not closely correlated, and without geographic pattern, and all possible combinations of the distinctive features can be found if enough specimens are examined. Consequently, the infraspecific taxa that Steyermark (1972) keyed and discussed are not recognized here. Chiococca auyantepuiensis Steyerm., Mem. New York Bot. Gard. 23: 374. 1972. Shrub to 1.5 m tall; leaves 3.5–5 × 1.5–3 cm; stipule length unknown; pedicels 5–8 mm long; calyx limb 1–1.3 mm long; corolla and fruit unknown. Thin, open scrub savannas on tepui summits, ca. 1800 m; Bolívar (Auyán-tepui). Endemic. Chiococca lucens Standl. & Steyerm., Fieldiana, Bot. 28(3): 571. 1953. Shrub to 4 m tall; leaves 3.5–8 × 1.5–4 cm; stipules 2–3 mm long; peduncles 1–3 cm long, pedicels 6–8 mm long; calyx limb 4–4.5 mm long; corolla tube ca. 9 mm long, lobes ca. 2 mm long; mature fruit unknown. Evergreen, densely forested slopes, 2100–2200 m; Bolívar (narrow ridge of Sororopán-tepui). Endemic. Chiococca nitida Benth., J. Bot. (Hooker) 3: 236. 1841. Shrub to 6 m tall, sometimes clambering; leaves 4.5–18 × 3–8 cm; stipules 1–3 mm long; inflorescences 1.5–6 cm long; calyx limb 0.5–2.5 mm long; corolla tube 5–12 mm long, lobes 1.2–2.5 mm long; fruit 7–9 × 4–5 mm. Venezuela, Guyana, French Guiana, Brazil; 3 varieties, all in the flora area.

546

R UBIACEAE

Key to the Varieties of C. nitida 1. Corolla ca. 14.5 mm long; inflorescence 3– 6 cm long, with basalmost axes 2–3 cm long ........................ var. chimantensis 1. Corolla 6.5–12 mm long; inflorescence 1.5–3 cm long, with basalmost axes 0.3– 1 cm long ........................................... 2 2. Corolla 6.5–7.5 mm long; leaf blades broadly ovate to lance-oblong, obtuse to rounded, truncate, or cordate at base .................................... var. amazonica 2. Corolla 8–12 mm long; leaf blades lanceoblong to oblong-elliptic, acute to obtuse at base ............................... var. nitida C. nitida var. amazonica Müll. Arg. in Mart., Fl. Bras. 6(6): 50. 1889. Evergreen and gallery forests, 400–1300 m; Bolívar (Altiplanicie de Nuria, Gran Sabana, Piacoa). Guyana, northeastern Brazil. C. nitida var. chimantensis Steyerm., Mem. New York Bot. Gard. 23: 376. 1972. Open rocky slopes, ca. 1400 m; Bolívar (Macizo del Chimantá [southwest-facing slopes of Toronó-tepui]). Endemic.

C. nitida var. nitida Chiococca erubescens Wernham, J. Bot. 51: 323. 1913. Evergreen and gallery forests, 500–1400 m; Bolívar (common in Altiplanicie de Nuria and Gran Sabana), Amazonas (Cerro Ualipano). Guyana, French Guiana, northeastern Brazil. ◆Fig. 439. Chiococca pubescens Humb. & Bonpl. ex Roem. & Schult., Syst. 5: 202. 1819. Shrub or tree to 4 m tall, often clambering; leaves 5–9 × 2–4.8 cm; stipules 1.5–2 mm long; inflorescences 3–6.5 cm long; calyx limb 0.8–1.5 mm long; corolla tube 8–9 mm long, lobes 2.5–3 mm long; fruit 5–6 × 5–6 mm. Evergreen forests, 50–900 m; Delta Amacuro (Manoa), Bolívar (Kavanayén, Santa Elena de Uairén). Sporadically but generally distributed elsewhere in Venezuela; Colombia, Trinidad, Tobago, Ecuador. Given the extensive morphological variation found among plants of Chiococca alba, it is conceivable that with rigorous re-evaluation of the taxonomy of Chiococca, the plants Steyermark separated as C. pubescens will be included in C. alba.

Fig. 439. Chiococca nitida var. nitida

Chomelia 547

15. CHOMELIA Jacq., Enum. Syst. Pl. 12: 1760. Anisomeris C. Presl, Symb. Bot. 2: 5, pl. 54. 1858 [1834]. by Charlotte M. Taylor and Julian A. Steyermark Shrubs, small trees, or lianas, terrestrial, sometimes armed with axillary spines. Leaves opposite, petiolate, venation sometimes lineolate; stipules interpetiolar, triangular, persistent or deciduous, acute to acuminate, twisted in bud. Inflorescence axillary, sessile to pedunculate, bracteate, capitate to shortly cymose or paniculate and multiflowered or sometimes with the flowers solitary. Flowers medium-sized, sessile to pedicellate, protandrous, distylous or perhaps sometimes homostylous. Hypanthium turbinate to ellipsoid. Calyx limb deeply lobed, the lobes 4, equal to unequal, without calycophylls; corolla salverform or slenderly funnelform, white or cream-colored, externally usually strigillose to sericeous, internally glabrous, lobes 4, valvate to valvate-conduplicate in bud (or reportedly sometimes subimbricate), with the margins sometimes crisped to appendaged. Stamens 4, inserted in corolla throat; anthers dorsifixed near middle, narrowly oblong, included or exserted. Ovary 2-locular; ovules solitary in each locule, pendulous from its apex. Fruit drupaceous, ellipsoid to cylindrical, fleshy, red to black; pyrene 1–3-locular, smooth to ridged. Seeds small, cylindric, pendulous. Central America, Antilles, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, tropical Asia; ca. 50 species, 13 in Venezuela, 9 of these in the flora area. Anisomeris was distinguished from Chomelia by its crisped to appendaged corolla lobes, but several species are intermediate and variable in this respect so this genus is no longer recognized (Steyermark 1967). The name Chomelia has been used incorrectly by some authors for several Old World Rubiaceae genera, notably Tarenna Gaertn. (ca. 200 species), and this usage has generated much confusion about the distribution and size of this genus. Chomelia is frequently and easily confused with Guettarda, Machaonia, and Malanea. The separation of Chomelia and Machaonia is discussed under Machaonia. Guettarda differs from Chomelia in its corolla lobes that are imbricate in bud; fruiting specimens of these genera are often difficult to assign to genus. Malanea has traditionally been separated from Chomelia (e.g., Steyermark 1974) by its relatively short tubular to campanulate corollas and climbing habit. However a few species of Chomelia are also climbers and may be difficult to separate from Malanea when the distinctive salverform corollas or axillary spines of Chomelia are not present on the individual plant. Key to the Species of Chomelia 1. 1. 2(1). 2. 3(2).

Higher-order leaf venation lineolate (i.e., closely parallel); bracts 0.5–5+ mm long .................................................................................................. 2 Higher-order leaf venation aerolate (i.e., irregular); bracts reduced or none ........................................................................................................ 5 Peduncles 1–6 cm long, rather stout; corolla tubes 30–35 mm long; fruits 20–25 × 10 mm; leaves 5–15 × 2.5–7.5 cm ........................ C. malaneoides Peduncles 0–35 cm long, slender; corolla tubes 10–30 mm long; fruits 8– 10 × 3–4 mm; leaves 1–9.5 × 0.5–5.5 cm .............................................. 3 Flowers 1–5 per axil, sessile or subsessile, the peduncles none or 1–5 per axil, to 3 mm long ................................................................... C. tenuiflora

548

3. 4(3). 4. 5(1). 5. 6(5). 6. 7(6). 7. 8(7). 8.

R UBIACEAE

Flowers 3–5 per peduncle, the peduncles 1 per axil, 15–30 mm long ..... 4 Calyx lobes 1.5–3 mm long, linear, equal ............................... C. monachinoi Calyx lobes 1.8–5 mm long, narrowly triangular to oblanceolate, usually unequal on an individual flower ...............................................C. volubilis Calyx lobes 2–5 mm long, narrowly elliptic, broadest above base, glabrescent to glabrous ......................................................................... C. stergiosii Calyx lobes 0.5–2 mm long, ligulate to triangular, broadest at base, strigillose to glabrous .................................................................................. 6 Inflorescences borne on lateral short shoots produced along developed long shoots, with peduncles 3–10 mm long ............................ C. caurensis Inflorescences mostly or all borne on developed long shoots, with peduncles 5–45 mm long ........................................................................... 7 Calyx and hypanthium glabrous; corolla with tube 2.5–3 mm long ............................................................................................... C. glabricalyx Calyx and hypanthium strigillose; corolla with tube 8–20 mm long ...... 8 Leaves 1.5–5.5 × 0.7–2.2 cm, with secondary veins plane adaxially ................................................................................................... C. delascioi Leaves 3.5–13.5 × 2–8 cm, with secondary veins impressed adaxially at least on larger leaves .............................................................. C. polyantha

Chomelia caurensis (Standl.) Steyerm., Mem. New York Bot. Gard. 17(1): 335. 1967. —Anisomeris caurensis Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 22: 172. 1940. Small tree or shrub to 8 m tall; leaves 3.5– 15 × 1.5–5.5 cm; secondary veins 3–6 pairs; higher-order venation not lineolate; petioles 2–5 mm long; stipules 1–2 mm long; peduncles 3–10 mm long; flowers 3–7, subcapitate; calyx lobes 1–1.5 mm long; corolla cream-colored, tube ca. 6 mm long, lobes ca. 2 mm long; fruit not seen. Riparian forests, 50–200 m; Bolívar (Guayapo, Río Caura). Endemic. Chomelia delascioi Steyerm., Mem. Soc. Ci. Nat. La Salle 113: 83. 1980. Shrub or small tree to 4 m tall; leaves 1.5– 5.5 × 0.7–2.2 cm; secondary veins 3–5 pairs; higher-order venation not lineolate; petioles 1–3 mm long; stipules 1–1.5 mm long; peduncles 5–30 mm long; flowers 2–5, subcapitate; calyx lobes 1–2 mm long; corolla yellow, tube 11–15 mm long, lobes 2.5–3 mm long; fruits oblanceolate, 7–9 × 3–4 mm, purple. Evergreen lowland forests, 100–200 m; Bolívar (Caño Negro, Isla Anacoco, basin of Río Cuyuní, San Martín de Turumbán southwest of Tumeremo). Endemic. Chomelia glabricalyx Steyerm., Ann. Missouri Bot. Gard. 74: 657. 1987.

Shrub ca. 1 m tall; leaves 4.5–9.5 × 1.5–4 cm; secondary veins 5 or 6 pairs; petioles 2–5 mm long; stipules ca. 0.5 mm long; higher-order venation apparently not lineolate; peduncles 14–20 mm long; flowers ca. 6, subcapitate to shortly pedicellate; calyx lobes 0.5–1 mm long; corolla color not noted, tube 2.5–3 mm long, lobes ca. 1.5 mm long; fruit narrowly cylindrical, ca. 10 × 3.5 mm. Riparian forests, 200–300 m; Bolívar (Río Caura upstream from Salto Pará). Endemic. No material of Chomelia glabricalyx was seen by C.M. Taylor for this treatment; the information here is taken from the original description. Based on the species to which Steyermark compared this, the higher-order venation is apparently areolate (or perhaps not visible). The corolla is remarkably small for Chomelia, and the generic placement of this species may need re-evaluation. Chomelia malaneoides Müll. Arg., Flora 58: 452. 1875, emend. Steyerm. in Lasser & Steyerm., Fl. Venez. 9(1): 861. 1974. —Anisomeris malaneoides (Müll. Arg.) Standl., Publ. Field Columbian Mus., Bot. Ser. 8: 361. 1931. Shrub or small tree to 7 m tall; leaves 5– 15 × 2.5–7.5 cm; secondary veins 6–8 pairs; higher-order venation lineolate; petioles 0.8– 2 cm long; stipules 5–8 mm; peduncles 1–6 cm long; flowers 10–20, subcapitate; calyx lobes 3–5 mm long; corollas cream-colored,

Chomelia 549

tube 30–35 mm long, lobes 5–6 mm long; fruits ellipsoid-oblong, 20–25 × 10 mm. Riparian forests, 100–200 m; Amazonas (La Esmeralda, Río Casiquiare, Río Siapa, Río Yureba). Guyana, Amazonian Ecuador, Peru, Brazil, Bolivia. ◆Fig. 440. Steyermark (1972, 372) distinguished Chomelia glabriuscula Steyerm. of French Guiana from C. malaneoides based only on its glabrous to glabrescent calyx limb, hypanthium, and leaves, and he reported C. glabriuscula from Amazonas in the headwaters of the Río Baría and the base of Sierra de la Neblina. However, plants from the flora area show continuous variation from glabrescent to densely strigillose on these structures, and consequently C. glabriuscula cannot be separated from C. malaneoides in the flora area and is not included here.

microloba Donn. Sm. of Central America and C. angustifolia Benth. of Guyana. Steyermark distinguished C. angustifolia by its glabrous hypanthium, corolla tube exterior, and leaves and its sparsely strigillose to glabrous calyx limb. Chomelia angustifolia has relatively small leaves, 2.5–6.5 × 1–3.5 cm, with the secondary veins plane adaxially and the higher-order venation not lineolate, inflorescences that may be borne on long shoots, short shoots, or both on the same branch, and corolla tubes ca. 10 mm long. Probably C. angustifolia and C. polyantha as well as C. stergiosii, C. caurensis, and C. delascioi are all closely related, and at least some of these may eventually be shown to be morphologically well marked but only partially isolated populations of one variable species rather than distinct taxa.

Chomelia monachinoi Steyerm., Mem. New York Bot. Gard. 23: 336. 1972. Scandent or clambering shrub to 3 m tall; leaves 1.5–10 × 1–4.5 cm; secondary veins 5 or 6 pairs; higher-order venation lineolate; petioles 3–10 mm long; stipules 3–10 mm long; peduncles 0.5–2.5 cm long; flowers 2 or 3, subcapitate; calyx lobes 1.5–3 mm long; corolla white, tube 10–16 mm long, lobes 4.5–5 mm long; fruits ca. 8 × 3 mm, ellipsoid-oblong. Mauritia palm swamps, riparian forests, 50–200 m; Bolívar (El Carmen, Río Parquaza). Apure, Guárico.

Chomelia stergiosii Steyerm., Ann. Missouri Bot. Gard. 74: 105. 1987. Shrub or small tree to 8 m tall; leaves 2.5– 9 × 1–4 cm; secondary veins 4 or 5 pairs, higher-order venation not lineolate; petioles 2–5 mm long; stipules 1–2 mm long; peduncles 1.5–2 mm long; flowers 3–7, congested cymose to subcapitate; calyx lobes 2–5 mm long, often unequal on an individual flower; corolla cream-colored to yellow, tube ca. 15 mm long, lobes 2–3 mm long; fruits oblanceoloid to cylindrical, ca. 10 × 4 mm. Evergreen lowland and riparian forests, 100–200 m; Bolívar (Isla Anacoco, mouth of Río Botanamo and Río Guarampín, basin of Río Cuyuní). Endemic. Chomelia stergiosii is similar to C. polyantha. They can be distinguished by their calyx lobes: these are equal, strigillose throughout, triangular, and broadest at the base in C. polyantha, versus subequal to strongly unequal, glabrescent, narrowly elliptic, and broadest near the middle in C. stergiosii. The type specimen of C. transiens Müll. Arg. [Glaziou 10940, B, photo (Rockefeller neg. #414) MO!] is aparently indistinguishable from C. stergiosii. However, this was reportedly collected near Rio de Janeiro, which if true suggests these taxa are perhaps related but not conspecific.

Chomelia polyantha S.F. Blake, Contr. U.S. Natl. Herb. 20: 532. 1924. —Anisomeris polyantha (S.F. Blake) Rusby, Bull. Torrey Bot. Club 52: 142. 1925. —Cruceta, Espuelito montañero, Guacharaca, Macollo, Punteral. Shrub or medium-sized tree to 15 m tall; leaves 3.5–13.5 × 2–8 cm; secondary veins 3– 5 pairs, higher-order venation not lineolate; petioles 3–5 mm long; stipules 2–4.5 mm long; peduncles 1–5.5 cm long; flowers (3)5– 17, subcapitate to congested-cymose; calyx lobes 1.5–2 mm long; corollas cream-colored to pale yellow, tube 8–15 mm long, lobes 3–4 mm long; fruits oblong-ellipsoid to oblanceoloid, 10–13 × 3–4 mm. Evergreen and semideciduous forests, 200–300 m; Bolívar (base of Serranía Tigre). Distrito Federal, Falcón, Miranda, Sucre, Yaracuy, Zulia; northeastern Colombia, northern Brazil. Chomelia polyantha is similar to C.

Chomelia tenuiflora Benth., J. Bot. (Hooker) 3: 235. 1841. —Anisomeris tenuiflora (Benth.) Pulle, Enum. Vasc. Pl. Suriname 466. 1906.

550

R UBIACEAE

Chomelia barbellata Standl., Field Mus. Nat. Hist., Bot. Ser. 13(6): 120. 1936. Shrub or small tree to 6 m tall; leaves 1– 13 × 0.5–6 cm; secondary veins 3–10 pairs; higher-order venation lineolate; petioles 2– 15 mm long; stipules 2–6 mm long; peduncles 0–3 mm long; flowers 1–5; calyx limb 3–4 mm long; corolla pale green to white, tube 13–30 mm long, lobes 5–6 mm long; fruits ellipsoid, ca. 10 × 4 mm. Evergreen forests, 100–600 m; Amazonas (Cerro Aracamuni, base of Cerro Duida, Río Mavaca, Río Siapa). Apure; Central America, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Bolivia, Brazil. Chomelia tenuiflora is circumscribed here a bit differently and more strictly than was done by Steyermark, who placed great weight on the presence of fused basal-most bracts in the inflorescence to distinguish this species but considered the inflorescence to vary from a sessile solitary flower to a pedunculate panicle. The type of C. tenuiflora has flowers that are subsessile and solitary in the leaf axils, and C. tenuiflora as circumscribed here is distinguished by this inflorescence arrangement. Steyermark additionally included in C. tenuiflora specimens with peduncles 1–4 cm long and more numerous flowers, to 30–35 in some cases. These specimens have inflorescences subtended by bracts that are sometimes fused, and thus readily key out to C. tenuiflora in Steyermark’s (1967; 1974) treatments. However, this second group of plants is here referred to other species. In particular, a number of plants from western Venezuela with peduncles 1–4 cm long are here treated as C. paniculata (Bartl. ex DC.) Steyerm., which

Fig. 440. Chomelia malaneoides

was been previously reported for Venezuela, and the plants from Guyana with peduncles 0.5–1.5 cm long are referred to C. schomburgkii Steyerm. Chomelia volubilis (Standl.) Steyerm., Mem. New York Bot. Gard. 17(1): 340. 1967. —Anisomeris volubilis Standl., Publ. Field Columbian Mus., Bot. Ser. 8: 58. 1930. Vining or clambering shrub to 6 m tall; leaves 2.5–8.5 × 1.5–4 cm; secondary veins 5 or 6 pairs; higher-order venation lineolate; petioles 2–6 mm long; stipules 3–5 mm long; peduncles 1.5–4 cm long; flowers 2–4, subcapitate; calyx lobes 1.8–5 mm long; corolla white, tube 15–24 mm long, lobes 4–5 mm long; fruits ellipsoid, ca. 8 × 3 mm. Evergreen lowland forests, ca. 100 m; Amazonas (Río Guainía). Endemic. Chomelia volubilis may not be distinct from C. monachinoi. These were distinguished by Steyermark primarily by the length of the calyx lobes, but the informative value of this character in Chomelia is not completely clear and may not be as high as Steyermark thought.

Cinchonopsis 551

16. CINCHONOPSIS L. Andersson, Ann. Missouri Bot. Gard. 82: 424. 1995. by Charlotte M. Taylor Trees, unarmed, terrestrial. Leaves opposite, petiolate, venation not lineolate; stipules interpetiolar or shortly united around the stem, caducous, lingulate to triangular, erect and flatly appressed in bud. Inflorescences terminal and in uppermost leaf axils, pedunculate, paniculate, expansive and multiflowered, bracteate. Flowers small, subsessile, homostylous, protandrous. Hypanthium turbinate. Calyx limb 5-lobed, without calycophylls; corolla campanulate, white, internally pubescent in the upper part of the tube and on the adaxial faces of the lobes, lobes 5, valvate in bud. Stamens 5, inserted near middle of corolla tube; anthers narrowly ob-

Fig. 441. Cinchonopsis amazonica

552

R UBIACEAE

long, dorsifixed near base, partially exserted, with the connective prolonged into a short apical extension. Ovary 2-locular; ovules numerous in each locule, on axile placentas. Fruit capsular, narrowly cylindrical to fusiform, papery, septicidal usually from the base. Seeds small, flattened, papery, margins winged and irregular. Colombia, Venezuela, Brazil, Peru; 1 species. This species was long included in Cinchona, but it is anomalous in that genus due to its relatively small campanulate white corollas. Because its generic relationships are not clear, Andersson and Taylor (1994) placed it in a monotypic genus. The genera of the flora area that are similar to Cinchonopsis and their separation are discussed under Ladenbergia. Cinchonopsis amazonica (Standl.) L. Andersson, Ann. Missouri Bot. Gard. 82: 424. 1995. —Cinchona amazonica Standl., Publ. Field Columbian Mus., Bot. Ser. 8: 334. 1931. Tree to 18 m tall; leaves 15–30 × 7–18 cm; stipules 1–1.5 cm long. Inflorescences 16– 30+ × 14–25+ cm; calyx limb ca. 2 mm long;

corolla with tube ca. 3 mm long, lobes ca. 1 mm long; capsules ca. 10 × 2 mm. Evergreen lowland to lower montane forests, 100–600 m; Bolívar (85 km south of Entre Ríos, Santa María de Erebato), Amazonas (Río Orinoco and tributaries). Colombia, Peru, Brazil (Acre, Amazonas, Roraima). ◆Fig. 441.

17. COCCOCHONDRA Rauschert, Taxon 31: 561. 1982. —Chondrococcus Steyerm., Mem. New York Bot. Gard. 23: 403. 1972, hom. illeg., non Kützing 1847. by Charlotte M. Taylor and Julian A. Steyermark Low shrubs, terrestrial, unarmed. Leaves opposite, sessile to subsessile, venation not visible; stipules interpetiolar, persistent, triangular, sometimes with setae attached medially, generally imbricate or valvate in bud. Inflorescence axillary, subsessile, subcapitate, bracteate, with the bracts fused in pairs. Flowers small, sessile, distylous, probably protandrous. Hypanthium turbinate to cylindrical. Calyx limb with short tube, the lobes 4, without calycophylls; corolla shortly funnelform, white, internally hirtellous in upper part of tube, lobes 4, valvate in bud. Stamens 4, inserted in corolla throat; anthers narrowly oblong, dorsifixed near middle, partially exserted. Ovary 2-locular; ovules 1 in each locule, basal. Fruit apparently drupaceous, ellipsoid, dry, indehiscent; pyrenes 2, 1-locular, crustaceous. Seeds small, planoconvex. Venezuela (endemic to the flora area); 1 species. Kirkbride and Robbrecht (1984) summarized the nomenclatural history of this genus and discussed its systematically informative characters. They, as well as Steyermark (1972, 403–404), described the fruits as indehiscent. However, Steyermark later (1974, 1021) described the fruits as “subcapsular” and septicidally dehiscent from the apex, without comment regarding the difference from the earlier descriptions. Coccochondra laevis (Steyerm.) Rauschert, Taxon 31: 561. 1982. —Chondrococcus laevis Steyerm., Mem. New York Bot. Gard. 23: 405, fig. 65. 1972. Shrub to 2 m tall, glabrous; leaves 1.7–4 × 0.6-1.6 cm; stipules 0.1–2 mm long; inflores-

cences 9–16-flowered; calyx limb 1–1.7 mm long; corolla tube 1.8–2.5 mm long, lobes 1.5– 2 mm long; fruit 3–5.5 × 2.5–3.5 mm. Tepuis, 2000–2100 m. Venezuela; 2 subspecies, both endemic to the flora area.

Coccocypselum 553

Key to the Subspecies of C. laevis 1. Youngest stem internodes 4–15 mm long; stipules smooth; leaves narrowly oblong to ligulate-oblong, 2.5–3.8 × 0.8–1.6 cm; calyx lobes 0.6–0.7 mm long ................... ........................................ subsp. laevis 1. Youngest stem internodes 3–5 mm long; stipules with 6–8 setae 0.6–1 mm long, these inserted medially; leaves elliptic, 1.7–2 × 0.6–0.8 cm; calyx lobes 1.2–1.6 mm long ............. subsp. maigualidae C. laevis subsp. laevis Exposed tepui summits and slopes, ca. 2000 m; Amazonas (summit of Cerro Parú). Endemic. ◆Fig. 442. C. laevis subsp. maigualidae J.H. Kirkbr., BioLlania Ed. Esp. 6: 393. 1997. Savannas on high granitic mountain tops, ca. 2100 m; Bolívar (summit of Sierra de Maigualida). Endemic.

Fig. 442. Coccochondra laevis subsp. laevis

18. COCCOCYPSELUM P. Browne, Civ. Nat. Hist. Jamaica 144, tab. 6, fig. 1 (miscited as fig. 2). 1756, nom. et orth. cons. Tontanea Aubl., Hist. Pl. Guiane 108, t. 42. 1775. Condalia Ruiz & Pav., Fl. Peruv. Prodr. 1: 54. 1794, nom. rej. Lipostoma D. Don, Edinburgh New Philos. J. 8: 168. 1830. by Cristina Bestetti Costa and Julian A. Steyermark Annual or perennial herbs, terrestrial, unarmed; stems prostrate, creeping, decumbent, or erect-ascending. Leaves opposite, petiolate, membranceous to carnose, sometimes purple or blue-lavender on lower surface, venation not lineolate; stipules interpetiolar, relatively small, triangular, entire, persistent, in bud generally valvate to imbricate. Inflorescence axillary, capitate or glomerate, sessile to pedunculate, 1–20-flowered, bracteate. Flowers small, sessile, distylous, protandrous. Hypanthium ovoid or turbinate. Calyx limb lobed, the lobes 4, usually narrow and elongated, generally unequal but without calycophylls; corolla funnelform, blue, purple, or white, internally glabrous, the lobes 4, valvate in bud. Stamens 4, inserted in the corolla tube; anthers included or exserted, dorsifixed near base, narrowly oblong; filaments short. Ovary 2-locular; ovules numerous in each locule, on axile placentas. Fruit baccate, ovoid or globose, spongy and often hollow, blue or purple, often very brightly colored. Seeds minute, orbicular, subangulate, or planoconvex; testa granulate. Mexico, Central America, West Indies, Colombia, Venezuela, Trinidad-Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Bolivia, Brazil, Paraguay; ca. 22 species, 8 in Venezuela, 6 of these in the flora area. Coccocypselum is often recognizable by its deep bright blue fruits that are rather dry in texture and hollow.

554

R UBIACEAE

Coccocypselum apurense Steyerm. was described based on a single collection from Apure (distrito Pedro Camejo, Caño El Cabello, 16 airline km northwest of Mata de Guanabana, Davidse & González 15825, MO). It probably occurs in the flora area, although as discussed under Coccocypselum condalia, below, it is part of the complex of forms generally included in that species and may not be distinct. Key to the Species of Coccocypselum 1. 1. 2(1). 2. 3(2).

3.

4(2). 4. 5(4). 5. 6(5).

6.

Stems, petioles, and leaves wholly glabrous or nearly so ........... C. condalia Stems, petioles, and/or leaves ± pubescent ............................................... 2 Stems with trichomes spreading, often woolly ......................................... 3 Stems with appressed pubescence ............................................................ 4 Calyx lobes, hypanthium, and fruit with loosely spreading hispidulous pubescence; petioles 8–27 mm long, 1/4–3/4 as long as to equal to the length of the leaf blade; leaf blades with the trichomes on the lower surface much shorter than those on the secondary veins and midrib, with the secondary veins impressed on the upper surface and manifest but not conspicuously elevated on the lower surface ............ C. guianense Calyx lobes conspicuously hirsute or hispid with cinereous or tawny trichomes, the hypanthium and fruit with loosely ascending to subappressed trichomes; petioles 4–7 mm long, 1/12–1/7 as long as the leaf blade; leaf blades densely hirsute on both surfaces with trichomes 1– 1.5 mm long, the trichomes on the lower surface as long as those on the secondary veins and midrib, with the secondary veins sulcate on the upper surface and conspicuously elevated on the lower surface .................................................................................................. C. hirsutum Inflorescences 1-flowered; stipules ca. 1.2 mm long ........................ C. huberi Inflorescences 3–20-flowered; stipules 2–9 mm long ............................... 5 Stems, leaves, peduncles, hypanthium, and calyx lobes with long, spreading or subappressed, conspicuous trichomes ......................... C. guianense Stems, leaves, peduncles, hypanthium, and calyx lobes with short villosulous or densely soft-tomentose pubescence ........................................ 6 Calyx lobes eventually squarrose or recurved, subfoliaceous, oblanceolate, obovate, or broadly elliptic, 0.7–2 mm wide; inflorescences with heads glomerate, 10–20-flowered, subsessile to usually pedunculate ............................................................................................. C. lanceolatum Calyx lobes erect, linear-lanceolate to lanceolate, 0.3–1 mm wide; inflorescences with heads capitate, 4–8-flowered, pedunculate to usually sessile or subsessile .................................................................... C. aureum

Coccocypselum aureum (Spreng.) Cham. & Schltdl., Linnaea 4: 139. 1829. —Schwenkfeldia aurea Spreng., Neue Entdeck. Pflanzenk. 1: 280. 1820. Rondeletia capitatum Benth., J. Bot. (Hooker) 3: 217. 1841. —Coccocypselum aureum var. capitatum (Benth.) Steyerm., Mem. New York Bot. Gard. 17: 307. 1967. Creeping perennial herbs; stems in upper part lavender- or purple-villose; leaf second-

ary veins 9–11 pairs; inflorescence capitate, sessile, subsessile, or with peduncles 3–5 cm long, with flowers 4–8; hypanthium villose; calyx lobes erect, linear-lanceolate to lanceolate, villose; fruits ellipsoid to obovoid, bright blue. Evergreen montane forests, savannas, streams, 800–1500 m; Bolívar (vicinity of Roraima-tepui and Kavanayén). Anzoátegui; Cuba, Colombia, Guyana, Peru, Brazil, Bolivia. ◆Fig. 443.

Coccocypselum 555

Steyermark (1967, 299–307) recognized two varieties of this species, with only one of them, var. capitatum, found in the flora area. These were distinguished by the inflorescence arrangement, sessile in var. aureum versus pedunculate in var. capitatum. However, peduncle length varies continuously and independently of other features, and this variety is not recognized here. Coccocypselum condalia Pers., Syn. Pl. 1: 132. 1805. —Condalia repens Ruiz & Pav., Fl. Peruv. 1: 54, t. 84, fig. a. 1798, not Coccocypselum repens Sw. 1788. Coccocypselum croatii Steyerm., Ann. Missouri Bot. Gard. 74: 106. 1987. Creeping perennial herb, glabrous or nearly so; leaf secondary veins 7 or 8 pairs; inflorescence capitate, pedunculate, with 7 or 8 flowers; hypanthium glabrous; calyx lobes oblanceolate to obovate, glabrous or ciliate; fruits ellipsoid to obovoid, bright blue. Sandy open places in dry upland valleys, forests, savannas, and lower forested slopes, 400–900 m; Bolívar (near El Paují, vicinity of Icabarú). Colombia, Ecuador, Peru, Brazil, Paraguay, Argentina. Coccocypselum condalia is widely distributed in South America. Steyermark described two new species from Venezuela, C. croatii (from Bolívar, vicinity of Icabarú), and C. apurense (from Apure, locality cited in the genus description, above). Both of these were distinguished based largely on differences in leaf and inflorescence sizes and the indument on the leaf blade. However C. croatii falls within the variation included in C. condalia, and C. apurense is very similar and when it is better known it will probably also be found to be better included in C. condalia. Coccocypselum condalia is closely related to C. guianense; C. condalia can be distinguished by its usually glabrous or ciliate stems, leaves, and peduncles, or its pubescence when present appressed and with short trichomes (versus loosely spreading with long trichomes in C. guianense); by its glabrous or ciliate hypanthium and calyx lobes (versus hispidulous or sericeous externally though glabrous at least in the lower part internally in C. guianense); and by its ovoid to ellipsoid blue fruit (versus globose and whitened to purple when young, becoming oblong and bright blue in C. guianense).

Coccocypselum guianense (Aubl.) K. Schum. in Mart., Fl. Bras. 6(6): 315. 1889. —Tontanea guianensis Aubl., Hist. Pl. Guiane 108, t. 42. 1775. —Nabajoso. Bellardia repens Willd., Sp. Pl. 1: 626. 1797. Coccocypselum tontanea Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 3: 406. 1818 [1820]. —Bellardia tontanea (Kunth) Schult. & Schult. f., Mant. 3: 147. 1827. Coccocypselum guianense var. patens Steyerm., Mem. New York Bot. Gard. 17: 303. 1967. Prostrate, creeping, or procumbent perennial herb, loosely strigillose, hirsutulous, or hispidulous; leaf secondary veins ca. 6 pairs; inflorescence capitate, pedunculate, with 3–6 flowers; hypanthium densely strigillose; calyx lobes lanceolate, externally sparsely sericeous, internally glabrous or pubescent in the upper half; fruits globose and whitened to purple when young, becoming oblong and bright blue. Forested slopes, mossy rocks, borders of savannas, along streams, 50–1300 m; Delta Amacuro (mouth of Caño Araguabisi), Bolívar (Altiplanicie de Nuria, Cerro Venamo west to Macizo del Chimantá, Gran Sabana, Roraima-tepui), Amazonas (near San Carlos de Río Negro). Trinidad, Guyana, Suriname, French Guiana, Brazil (Pará, Roraima). ◆Fig. 444. The corollas of Coccocypselum guianense range from lavender or blue to pale blue or white. It is closely related to C. condalia; see comments under this latter species, above. Two varieties of Coccocypselum guianense have been recognized, both from the flora area, based on the form of the pubescence (appressed in the typical variety versus spreading in var. patens). However, in the Venezuelan plants there is extensive variation in the form of the pubescence. Young plants may be covered with trichomes variously short and purplish to long, spreading, and conspicuous, with the pubescence, strigillose, subappressed, hirsute, or hispid; the leaves of most plants, both young and adult, are densely pubescent especially on the veins. Thus, these varieties actually cannot be separated and are not recognized here. Coccocypselum hirsutum Bartl. ex DC., Prodr. 4: 396. 1830.

556

R UBIACEAE

Fig. 443. Coccocypselum aureum

Coccocypselum brevipetiolatum Steyerm., Mem. New York Bot. Gard. 17(7): 304. 1967. Creeping perennial herb, with stems, petioles, and peduncles densely hirsute or hispid with tawny trichomes; leaf secondary veins 6 or 7 pairs; inflorescence capitate and subglobose, pedunculate, with 3–several flowers; hypanthium densely hirsute; calyx lobes linear, externally hirsute or hispid; fruits oblanceoloid, blue. Treeless moist savannas, borders of gallery forests along streams, 1200–1300 m; Bolívar (Km 167 south of El Dorado, Gran Sabana, region of Kavanayén, San Ignacio). Colombia, Guyana, Peru, Brazil, Bolivia. Coccocypselum huberi Steyerm., Ann. Missouri Bot. Gard. 74: 657. 1987. Creeping perennial herb, sparsely strigillose; stems slender, reddish; leaf secondary veins 5–7 pairs; inflorescence 1-flowered, pedunculate; hypanthium densely strigose; calyx lobes lanceolate, sparsely strigillose on both surfaces; fruits ellipsoid or obovoid, bright

Fig. 444. Coccocypselum guianense

blue. Upland forests, 1100–1200 m; Bolívar (Caco-tepui west of San Ignacio, northwest of San Ignacio de Yuruaní). Endemic. Coccocypselum lanceolatum (Ruiz & Pav.) Pers., Syn. Pl. 1: 132. 1805. —Condalia lanceolata Ruiz & Pav., Fl. Peruv. 1: 54. 1798. Coccocypselum canescens Willd. ex Schult. & Schult. f., Mantissa 3: 130. 1827. Perennial decumbent, spreading, or erectascending herb, villosulous; stems purplish or green; leaf secondary veins 7–12 pairs; inflorescence globose, pedunculate, with flowers 10–20; hypanthium villous; calyx lobes oblanceolate, obovate, or broadly elliptic and eventually squarrose or recurved, villous; fruits ellipsoid or obovoid, bright blue. Tepui slope forests, forests bordering savannas, 800–2300 m; Bolívar (Gran Sabana, Roraima-tepui, Sororopán-tepui, Uaipán-tepui), Amazonas (Simarawochi). Common elsewhere throughout Venezuela; Central America, Colombia, Guyana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina.

19. COFFEA L., Sp. Pl. 172. 1753. by Charlotte M. Taylor and Julian A. Steyermark Shrubs or small trees, unarmed, terrestrial, sometimes resinous. Leaves opposite or verticillate, sessile to petiolate, venation not lineolate, abaxially sometimes with foveolate domatia in the axils of the secondary veins; stipules persistent, interpetiolar or sometimes united around the stem, generally imbricate in bud, tri-

Coffea 557

angular, acute to acuminate or aristate. Inflorescences axillary, subcapitate to congested-cymose and several-flowered or rarely with flowers solitary, sessile to shortly pedunculate, bracteate with bracts united in pairs into a calyculus (i.e., a cupule). Flowers subsessile to pedicellate, medium to small, homostylous, protandrous, fragrant. Hypanthium turbinate to ellipsoid. Calyx limb reduced or truncate to lobed, the lobes 5, without calycophylls; corolla salverform or funnelform, white, internally glabrous or villous in the throat, the lobes 4–8, convolute in bud. Stamens 4–8, inserted in corolla throat; anthers narrowly oblong, dorsifixed near middle or near base, included or partially to fully exserted. Ovary 2-locular; ovules 1 in each locule, axile. Fruit subglobose to ellipsoid, drupaceous, carnose, red to black; pyrenes 2, chartaceous to coriaceous, 1-locular. Seeds rather large, subglobose to hemispherical. Neotropics (cultivated and persisting), Old World tropics; ca. 100 species, 2 in Venezuela, both in the flora area. Most species are native to Africa and Madagascar. Several species and hybrids are widely cultivated and sometimes locally persisting throughout the tropics, including in humid zones in the Neotropics. The roasted seeds (“beans”) of Coffea are of course the source of the beverage coffee, and an important world agricultural commodity. The taxonomy of Coffea in general and also the cultivated coffees is complex. Many of the cultivated races are hybrids that have been subjected to extensive selection. Bridson (1988) provides a very useful treatment of this group, including the cultivated lines. Coffea canephora Pierre ex A. Froehner is occasionally cultivated in the Neotropics and might be found in the flora area also. It is distinguished from the other cultivated coffees by its shiny leaves with long acuminate apices and cuneate to rounded bases and its corollas with 5 or occasionally 6 lobes. Key to the Species of Coffea 1.

1.

Leaves 8–15 × 2.5–9 cm, chartaceous when dry, acuminate to caudateacuminate and usually falcate at apex, abaxially with foveolate domatia 0.1–0.4 mm long; stipules acute to aristate; corolla lobes 5; fruits 10–16 × 10–14 mm ....................................................... C. arabica Leaves 12–30 × 3–13 cm, subcoriaceous when dry, acute, obtuse or abruptly and shortly acuminate at apex, abaxially with foveolate domatia 0.2–0.8 mm long; stipules obtuse; corolla lobes 6–8; fruits 15– 25 × 12–18 mm ............................................................................ C. liberica

Coffea arabica L., Sp. Pl. 172. 1753. —Arabica coffee, Café. Shrub to 3(–8) m tall; stems glabrescent; leaves 8–19.5 × 2.5–9 cm; stipules 3–12 mm long; inflorescences subcapitate; calyx limb 0.1–0.2 mm long; corolla tube 12–14 mm long, lobes 5, 14–20 mm long; fruit red to orange, 10–16 × 10–14 mm. Forests, gardens, 100–200 m; Bolívar (scattered localities), Amazonas (vicinity of San Carlos de Río Negro). Cultivated and locally persistent elsewhere in Venezuela, particularly in cooler zones and especially at middle elevations;

cultivated in similar areas in Mexico, Central America, the Antilles, Colombia, Ecuador, Peru, Bolivia, southern Brazil, and widely in other tropical regions. ◆Fig. 445. Coffea liberica Bull ex Hiern., Trans. Linn. Soc. London, Bot. 2, 1: 171, t. 24. 1876. —Café criollo, Café robusta. Tree to 8(–15) m tall; stems glabrous; leaves 12–30 × 3–13 cm; stipules 3–6 mm long; inflorescences subcapitate; calyx limb ca. 0.1 mm long; corolla with tube 10–14 mm long, lobes 6–8, 8–15 mm long; fruit red, 15–

558

R UBIACEAE

Fig. 445. Coffea arabica

25 × 12–18 mm. Forests, gardens, ca. 50 m; Delta Amacuro (Caño Araguaito near Isla Tórtola, Caño Joba-Suburu east of Caño Sacupana). Probably cultivated and locally persistent elsewhere in Venezuela, particularly in moist lowlands; cultivated in similar areas in Mexico, Central America, the Antilles, Colombia, Brazil, and perhaps elsewhere in the Neotropics, and widely in other tropical regions.

This coffee prospers at lower altitudes than Coffea arabica, although its beans are not as highly valued commercially for flavor as those of C. arabica.

20. CORDIERA A. Rich. in DC., Prodr. 4: 445. 1830. Gardeniola Cham., Linnaea 9: 247. 1834. Scepseothamnus Cham., Linnaea 9: 248. 1834. Garapatica H. Karst., Fl. Columb. 1: 57, t. 28. 1860. by Claes Persson, Piero G. Delprete, and Julian A. Steyermark Shrubs, treelets, trees, or rarely subshrubs or geofrutices (xylopodial low shrubs), glabrous or pubescent, dioecious, unarmed, terrestrial. Leaves opposite, papyraceous to subcoriaceous or rarely coriaceous, petiolate, venation brochidodromous, tertiary venation not lineolate; stipules interpetiolar and shortly connate intrapetiolarly at the base, persistent, ovate, triangular or transverse-oblong, acute, acuminate, rounded or truncate at apex, valvate in bud. Inflorescence terminal, bracteate, sessile or rarely subsessile, the staminate fasciculate or rarely trichotomously branched or with flowers solitary, the pistillate with flowers solitary or rarely to 5. Flowers small to medium-sized, sessile to shortly pedicellate, unisexual. Staminate flowers: hypanthium reduced; calyx limb cup-shaped, truncate or minutely dentate, internally equipped with colleters; corolla salverform, white or the tube sometimes greenish, internally glabrous, sparsely to densely puberulent, sericeous, or rarely canescent or hirsute, internally glabrous or pubescent, the tube

Cordiera 559

sometimes constricted below the lobes, lobes (3)4 or 5(–7), contorted in bud; stamens (3)4 or 5(–7), inserted in the lower to the upper portion of the corolla tube; anthers included or rarely the tips shortly exserted, narrowly oblong, dorsifixed near base or rarely in the middle, sessile or subsessile; pollen 3-colporate; pistillode present, slender. Pistillate flowers: hypanthium globose; calyx and corolla similar to the staminate except corolla usually shorter and equipped with 1 additional lobe; staminodes present, similar to anthers but shorter; ovary 2- or 3(–5)-locular; ovules 3–20 per locule, on axial placentas; styles with 2 or 3(–5) branches, abaxially with a revolute stigmatic area. Fruit baccate, globose, turbinate, or rarely slightly compressed, 2–many-locular, often with false septa, pulp fleshy, pericarp fleshy, usually black, sometimes yellow or reddish, or rarely beige-brown. Seeds 3–20, wedgeshaped or rarely sublenticular or ellipsoid, testa firm of elongated cells with secondary thickenings in the radial walls. Central America, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 24 species, 2 in Venezuela, both in the flora area. Cordiera is closely related to Alibertia, from which it differs in having generally smaller flowers, colporate pollen grains, and smaller fruits with a fleshy and juicy pericarp. Cordiera has been included in Alibertia by many previous authors, but the results of Persson (2000a) support its separation. Key to the Species of Cordiera 1.

1.

Corolla tube cylindrical or fusiform, not constricted below the lobes; stipules 0.8–2 mm long, united around the stem into a short tubular sheath or sometimes only shortly connate intrapetiolarly, truncate or rounded at apex ..................................................................... C. myrciifolia Corolla tube narrowly conical and constricted just below the lobes; stipules 1.3–6 mm long, deltoid to narrowly triangular and shortly connate intrapetiolarly, acute, acuminate, or aristate at apex ....... C. triflora

Cordiera myrciifolia (Spruce ex K. Schum.) C. Perss. & Delprete, comb. nov. —Alibertia myrciifolia Spruce ex K. Schum. in Mart., Fl. Bras. 6(6): 393. 1889. Alibertia uniflora Standl., Field Mus. Nat. Hist., Bot. Ser. 11: 179. 1936. Alibertia triloba Steyerm., Mem. New York Bot. Gard. 12: 227. 1965.

Fig. 446. Cordiera myrciifolia

560

R UBIACEAE

Cordiera myrciifolia var. tepuiensis Steyerm., Mem. New York Bot. Gard. 12: 226. 1965. Shrub 0.5–3 m or tree 1–8(–22) m tall; leaves 2.5–15.5 × 1.5–6.7 cm, acute to rounded or rarely cordate at base, acute to rounded at apex with the acumen deltoid to narrowly triangular, falciform, or oblong, 0.5–2.5 cm long. Evergreen lowland forests, shrublands, shrub savannas (caatingas}, gallery forests, savannas, 100–1300 m; Bolívar (widespread), Amazonas (near Puerto Ayacucho and Samariapo). Apure; Panama, Colombia, Guyana, Suriname, French Guiana, Brazil, Bolivia. ◆Fig. 446. Cordiera myrciifolia is a widespread species that is recognized by having densely puberulent corollas and relatively short stipules with a truncate or rounded apex, which are also united around the stem into a short tubular sheath or sometimes shortly connate intrapetiolarly.

Cordiera triflora A. Rich. in DC., Prodr. 4: 445. Sept. 1830. —Cordiera trifolia Steud., Nomencl. Bot. ed. 2, 1: 419. 1840, orth. var. —Alibertia triflora (A. Rich.) K. Schum. in Mart., Fl. Bras. 6(6): 392. 1889. Alibertia tenuifolia K. Krause, Verh. Bot. Vereins Prov. Brandenburg 50: 106. 1908. Alibertia steinbachii Standl., Publ. Field Columbian Mus., Bot. Ser. 4: 287. 1929. Alibertia benensis Standl., Publ. Field Columbian Mus., Bot. Ser. 7: 291. 1931. Shrub 0.8–3 m tall or tree 2–14 m tall; leaves 3.5–14 × 1.4–4.3 cm, acute to decurrent at base, the apex acute and commonly tapering into a deltoid to narrowly triangular acumen 0.8–1.3 cm long. Montane forests, 100–1900 m; Amazonas (Sierra de la Neblina). Guyana, Suriname, French Guiana, Amazonian Ecuador, Peru, Brazil, and Bolivia.

21. CORYPHOTHAMNUS Steyerm., Mem. New York Bot. Gard. 12(3): 264, fig. 36C–H. 1965. by Charlotte M. Taylor and Julian A. Steyermark Low pachycaulous shrubs, terrestrial, unarmed. Leaves 3- or 4-verticillate but often closely grouped at stem apices, subsessile, venation not lineolate; stipules interpetiolar, persistent, generally erect and appressed in bud, narrowly triangular, setose. Inflorescence terminal, pedunculate, multiflowered, capitate, bracteate with the external bracts involucral. Flowers small, sessile, floral biology unknown. Hypanthium turbinate. Calyx limb lobed, the lobes 3–5, without calycophylls; corolla funnelform, white sometimes flushed with purple, internally densely papillose-pilose in upper part of tube and on lobes, lobes 4, valvate in bud. Stamens 4, inserted in corolla throat; anthers elliptic, dorsifixed near middle, exserted. Ovary semi-inferior, 2-locular; ovules 6–8 in each locule, on axile placentas. Fruit capsular, superior, obovoid, woody, septicidal from the apex, the valves often later splitting from apex. Seeds small, ellipsoid to subglobose, rugulose. Venezuela (endemic to the flora area); 1 species. Coryphothamnus is similar to and apparently sympatric with Aphanocarpus and Pagameopsis. Coryphothamnus can generally be recognized by its hirsute-pilose pubescence, 4-lobed calyx and corolla, and terminal inflorescences, whereas the other two genera have sericeous to strigose pubescence, (except P. garryoides often has spreading-ciliate hairs on the leaf margins), 5- or 6-lobed calyx and corolla, and often axillary or pseudoaxillary inflorescences. Coryphothamnus auyantepuiensis (Steyerm.) Steyerm., Mem. New York Bot. Gard. 12(3): 264, fig. 36C–H. 1965. —Pagamea auyantepuiensis Steyerm., Fieldiana, Bot. 28: 584, fig. 129. 1953.

Shrub to 0.5 m tall; leaves 10–17 × 2.5–3 mm, pilose on midrib and margins at least when young; stipules 2.5–3 mm long, pilose when young, the sheath reduced, the setae 5–11; peduncles 16–23 cm long; heads ca. 1

Cosmibuena 561

cm diameter; calyx limb 2–4 mm long; corolla tube 2–4.5 mm long, lobes 3–3.5 mm long; capsules 3.5–4 × 2 mm. Open places, tepui summits, 1700–2300 m; Bolívar (Auyántepui). Endemic. ◆Fig. 447. The internodes and leaves of young plants of Coryphothamnus auyantepuiensis are more conspicuously hirsute-pilose than those of mature plants. Although Steyermark

Fig. 447. Coryphothamnus auyantepuiensis

(1974) described the capsule as 3.5–4 mm long, his accompanying figure depicts them as ca. 10 × 4 mm. This appears to be an error in the figure, which also depicts the anthers as ca. 5 mm long (versus 0.9 mm long on Holst 3004, MO), and depicts the calyx limb plus hypanthium as ca. 1.5 cm long (versus ca. 4.5 mm long on Holst 3004, MO).

Fig. 448. Cosmibuena grandiflora

562

R UBIACEAE

22. COSMIBUENA Ruiz & Pav., Fl. Peruv. 3: 2. 1802, nom. cons., non Cosmibuena Ruiz & Pav. 1794. Buena Pohl, Pl. Bras. Icon. Descr. 8, t. 8. 1827, nom. illeg., non Buena Cav. 1800. by Charlotte M. Taylor and Julian A. Steyermark Glabrous succulent trees or shrubs, usually epiphytic, unarmed. Leaves opposite, drying thickly coriaceous, venation usually not visible; stipules interpetiolar, oblanceolate to obovate, in bud erect and flatly appressed, caducous. Inflorescences terminal, pedunculate to subsessile, umbelliform or corymbiform and few-flowered or with flowers solitary, ebracteate or bracts reduced. Flowers pedicellate, large, showy, homostylous, protandrous, nocturnal, fragrant. Hypanthium ellipsoid to cylindrical. Calyx limb with short tube, the lobes (4)5 or 6(7), without calycophylls; corolla salverform with the tube well developed, white becoming yellowed with age, internally glabrous, the lobes (4)5 or 6(7), in bud convolute (C. valerii) or imbricate with 3 lobes external. Stamens 5 or 6, inserted just below the corolla mouth; anthers narrowly oblong, basifixed, included, sometimes shortly bicaudate at the base. Ovary 2-locular; ovules numerous in each locule, on axile placentas. Fruit capsular, oblong or cylindric, woody, septicidal from the apex. Seeds small, flattened, papery, winged, the wings entire to erose. Southern Mexico, Central America, Colombia, Venezuela, Guyana, Suriname, Ecuador, Peru, Brazil, Bolivia; 4 species, 1 in Venezuela. Cosmibuena is similar to Hillia; their distinctions are discussed under Hillia. Cosmibuena grandiflora (Ruiz & Pav.) Rusby, Bull. New York Bot. Gard. 4: 368. 1907. —Cinchona grandiflora Ruiz & Pav., Fl. Peruv. 2: 54, t. 198. 1799. Epiphytic or terrestrial tree 4–15 m tall; leaves 6.5–19.5 × 3.6–16 cm; stipules 8–30 mm long; corolla tube 5–9(–10) cm long, lobes elliptic-oblong, 10–40 mm long; capsules 40–

65 × 6–13 mm. Moist forested slopes, lowland riparian forests, 50–1300 m; Delta Amacuro (Curiapo, Río Amacuro), Bolívar (Altiplanicie de Nuria and eastern part of Cerro Venamo and Cerro Uroi west to Salto Pará on Río Caura). Barinas, Zulia; Central America, Colombia, Guyana, Suriname, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 448.

23. COUSSAREA Aubl., Hist. Pl. Guiane 98, pl. 38. 1775. by Charlotte M. Taylor and Julian A. Steyermark Trees or shrubs, unarmed, terrestrial, sometimes resinous. Leaves opposite or rarely ternate, sessile to petiolate, venation not lineolate, often with pilosulous and/ or foveolate domatia in the abaxial axils of the secondary veins; stipules usually caducous, generally triangular to broadly so, interpetiolar to united around the stem, and generally imbricate in bud or occasionally calyptrate (i.e., fused into a conical cap). Inflorescence terminal and uncommonly also axillary, sessile to pedunculate, multiflowered and subcapitate to congested-cymose, fasciculate, paniculate, or racemiform or with the flowers solitary, bracteate or the bracts sometimes reduced. Flowers sessile or pedicellate, medium-sized to large, fragrant, at least sometimes nocturnal, protandrous and homostylous or distylous or reportedly rarely unisexual on dioecious plants. Hypanthium subglobose to turbinate. Calyx limb truncate to denticulate or lobed, teeth or lobes 4, without calycophylls; corolla salverform to funnelform, white sometimes becoming yellow with age, internally usually glabrous, lobes 4, valvate in bud, often fleshy and triangular in cross section. Stamens

Coussarea 563

4, inserted in upper part of the corolla tube; anthers narrowly oblong, dorsifixed, included or partially exserted, subsessile or the filaments developed but short. Ovary incompletely 2-locular; ovules solitary in the ovary or 2 and fused, basal. Fruit ellipsoid to subglobose, baccate, coriaceous to spongy or carnose, white to yellow or infrequently orange, smooth, endocarp papery to chartaceous or infrequently woody. Seeds solitary, ellipsoid to subglobose, smooth. Mexico, Central America, Colombia, Venezuela, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina; ca. 100 species, 16 in Venezuela, 13 of these in the flora area. Coussarea is frequently overlooked, and its species confused with Faramea and Psychotria. Coussarea differs from Psychotria in its single-seeded fruits with the endocarp papery, versus drupaceous with 2–5 hard pyrenes in Psychotria. Coussarea can be separated from most species of Psychotria by its stipules that are not bilobed and its corollas with 4 lobes; most species of Psychotria have bilobed stipules and 5 corolla lobes. Faramea can be separated from Coussarea by its usually aristate stipules, often blue corollas, and its usually blue to black fruits. Key to the Species of Coussarea 1. 1.

2(1). 2. 3(2). 3. 4(3). 4. 5(2). 5. 6(5). 6. 7(1).

7.

8(7). 8.

Leaves on lower surface without domatia in the axils of the secondary veins ....................................................................................................... 2 Leaves on lower surface with pilosulous and/or foveolate domatia in the axils of the secondary veins (more than one leaf should be examined) ................................................................................................................ 7 Inflorescences umbelliform or fasciculate, with 1–several peduncles terminating in a single, usually unbranched group of flowers ................ 3 Inflorescences paniculate, with 1 peduncle and the flowering portion branched to several orders .................................................................... 5 Inflorescences with the flowers racemose, the pedicels borne from a developed (at least shortly) axis ......................................................... C. sp. A Inflorescences with the flowers capitate to fasciculate or umbellate, the flowers or pedicels borne directly from the top of the peduncle .......... 4 Calyx limb 2–3 mm long; corolla with tube 10–15 mm long, lobes 10– 15 mm long ............................................................................. C. brevicaulis Calyx limb 1.5–2 mm long; corolla with tube 4.5–5.5 mm long, lobes 3.5– 5 mm long ................................................................................... C. violacea Calyx limb ca. 1.5 mm long; peduncles 1–4 cm long ........................ C. ovalis Calyx limb 2–2.5 mm long; peduncles 4–7.5 cm long ............................... 6 Calyx limb ca. 2 mm long; stipules 5–8 mm long ........................... C. evoluta Calyx limb ca. 2.5 mm long; stipules 3–4 mm long ........................ C. liesneri Lower surface of leaves glabrescent to strigose or pilosulous at least on the veins; calyx, corolla, and fruit densely velutinous to pilosulous ................................................................................................... C. leptoloba Lower surface of leaves glabrous except sometimes pubescent in the axils of the secondary veins; calyx, corolla, and fruit puberulous to glabrous ................................................................................................................ 8 Calyx limb 4–5 mm long; corolla with tube 25–35 mm long, lobes 10–15 mm long; fruit 2.5–3 × 1.5–2 cm, with the endocarp woody ............ C. grandis Calyx limb 1–3 mm long; corolla with tube 5–6 mm long, lobes 5–6.5 mm long; fruit 1–2 × 1–1.5 cm, with the endocarp chartaceous ................. 9

564

R UBIACEAE

9(8). 9. 10(9). 10. 11(10). 11. 12(11). 12. 13(11). 13.

Inflorescence umbelliform to fasciculate, with 1–3 axes terminating in a group of flowers .......................................................................... C. violacea Inflorescences pedunculate and branched to several orders ................. 10 Calyx limb 2.5–3 mm long, with teeth 0.5–0.7 mm long ................. C. lasseri Calyx limb 1–2 mm long, truncate or with teeth to 0.3 mm long .......... 11 Inflorescences with peduncles 1–3.5 cm long and branched portion 3.5– 8 × 4–8 cm ............................................................................................ 12 Inflorescences with peduncles 0.5–2 cm long and branched portion 1.5– 4 × 2.5–4 cm ......................................................................................... 13 Corolla with tube 16–20 mm long and lobes ca. 8 mm long, these 1/3–1/2 as long as the tube ........................................................................ C. hirticalyx Corolla with tube 5–6 mm long and lobes ca. 6 mm long, about as long as the tube .................................................................................. C. paniculata Calyx, hypanthium, and corolla tube externally puberulous ............... ........................................................................................... C. leptophragma Calyx, hypanthium, and corolla tube externally glabrous ........... C. revoluta

Coussarea brevicaulis K. Krause, Bot. Jahrb. Syst. 40: 142. 1907, nom. nud.; Verh. Bot. Vereins Prov. Brandenburg 50: 117. 1908. Shrub to 4.5 m tall; leaves 21–23 × 4–9 cm; stipules 2–3 mm long; inflorescences terminal and sometimes also axillary; calyx limb 2–3 mm long; corolla tube 10–15 mm long, lobes 10–15 mm long; fruit 20–22 mm diameter. Evergreen lowland forests, 100– 200 m; Amazonas (near San Carlos de Río Negro). Amazonian Peru, Brazil, and Bolivia. Steyermark cited this species for the flora area, but no material of it has yet been seen by C.M. Taylor. Coussarea brevicaulis is similar to C. longiflora Müll. Arg. of the central and southwestern Amazon basin; C. longiflora has terminal inflorescences, flowers with pedicels 0.5–1.5 mm long, and funnelform calyx limb, versus subsessile and tubular in C. brevicaulis. Coussarea longiflora was reported from the region of San Carlos de Río Negro by H. Clark et al. (2000) but no material of this species has been seen for the preparation of this flora. Coussarea evoluta Steyerm., Ann. Missouri Bot. Gard. 75: 334, fig. 8. 1988. Tree to 8 m tall; leaves 18–29 × 5.5–13 cm; stipules 5–8 mm long; calyx limb ca. 2 mm long; corolla tube 12.5–13 mm long, lobes 7– 10 mm long; fruit unknown. Lower tepui slope forests, 200–400 m; Amazonas (base of Sierra de la Neblina). Endemic. Coussarea evoluta may not be entirely distinct from C. liesneri, but as yet neither of these species is well known.

Coussarea grandis Müll. Arg. in Mart., Fl. Bras. 6(5): 100. 1881. —Curello, Palo de pereze, Vanallo. Shrub or tree to 10 m tall; leaves 15–27 × 4.5–11.5 cm; stipules 2–3 mm long; calyx limb 4–5 mm long; corolla tube 25–35 mm long, lobes 10–15 mm long; fruit 2.5–3 × 1.5– 2 cm. Evergreen lowland forests, 100–200 m; Amazonas (Capibara, Maroa, San Carlos de Río Negro). Endemic. Coussarea hirticalyx Standl., Publ. Field Columbian Mus., Bot. Ser. 8: 175. 1930. Shrub to 2 m tall; leaves 12–30 × 7–22 cm; stipules 3–5 mm long; calyx limb 1–1.5 mm long; corolla tube 16–20 mm long, lobes ca. 8 mm long; fruit ca. 18 × 14 mm. Wet forests on granitic hills, 200–300 m; Amazonas (Río Cunucunuma). Colombia, Ecuador, Peru. The one collection seen from the flora area (A. Fernández 7544, MO, PORT) is here assigned to this species based on Steyermark’s species delimitations in Coussarea. This specimen is also similar to C. mapourioides Bremek. of the Guianas. Steyermark separated these only by their glabrous (C. mapourioides) versus strigose to pilose (C. hirticalyx) upper surfaces of the leaf midribs. However, the specimens treated by Steyermark as C. mapourioides have corolla tubes 15–20 mm long, while plants of C. hirticalyx from western South America have corolla tubes 33–34 mm long. Coussarea hirticalyx may actually vary notably in corolla tube length across South America, or the delimitation of these species may deserve re-evaluation.

Coussarea 565

Steyermark (1967) recognized two varieties of Coussarea hirticalyx: var. hirticalyx with hispid and/or strigose to strigillose pubescence on the pedicels and calyx, and var. glabrior Steyerm. with puberulous to sparsely hirtellous pubescence on these structures. These varieties were not recognized by Taylor (1999). Coussarea lasseri Steyerm., Mem. New York Bot. Gard. 17(1): 366. 1967. Small tree; leaves 11–14.5 × 5.5–7.5 cm; stipules ca. 3 mm long; calyx limb 2.5–3 mm long with teeth 0.3–0.7 mm long; corolla in bud with tube to 8 mm long, lobes to 6 mm long; fruit unknown. Gallery forests, ca. 800 m; Bolívar (Río Uairén in Río Kukenán basin). Endemic. Coussarea leptoloba (Benth. & Hook. f.) Müll. Arg., Flora 58: 468. 1875. —Faramea leptoloba Benth. & Hook. f., Gen. Pl. 2: 121. 1873. —Paripari. Shrub or tree to 5 m tall; leaves 13–23 × 3–8 cm; stipules 1.5–3 mm long; calyx limb with tube 5–6 mm long, lobes 2–3 mm long; corolla tube 12–20 mm long, lobes 8–17 mm long; fruit ca. 18 × 12 mm, orange. Evergreen lowland forests, 100–200 m; Amazonas (headwaters of Río Yatúa, San Carlos de Río Negro). Colombia, Guyana, French Guiana, Brazil. Steyermark (1967) recognized two varieties of this species, var. leptoloba with sessile or subsessile inflorescences, and var. mutisii (Standl.) Steyerm. with shortly pedunculate inflorescences. Var. mutisii is known only from the type, which was collected in an unknown location in Colombia. There are several additional species of Coussarea in Colombia that are similar to C. leptoloba (e.g., C. enneantha Standl.) that Steyermark (1967) did not consider in his study of this genus, so the identity of C. mutisii Standl. (and thus Steyermark’s variety) is considered unclear until it is compared with these other species as well. Coussarea leptophragma Müll. Arg., Flora 58: 466. 1875. —Cachicamo, Café, Café orillera. Tree to 7 m tall; leaves 8–20 × 3.5–8 cm; stipules 2–3 mm long; calyx limb 1.5–2 mm long; corolla tube ca. 5 mm long, lobes ca. 5 mm long; fruit 11–12 × 7–8 mm. Evergreen lowland forests, 100–200 m; Amazonas (Pie-

dra Cocuy, vicinity of San Carlos and Santa Barbara). Amazonian Brazil. ◆Fig. 451. Coussarea leptophragma may not be distinct from Coussarea paniculata, which is variable and widespread. It is maintained here provisionally pending more detailed study of Amazonian Coussarea. Coussarea liesneri Steyerm., Ann. Missouri Bot. Gard. 76: 967. 1989. Tree to 8 m tall; leaves 22–33 × 7–14 cm; stipules 3–4 mm long; calyx limb ca. 2.5 mm long; corolla in bud with tube to 6 mm long, lobes to 3 mm long; fruit unknown. Slope forests, ca. 600 m; Amazonas (Cerro Aracamuni). Northern Brazil. Coussarea ovalis Standl. Publ. Field Columbian Mus., Bot. Ser. 8: 367. 1931. —Oido de picure. Tree to 6 m tall; leaves 14–17 × 7–9.5 cm; stipules ca. 8 mm long; calyx limb ca. 1.5 mm long; corolla in bud with tube to 6 mm long, lobes to 4.5 mm long; fruit unknown. Evergreen lowland forests, 100–200 m; Amazonas (near mouth of Caño Marimajare south of San Carlos de Río Negro). Colombia, Ecuador, Peru. The specimens cited by Steyermark of this species from the flora area have not been seen by C.M. Taylor, and the application of Standley’s name in general and in Steyermark’s usage is not completely clear. Coussarea paniculata (Vahl) Standl., J. Wash. Acad. Sci. 18: 282. 1928. —Billardiera paniculata Vahl, Eclog. Amer. 1: 13, pl. 10, fig. 3. 1796 [1797]. —Naranjillo. Shrub or small tree to 8 m tall; leaves 15– 22 × 5.5–11 cm; stipules 2–5 mm long; calyx limb 1.5–2 mm long; corolla with tube 5–6 mm long, lobes 6–6.5 mm long; fruit 12–16 × 10–15 mm. Evergreen lowland to lower montane forests, 100–500 m; Bolívar (basins of Río Caroní, Río Caura, and Río Cuyuní), Amazonas (Río Cunucunuma). Anzoátegui, Barinas, Mérida, Sucre, Táchira; Panama, Trinidad, Guyana, Suriname, French Guiana, Amazonian Ecuador, Peru, and Bolivia, northern Brazil. ◆Fig. 450. Coussarea revoluta Steyerm., Mem. New York Bot. Gard. 17(1): 368. 1967. Shrub to 2.5 m tall; leaves 8.5–17 × 3.5–8 cm; stipules 2–3 mm long; calyx limb 1–2 mm

566

R UBIACEAE

long; corolla tube ca. 6 mm long, lobes ca. 6 mm long; fruit 13–18 × 7–9 mm. Riparian and evergreen lowland forests, 100–200 m; Amazonas (Río Cunucunuma, Río Ocamo, Río Padamo). Endemic. ◆Fig. 449. Coussarea revoluta may not be distinct from C. paniculata, which is variable and widespread. It is maintained here provisionally pending more detailed study of Amazonian Coussarea. Coussarea violacea Aubl., Hist. Pl. Guiane 98, pl. 38. 1775. Faramea schomburgkiana Benth., Linnaea 23: 450. 1850. —Coussarea schomburgkiana (Benth.) Benth. & Hook. f., Gen. Pl. 2: 121. 1873.

Fig. 449. Coussarea revoluta

Tree to 9 m tall; leaves 8–20 × 3.5–9 cm; stipules 2–3 mm long; calyx limb 1.5–2 mm long; corolla tube 4.5–5.5 mm long, lobes 3.5– 5 mm long; fruit ca. 22 × 11 mm. Evergreen lowland forests, 100–200 m; Amazonas (Río Casiquiare, upper Río Orinoco). Guyana, French Guiana. In French Guiana, Aublet reports that the fruits of this species are violet, and Boom and Delprete (2002, 616) report that they are bright red-orange. Either of these colors is quite unusual in Coussarea, although some species do have fruits reported to be pale to medium orange. However, collections of C. violacea from Venezuela (e.g., Stergios and Aymard 9167, MO) describe the mature fruits as white.

Coussarea 567

Fig. 450. Coussarea paniculata

Fig. 451. Coussarea leptophragma

Coussarea sp. A Tree to 7 m tall; leaves 15–28 × 5–10 cm; stipules 2–4 mm long; flowers unknown; fruit 18–20 × 14–16 mm. Periodically flooded forests, 100–200 m; Amazonas (base of Sierra de la Neblina near Río Mawarinima). Endemic. This species cannot be identified until its

flowers are found, and probably until Amazonian Coussarea is studied in more detail. Currently, it is based on Stein et al. 1712A (MO), W.W. Thomas 3282 (MO, NY), Liesner 17467 (MO, NY), Boom and Weitzman 5924 (MO), Liesner and Funk 15829 (MO, NY)

568

R UBIACEAE

24. COUTAREA Aubl., Hist. Pl. Guiane 314, pl. 122. 1775. by Charlotte M. Taylor and Julian A. Steyermark Trees or shrubs, terrestrial, unarmed. Leaves opposite, petiolate, venation not lineolate; stipules interpetiolar and sometimes shortly united intrapetiolarly, generally imbricate in bud, persistent, triangular, acute. Inflorescence terminal, cymose, bracteate, 3–15-flowered. Flowers pedicellate, large, showy, homostylous, protandrous. Hypanthium ellipsoid to turbinate. Calyx limb deeply lobed, lobes 5 or 6(7), linear or lanceolate-setaceous, without calycophylls; corolla funnelform and often somewhat zygomorphic, roseate, reddish lavender, or magenta marked with cream or suffused with white or pale green, internally glabrous, lobes 5 or 6(7), imbricate in bud. Stamens 6(7), inserted at the base of the corolla tube, zygomorphic due to the filaments curving to position the anthers together on the lower side of corolla; anthers narrowly oblong, dorsifixed near base, included or exserted. Ovary 2-locular; ovules numerous in each locule, on axile placentas. Fruit capsular, obovoid to ellipsoid, strongly flattened perpendicular to septum, chartaceous to woody, septicidal from the apex then loculicidal. Seeds small, flattened, elliptic, papery, broadly winged. Mexico, Central America, Lesser Antilles, Colombia, Venezuela, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Paraguay, Argentina; 3 species, 2 in Venezuela, 1 of these in the flora area. Similar large, funnelform, somewhat zygomorphic corollas are found in Platycarpum, Gleasonia, and Henriquezia. The distinctions among these are outlined under Henriquezia. The other Venezuelan species of Coutarea, C. alba Griseb., is distinguished by its corollas uniformly expanded in the tube and its fruits with numerous circular lenticels. Coutarea hexandra (Jacq.) K. Schum. in Mart., Fl. Bras. 6(6): 196. 1889. —Portlandia hexandra Jacq., Select. Stirp. Amer. Hist. 63, pl. 182, fig. 20. 1763. —Bonita de noche, Cabrito negro, Guatacare amarillo. Coutarea speciosa Aubl., Hist. Pl. Guiane 314, pl. 122. 1775. Coutarea campanilla DC., Prodr. 4: 350. 1830. —Coutarea hexandra var. campanilla (DC.) Steyerm., Mem. New York Bot. Gard. 23: 298. 1972. Coutarea pubescens Pohl in Mart., Fl. Bras. 2: 148, pl. 200. 1833. —Coutarea hexandra var. hexandra f. pubescens (Pohl) Steyerm., Mem. New York Bot. Gard. 23: 298. 1972. Coutarea hexandra var. pubescens K. Schum. in Mart., Fl. Bras. 6(6): 198. 1889. Coutarea lindeniana Baill., Adansonia 12: 300. 1879. Shrub or small tree to 6 m tall; leaves 2– 16 × 1.5–8 cm; stipules 1.5–2 mm long; inflo-

rescences 3–8 × 3–8 cm (not including the corollas); calyx limb 4–12 mm long; corolla with tubes 2–7 cm long, lobes 0.7–2 cm long; capsules 2–4 × 1.5–2.5 cm; seeds 1–1.5 × 0.5–0.8 cm. Deciduous to evergreen lowland forests and scrub, 50–400 m; Bolívar (Altiplanicie de Nuria, northeast of Ciudad Piar, between El Dorado and Km 88, El Manteco, Represa Guri, near Río Supamo, near Roraima-tepui), Amazonas (Caño Garcitas, south of Puerto Ayacucho). Widespread elsewhere in Venezuela; other distribution as in genus. ◆Fig. 452. Coutarea hexandra is a widespread, morphologically variable species. Several varieties and forms have been recognized by some previous authors, notably Steyermark (1972, 297–299), based on the pubescence of the vegetative parts of the plants [i.e., pilosulous or hirtellous in var. (or f.) pubescens], the size of the flowers (i.e., markedly smaller in var. campanilla), and the shapes of the leaves and flowers (i.e., markedly narrower in var. fluminensis K. Schum).

Declieuxia 569

Ochoterena (1994) reviewed this species in detail and noted that these and other distinctive characters are found in occasional plants from most parts of the range of this species, that these distinctive characters are extremes of continuous variation, that the various characters can be found in all combinations, and that the varieties as circumscribed morphologically lack geographic coherence. Consequently, these varieties are not recognized here.

Fig. 452. Coutarea hexandra

25. DECLIEUXIA Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 3: 352, pl. 281. 1818 [1819]. by Charlotte M. Taylor and Julian A. Steyermark Perennial herbs or subshrubs from a xylopodium, unarmed, terrestrial. Leaves opposite or verticillate, sessile or petiolate, venation not lineolate; stipules interpetiolar, generally valvate to imbricate in bud, subulate to setose, and persistent or sometimes obsolete. Inflorescence axillary and/or terminal, pedunculate, multiflowered, cymose with the axes often dichotomous or scorpioid, bracteate. Flowers sessile or subsessile, rather small, usually distylous, apparently diurnal. Hypanthium ellipsoid to flattened. Calyx limb lobed, lobes (2)4, generally equal, without calycophylls; corolla salverform, blue, purple, or white, internally sericeous in upper part of tube, lobes 4, valvate in bud. Stamens 4, inserted in upper part of tube; anthers narrowly oblong, dorsifixed below middle, included or exserted. Ovary 2-locular; ovules 1 per locule, basal. Fruit schizocarpous, didymous, leathery, strongly compressed laterally; mericarps 2, orbicular, indehiscent. Seeds small, compressed, lenticular. Mexico, Central America, Cuba, Colombia, Venezuela, Guyana, Suriname, Brazil, Bolivia, Paraguay; 27 species, 2 in Venezuela, both in the flora area.

570

R UBIACEAE

Key to the Species of Declieuxia 1. 1.

Leaves opposite or usually verticillate; corolla 4–6 mm long; fruit usually glabrous ..................................................................................... D. fruticosa Leaves opposite; corolla 2.5–3 mm long; fruit papillate-puberulent ................................................................................................. D. tenuiflora

Declieuxia fruticosa (Willd. ex Roem. & Schult.) Kuntze, Revis. Gen. Pl. 1: 279. 1891. —Houstonia fruticosa Willd. ex Roem. & Schult., Syst. Veg. 3: 527. 1818. Declieuxia chiococcoides Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 3: 354. pl. 281. 1818 [1819]. Declieuxia mexicana DC., Prodr. 4: 479. 1830. —Declieuxia fruticosa var. mexicana (DC.) Standl., Field Mus. Nat. Hist., Bot. Ser. 12: 378. 1936. Declieuxia chiococcoides var. guyanensis Müll. Arg. in Mart., Fl. Bras. 6(5): 442. 1881. —Declieuxia fruticosa var. guyanensis (Müll. Arg.) Standl., Publ. Field Columbian Mus., Bot. Ser. 8(5): 369. 1931. Declieuxia alfredii Ernst, Revista Cient. Mens. Univ. Central Ven. 1, no. 7: 222. 1887, “alfredi.” Subshrub to 0.7(–1.5) m tall; leaves (2)3(4) per node, 1–8 × 0.3–5 cm; stipules 3–8 mm long; inflorescences 1–6 × 1–6 cm; calyx lobes 0.1–1 mm long; corollas white or blue, tubes 1–7 mm long, lobes 1–3 mm long; mericarps 1–3 mm diameter. Savannas, open rocky places, sandstone or igneous outcrops, 50– 1300 m; Bolívar (Altiplanicie de Nuria, Auyán-tepui, Cerro Toribio, Gran Sabana, Río Paragua), Amazonas (Puerto Ayacucho and surrounding area, Sierra Parima). Aragua, Carabobo, Distrito Federal, Lara, Mérida, Miranda, Monagas, Sucre, Yaracuy, Trujillo, Zulia; other distribution as in the genus. ◆Fig. 453. Declieuxia fruticosa is a markedly variable species in leaf size, shape, and pubescence. Kirkbride (1976) revised the genus and reduced numerous varieties and species to synonymy under D. fruticosa. His treatment has been followed here. Declieuxia tenuiflora (Willd. ex Roem. & Schult.) Steyerm. & J.H. Kirkbr., Mem. New York Bot. Gard. 23: 399. 1972. —Tournefortia tenuiflora Willd. ex Roem. & Schult., Syst. Veg. 4: 540. 1819.

Knoxia brasiliensis Spreng., Syst. Veg. 1: 406. 1825. —Declieuxia brasiliensis (Spreng.) Müll. Arg., Flora 59(28): 434. 1876. Subshrub to 0.7 m tall; leaves paired, 1–7 × 0.5–2.5 cm; stipules 1–5 mm long; inflorescences 2.5–15 × 0.5–20 cm; calyx lobes ca. 0.5 mm long; corollas white or blue, tube 1.5–2.5 mm long, lobes 0.5–1 mm long; mericarps 1– 2 mm diameter. Deciduous to evergreen forests, borders of forest clearings, igneous outcrops, lowland savanna borders, 100–600 m; Bolívar (Cerro Altamira, Río Caura, Río Suapure, Serranía Los Pijiguaos), Amazonas (Caño San Miguel, Río Cuao, Río Ventuari, Río Yatúa). Sucre; northern to southeastern Brazil.

Fig. 453. Declieuxia fruticosa

Dendrosipanea 571

26. DENDROSIPANEA Ducke, Arch. Inst. Biol. Veg. Rio de Janeiro 2: 69. 1935. by Piero G. Delprete and Julian A. Steyermark Shrubs or small trees, terrestrial, unarmed. Leaves opposite, petiolate to subsessile, venation not lineolate; stipules interpetiolar, triangular, acute to acuminate or bifid, mainly persistent, generally imbricate to valvate in bud. Inflorescence terminal and in the uppermost leaf axils, subsessile to pedunculate, multiflowered, a trichotomously branched cincinnate cyme, bracteate with the bracts reduced. Flowers medium-sized, sessile to pedicellate, homostylous, protandrous. Hypanthium turbinate. Calyx limb lobed, lobes 5, equal or with one enlarged but without calycophylls; corolla salverform, white, internally villous in tube except glabrous at its base and densely yellow-barbate in its mouth, lobes 5, contorted in bud. Stamens 5, inserted near middle of the corolla tube; anthers dorsifixed, linear-oblong, apiculate, sub-sagittate, included; filaments short. Ovary 2-locular; ovules numerous in each locule, axile; style glabrous. Fruit capsular, ellipsoid, subligneous to chartaceous, septicidal from the apex. Seeds small (1–2 mm long), irregularly polyhedric, angled, finely scrobiculate. Southwest Venezuela, northern Brazil; 2 species, both in the flora area. Key to the Species of Dendrosipanea 1.

1.

Corolla tube 9–13 mm long; corolla lobes 7–11 mm long; leaf blades mostly rounded or broadly obtuse at apex, oblong- to oblanceolatespatulate, mainly broadest above the middle, generally revolute-margined, with the secondary veins and midrib of upper surface markedly sulcate ........................................................................................ D. revoluta Corolla tube 15–20 mm long; corolla lobes 12–16 mm long; leaf blades acute to acuminate at apex, elliptic-lanceolate, oblong-lanceolate, or lance-ovate, broadest at the middle, scarcely or not at all revolute-margined, with the secondary veins and midrib of upper surface not sulcate ....................................................................................... D. spigelioides

Dendrosipanea revoluta Steyerm., Mem. New York Bot. Gard. 10: 196, fig. 71. 1963. Shrub 0.5–4 m tall; leaves subsessile to short-petiolate, blades 3–10 × 1–3.5 cm; stipules ovate-triangular, 2–4 mm long; calyx lobes subequal to unequal, 2–17 mm long; corolla tube 9–13 mm long, the lobes 7–11 × 4– 5.5 mm; capsules (5-)7–10 × 3–5 mm. Whitesand savannas near rivers, 100–200 m; Amazonas (Río Atabapo, Río Pacimoni). Endemic. ◆Fig. 454. Dendrosipanea spigelioides Ducke, Arch. Inst. Biol. Veg. Rio de Janeiro 2: 70. 1935. Dendrosipanea wurdackii Steyerm., Mem. New York Bot. Gard. 10: 198, fig. 72A–G. 1963.

Shrub 2–4 m tall; leaves short-petiolate, blades 4.5–13 × 1–5 cm; stipules ovate-triangular, 2.5–3 mm long; calyx lobes subequal, 3–3.5 mm long; corolla tube 15–20 mm long, the lobes 12–16 × 4.5–7 mm; capsules 8– 10 × 4–5 mm. White-sand savannas, along streams, and adjacent seasonally flooded forests, usually in hummocks called “tatucos,” 100–200 m. Amazonas (above Pimichín, Río Guainía, Sabana El Venado, 15–20 m south of Tama Tama). Amazonian Brazil (Amazonas: Rio Curicuriari). Dendrosipanea wurdackii was separated from D. spigelioides based on the shape and size of the calyx lobes and details of the pubescence of both vegetative and reproductive structures, but the variation in these features can now be seen to be continuous.

572

R UBIACEAE

Fig. 454. Dendrosipanea revoluta

Fig. 455. Didymochlamys connellii

27. DIDYMOCHLAMYS Hook. f. in Hook., Icon. Pl. pl. 1122. 1872. by Charlotte M. Taylor and Julian A. Steyermark Small, rather fleshy herbs, epiphytic, unarmed. Leaves opposite but markedly anisophyllous and strongly distichous, often appearing alternate, shortly petiolate, venation not lineolate; stipules interpetiolar, triangular, generally imbricate to valvate in bud, caducous. Inflorescence terminal, pedunculate, few-flowered, shortly fasciculate, enclosed by 2 complanate ovate bracts. Flowers shortly pedicellate to subsessile, rather small, apparently homostylous and protandrous. Hypanthium turbinate. Calyx limb deeply divided, lobes 5, equal; corolla funnelform, white to blue, internally glabrous or with a ring of trichomes near the base, lobes 5, valvate in bud, the margins sometimes crisped to lobulate or incised. Stamens 5, equal or 2 of them larger, inserted near base of the corolla tube; anthers dorsifixed near the base, narrowly oblong, included. Ovary 2-locular; ovules numerous in each locule, borne on a central basal placenta. Fruit cupuliform, a leathery berry or capsule opening at the apex, apparently green. Seeds small, flattened, ovate, thinly sericeous and with a tuft of long trichomes attached at one end. Central America, Colombia, Venezuela, Guyana; 2 species, 1 in Venezuela. Didymochlamys is sometimes not recognized in the field as a Rubiaceae, because the marked anisophylly of the leaves often makes them appear alternate and the stipules are quickly caducous so the reduced leaf at each node is occasionally

Diodia 573

misinterpreted as a stipule situated opposite the leaf. Didymochlamys whitei Hook. f. is found in southern Central America and northwestern South America, so the two species of this genus are widely allopatric. Didymochlamys connellii N.E. Br., Trans. Linn. Soc. London, Bot. 6: 34, pl. 5, figs. 1–9. 1901. Herb to 15 cm tall; larger leaves 20–40 × 4–10 mm, oblanceolate, reduced leaves 5–10 × 1–2 mm, sometimes caducous; stipules 0.5– 1 mm long; peduncles 1–3 mm long; involu-

cral bracts 1–1.5 × 0.5–1 cm; calyx lobes 2– 2.5 mm long; corolla tube 12–14 mm long, the lobes 3–4.5 mm long, subentire; fruits not seen. Epiphytic and on boulders in damp evergreen forests, 800–1100 m; Bolívar (slopes of Roraima-tepui). Guyana. ◆Fig. 455.

28. DIODIA Gronov. ex L., Sp. Pl. 104. 1753. Hemidiodia K. Schum. in Mart., Fl. Bras. 6(6): 29. 1888. by Charlotte M. Taylor and Julian A. Steyermark Annual or perennial herbs or low shrubs, terrestrial, unarmed, often weedy. Leaves opposite, sometimes developed from an axillary buds and apparently verticillate, subsessile to petiolate, venation not lineolate; stipules interpetiolar and fused to petiole bases, generally erect and appressed to valvate in bud, persistent, sheath truncate to rounded or triangular, setose. Inflorescences axillary and sometimes also terminal, glomerulate or capitate, few-flowered to multiflowered, sessile, bracteate with the bracts setose or chaffy, the flowering portion of the stems sometimes spiciform due to the reduction of the leaves subtending the inflorescences. Flowers small, homostylous, protandrous, sessile. Hypanthium turbinate to subglobose. Calyx limb deeply 2- or 4(–6)-lobed, lobes equal or sometimes unequal in pairs, without calycophylls, the sinuses sometimes setose or with small triangular projections; corolla salverform to funnelform or rotate, white, pink, or pale purple, internally glabrous or variously pubescent, lobes 4(–6), valvate in bud. Stamens 4, inserted in the corolla throat; anthers narrowly oblong, dorsifixed, exserted. Ovary 2-locular; ovules solitary in each locule, axile. Fruit schizocarpous, ellipsoid to obovoid, papery to cartilaginous, bony, or corky, often shortly stipitate; mericarps or cocci remaining connected or usually separating, indehiscent or occasionally later dehiscent along the adaxial face. Seeds small, flattened to hemispherical or ellipsoid. Warm-temperate eastern U.S.A., Mexico, Central America, West Indies, Colombia, Venezuela, Trinidad-Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina, also in Africa and adventive in Asia and the Pacific; ca. 50 species, 9 in Venezuela, all in the flora area. Diodia is similar to and often confused with several other genera of its tribe, in particular Borreria. These genera are separated by their mode of fruit dehiscence, in particular with the two segments dehiscent in Borreria but indehiscent in Diodia, or tardily so in D. ocymifolia. Because of the difficulty of identifying plants with flowers or immature fruits, the species of Diodia are all included in the key to Borreria as well. As with the other members of its tribe, Diodia has been circumscribed differently by various authors. The systematics of this group are currently controversial and probably rather far from stable resolution. Therefore, the generic limits used by Steyermark (1972; 1974) are followed here provisionally, for consistency and as a gateway to those significant works. A useful study of many of our species of this genus in this circumscription was done by Bruza (1982).

574

R UBIACEAE

Diodia has recently been treated with a much more restricted circumscription by Bacigalupo and Cabral (1999), who have studied this group in detail in central and southeastern South America. They consider Diodia a genus of 5 American species with indehiscent corky fruits; this group is represented in the flora area only by D. kuntzei. They include the remaining Diodia species variously in Borreria (in the flora area this group comprises D. hyssopifolia, D. multiflora, D. ocymifolia, and D. spicata), and Diodella Small (in the flora area this group comprises D. apiculata, D. teres, D. radula, and D. sarmentosa). In their ongoing studies of this and related genera, Bacigalupo and Cabral have been to a large extent sequentially subdividing the traditional genera, in many cases with very strong justification, and as their work continues further changes will likely be published. Key to the Species of Diodia 1.

1.

2(1).

2. 3(2). 3.

4(3). 4. 5(4). 5. 6(3). 6. 7(6). 7. 8(7).

Plants creeping, usually rooting at the nodes, growing in muddy sites especially where water is drying up; fruits rather corky, solitary in the leaf axils ...................................................................................... D. kuntzei Plants weak-stemmed to erect, not rooting at the nodes, growing in various habitats, from moist to dry; fruits cartilaginous to bony or chartaceous, 1 or usually several in each leaf axil ............................... 2 Leaves dimorphic, those at flowering nodes reduced, half or less the size of the vegetative leaves, giving the flowering apex of the stem a spiciform appearance; vegetative leaves 17–110 × 7–35 mm ............ D. spicata Vegetative and reproductive leaves similar, the flowering portion of the stem thus similar to the vegetative; leaves 10–90 × 1–23 mm ........... 3 Leaves elliptic, 5–23 mm wide, adaxially with the secondary veins visible and usually sulcate ................................................................................ 4 Leaves narrowly ligulate to narrowly lanceolate, narrowly oblong, narrowly elliptic, linear, or narrowly oblanceolate, 1–7 mm wide, adaxially with the secondary veins usually not evident or visible and plane, to sometimes impressed on the oldest leaves ....................................... 6 Calyx lobes ca. 0.5 mm long; fruits ellipsoid-oblong, 3.5–5 × 1.5–2 mm, hirtellous ................................................................................. D. ocymifolia Calyx lobes 1.2–5 mm long; fruits subglobose to ellipsoid, 3–5 × 2.5– 4 mm, hirtellous to glabrous ................................................................. 5 Calyx lobes 1.5–5 mm long, at least some of them more than 3 mm long; corolla tube 5–7 mm long ............................................................ D. radula Calyx lobes 1.2–2.5 mm long; corolla tube 1.5–2 mm long ..... D. sarmentosa Leaves ligulate to narrowly elliptic or oblanceolate, adaxially with the secondary veins sulcate .......................................................... D. multiflora Leaves narrowly elliptic, linear, triangular, or lanceolate, adaxially with the secondary veins plane ..................................................................... 7 Corolla tube 1.5–2.2 mm long; fruit ellipsoid-oblong, 1–2 mm diameter; stipule setae glabrous .......................................................... D. hyssopifolia Corolla tube 2.5–8 mm long; fruit obovoid to subglobose; stipule setae glabrous or pubescent ................................................................................. 8 Plants perennial, woody at the base, the roots usually not collected with the stems; calyx lobes equal or 2 of them notably longer; each segment of fruit with 3–5 well-developed ribs on the dorsal (abaxial) face; corolla tube 4–8 mm long ............................................................ D. apiculata

Diodia 575

8.

Plants annual, soft or sometimes woody at base, the roots usually collected with the stems; calyx lobes equal to subequal; each segment of fruit smooth or with 1 well-developed rib and sometimes 2 shorter ribs on the dorsal (abaxial) face; corolla tube 2.5–4.5 mm long ............ D. teres

Diodia apiculata (Willd. ex Roem. & Schult.) K. Schum., Bot. Jahrb. Syst. 10: 313. 1889. —Spermacoce apiculata Willd. ex Roem. & Schult., Syst. Veg. 3: 531. 1818. Spermacoce rigida Willd. ex Roem. & Schult., Syst. Veg. 3: 531. 1818, nom. illeg., not Spermacoce rigida Salisb. 1796. —Diodia rigida Cham. & Schltdl., Linnaea 3: 341. 1828. —Diodella rigida (Cham. & Schltdl.) Small, Fl. Miami 177. 1913. Spermacoce rigida Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 3: 342. 1818 [1819], nom. illeg., not Spermacoce rigida Salisb. 1796. Diodia setigera DC., Prodr. 4: 563. 1830. Herb or subshrub to 0.5 m tall, hirtellous or pilose to glabrescent; leaves 10–30 × 2–7 mm, cuneate to rounded at base; stipule sheath 1.5–3 mm long, setae 7–8, 3–10 mm long; flowers several per axil; calyx lobes 4, 0.5–2.5 mm long; corolla pale to deep pink, tube 4–8 mm long, lobes 2–5 mm long; fruit obovoid, 2.5–4 × 1.5–2.5 mm. Savannas, open rocky places, along roadsides, 100–800 m; eastern Bolívar, Amazonas (Los Castillos de Guayana, Piacoa). Common in northern and central Venezuela; Mexico, Central America, West Indies, Colombia, Trinidad-Tobago, Guyana, Suriname, French Guiana, Brazil. ◆Fig. 456. Diodia apiculata is highly variable in leaf shape, size, and pubescence. The distinctions between it and D. teres are often subtle, but previous authors including in particular Bruza (1982) have considered them separate. The leaves often have cartilaginous margins that are generally as well developed as the midrib on the abaxial (lower) surface; when present this character helps distinguish D. apiculata and D. teres from other species of Diodia. Diodia hyssopifolia (Willd. ex Roem. & Schult.) Cham. & Schltdl., Linnaea 3: 350. 1828. —Spermacoce hyssopifolia Willd. ex Roem. & Schult., Syst. Veg. 3: 532. 1818. —Borreria hyssopifolia (Willd. ex Roem. & Schult.) Bacigalupo &

E.L. Cabral, Opera Bot. Belg. 7: 307. 1996. Spermacoce linearis Willd. ex Roem. & Schult., Syst. Veg. 3: 532. 1818. —Diodia hyssopifolia var. linearis (Willd. ex Roem. & Schult.) Steyerm., Mem. New York Bot. Gard. 23: 796. 1972. Diodia hyssopifolia var. hyssopifolia f. psiloclada Steyerm., Mem. New York Bot. Gard. 23: 795. 1972. Diodia hyssopifolia var. linearis f. glabriuscula Steyerm., Mem. New York Bot. Gard. 23: 796. 1972. Herb or subshrub to 1(–4) m tall, hirtellous to glabrescent; leaves 20–60 × 1–8 mm, acute to attenuate at base; stipule sheath 2– 3 mm long, setae 5–7, 0.5–4 mm long; flowers several per axil; calyx lobes 4, 0.2–1.5 mm long; corolla white, tube 1.5–2.2 mm long, lobes 1.5–2.5 mm long; fruit oblong-ellipsoid, 2–4.5 × 1–2 mm. Riparian and swampy habitats, sea level to 200 m; Delta Amacuro (Boca de Macareo, Caño Manamo), Bolívar (Río Aro, Río Caroní, Río Orinoco, Río Cuchivero, Río Cuyuní, Río Paragua), Amazonas (Isla Maricapana, Río Negro basin, Río Orinoco). Apure; Colombia, Guyana, Suriname, French Guiana, Brazil, Bolivia. ◆Fig. 457. Steyermark recognized two sympatric varieties and two sympatric forms of this species, based on partly overlapping ranges of leaf size and shape, pubescence pattern, calyx lobe length and shape, fruit size, and degree of development of the stem internodes. However, as discussed by Bruza (1982), this species is widely variable morphologically and these varieties and forms cannot be separated in a meaningful way. Diodia kuntzei K. Schum. in Mart., Fl. Bras. 6(6): 15. 1888. Glabrous herb; leaves ligulate, 10–30 × 2– 6 mm, truncate to cordulate at base; stipule sheath 2–3 mm long, setae 5–6, 2–4.5 mm long; flowers 1 or 2 per axil; calyx lobes 2–4, 2–3 mm long; corolla white, tube 9–14 mm long, lobes 3–4.5 mm long; fruit 4–5.5 × 3–4 mm. Wet places in savannas, ca. 100 m; Bolívar (southwest of Caicara del Orinoco).

576

R UBIACEAE

Guárico; Brazil, Bolivia, Paraguay, Argentina. Diodia multiflora DC., Prodr. 4: 564. 1830. —Borreria multiflora (DC.) Bacigalupo & E.L. Cabral, Opera Bot. Belg 7: 307. 1996. Herb or subshrub to 0.5 m tall, scabrous and hirtellous to glabrescent; leaves 12–62 × 5–20 mm, obtuse to rounded at base; stipule sheath 1.5–2 mm long, setae 5–7, 1–5 mm long; flowers several per axil; calyx lobes 4, 1–1.5 mm long; corolla white, tube 2–2.5 mm long, lobes 1.5–2 mm long; fruit subglobose to obovoid, ca. 3 × 2 mm. Infrequent in grasslands, near sea level to 100 m; Delta Amacuro (Clavellina). Anzoátegui, Guárico, Monagas; Colombia, Brazil, Bolivia, Paraguay. Diodia ocymifolia (Willd. ex Roem. & Schult.) Bremek., Recueil Trav. Bot. Néerl. 31: 303. 1934. —Spermacoce ocymifolia Willd. ex Roem. & Schult., Syst. Veg. 3: 530. 1819. —Hemidiodia ocymifolia (Willd. ex Roem. & Schult.) K. Schum. in Mart., Fl. Bras. 6(6): 30., pl. 72. 1888. —Borreria ocymifolia (Willd. ex Roem. & Schult.) Bacigalupo & E.L. Cabral, Opera Bot. Belg. 7: 307. 1996. Subshrub to 1.5 m tall, hirtellous or puberulous to glabrescent; leaves 30–90 × 5– 30 mm, acute at base; stipule sheath 3–5 mm long, setae 6–8, 1.5–5 mm long; flowers several per axil; calyx lobes 4, ca. 0.5 mm long; corolla white, tube 1.5–2 mm long, lobes 1–2 mm long; fruit ellipsoid-oblong, 3.5–4 × 1.5–2 mm. Wooded thickets and along streams, near sea level to 1300 m; common in Delta Amacuro (northeast of El Palmar), Bolívar (El Foco near Upata, base of Perai-tepui), Amazonas (Río Mavaca, upper Río Orinoco). Widespread though scattered elsewhere in Venezuela; Mexico, Central America, Antilles, Colombia, Trinidad-Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. The name of this species has been frequently misspelled as “ocimifolia,” including by Steyermark (1974). As evident from its nomenclature, the generic placement of this species has been controversial, due to varied interpretations of the mode of dehiscence of its fruit. Diodia radula (Willd. & Hoffmanns. ex Roem. & Schult.) Cham. & Schltdl.,

Linnaea 3: 342. 1828. —Spermacoce radula Willd. & Hoffmanns. ex Roem. & Schult., Syst. Veg. 3: 531. 1818. Venezuela and eastern to southeastern Brazil; 2 rather widely disjunct subspecies, 1 in Venezuela. Both Steyermark (1974) and Bruza (1982) considered the Venezuelan plants conspecific with the Brazilian ones, though these are notably disjunct geographically. D. radula subsp. venezuelensis Steyerm., Acta Bot. Venez. 6: 181. 1971. Subshrub to 0.8 m tall, pilosulous or pilose to glabrescent; leaves 20–55 × 10–18 mm, acute at base; stipule sheath 2–3 mm long, setae 6–12, 1.5–7 mm long; flowers several per axil; calyx lobes 4, 1.5–5 mm long, usually markedly unequal; corolla white often flushed with pink, tube 5–7 mm long, lobes 2.5–4 mm long; fruit subglobose to ellipsoid, 3–5 × 3–4 mm. Dry wooded slopes, 200–500 m; Bolívar (northeast of Icabarú, Cerro Los Matachines south of Santa Rosalina). Aragua, Falcón, Miranda, Sucre, Trujillo. Diodia sarmentosa Sw., Prodr. 30. 1788. Subshrub, usually clambering to 1–4 m high on other vegetation, scabrous and hirtellous to glabrescent; leaves 25–65 × 5–25 mm, acute at base; stipule sheath 1.5–2 mm long, setae 7–8, 3–6 mm long; flowers several per axil; calyx lobes 2 or 4, 1.2–2.5 mm long; corolla white, tube 1.5–2 mm long, lobes 1– 1.5 mm long; fruit subglobose to ellipsoid, 3.5–4 × 2.5–3 mm. Wet thickets, along streams in wet places, moist rocks, near sea level to 800 m; Delta Amacuro (Caño Araguabisi, Serranía de Imataca), Bolívar (Río Paramichí), Amazonas (Sierra Parima). Anzoátegui, Aragua, Distrito Federal, Guárico, Miranda, Sucre; Mexico, Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Brazil, also in tropical Africa, the Malayan Peninsula, and the Pacific islands. Diodia spicata Miq., Stirp. Surinam. Select. 179. pl. 52. 1850 [1851]. —Borreria spicata (Miq.) Bacigalupo & E.L. Cabral, Opera Bot. Belg. 7: 307. 1996. Stiff herb or subshrub to 1.5 m tall, scaberulous and/or puberulous to glabrescent; vegetative leaves 17–110 × 7–35 mm, acute at base; stipule sheath 3–4 mm long,

Diodia 577

setae 7–8 mm, 3–8 mm long; flowers several per axil; calyx lobes 4, ca. 0.5 mm long; corolla rotate, white, ca. 1 mm long, lobed for ca. 1/2; fruit ca. 1 mm diameter. Wet low ground along streams, 200–400 m; Amazonas (uppermost Río Orinoco above Platanal). Guyana, Suriname, French Guiana, Brazil (Amazonas: Rio Cauaburi, Serra da Neblina). Diodia teres Walter, Fl. Carol. 87. 1788. —Diodella teres (Walter) Small, Fl. Lancaster Co. 271. 1913. Diodia prostrata Sw., Prodr. 30. 1788. —Diodia teres subsp. prostrata (Sw.) Steyerm., Acta Bot. Venez. 6: 185. 1971. Diodia teres var. angustata A. Gray, Syn. Fl. N. Amer. ed. 2, 1(2): 35. 1884. —Diodia teres subsp. angustata (A. Gray) Steyerm., Acta Bot. Venez. 6: 184. 1971. Diodia teres subsp. angustata var. angustata f. latior Steyerm., Acta Bot. Venez. 6: 184. 1971. Diodia teres subsp. prostrata var. prostrata f. latifolia Steyerm., Acta Bot. Venez. 6: 186. 1971. Diodia teres subsp. prostrata var. prostrata f. leiocarpa Steyerm., Acta Bot. Venez. 6: 187. 1971.

Fig. 456. Diodia apiculata

Annual herb or subshrub to 0.4 m tall, pilosulous, hirtellous, pilose, or hirsute to glabrescent; leaves 10–40 × 5–9 mm, acute to rounded at base; stipule sheath 1–3 mm long, setae 6–7, 2–8 mm long; flowers 1–several per axil; calyx lobes 4, 0.5–3 mm long; corolla pale to deep pink, tube 2.5–4.5 mm long, lobes 1.5–2.5 mm long; fruit obovoid, 2–5 × 2–2.5 mm. Dry open savannas, sandstone or igneous outcrops, disturbed usually sandy ground, 50–300 m; Bolívar (east of La Paragua, Maripa-Aripao), Amazonas (area of Puerto Ayacucho). Widespread in lowland Venezuela; eastern and southern U.S.A., Mexico, Central America, West Indies, Colombia, Trinidad-Tobago, Guyana, Ecuador, Brazil, Bolivia, and not seen but probably in Suriname, French Guiana, and Peru. Among the Venezuelan plants of this species Steyermark (1974) recognized two varieties and a number of subspecies based on variation in leaf shape and pubescence. However, this species is notably variable morphologically in local populations throughout its range. Bruza (1982) noted that all the infraspecific taxa that have been recognized are linked by many intermediate forms, and he did not formally recognize any taxa within Diodia teres.

Fig. 457. Diodia hyssopifolia

578

R UBIACEAE

29. DUIDANIA Standl., Publ. Field Columbian Mus., Bot. Ser. 7: 408. 1931, emend. Steyerm., Mem. New York Bot. Gard. 17(7): 230, fig. 29A–F. 1967. by Charlotte M. Taylor and Julian A. Steyermark Shrubs or small trees, terrestrial, unarmed. Leaves opposite, subsessile to petiolate, venation not lineolate; stipules interpetiolar and often shortly united around the stem, persistent, generally imbricate to valvate in bud, triangular, aristate to 3-setose. Inflorescence terminal, subsessile, cymose-corymbose, manyflowered, bracteate. Flowers medium to small, subsessile, floral biology unknown. Hypanthium turbinate to hemispherical. Calyx limb deeply lobed, lobes 5 or 6, usually unequal, usually one or more flowers in each cyme with 1 or 2 lobes longer than the others, these longer lobes sometimes enlarged or expanded into purple petaloid calycophylls; corolla tubular, yellow, internally densely hirtellous in upper part of tube and on lobes, lobes 5, valvate in bud. Stamens 5, inserted in corolla throat; anthers narrowly oblong, dorsifixed, partially exserted. Ovary 2-locular; ovules 4 in each locule, on axile placentas. Fruit capsular, subglobose, woody, loculicidal from apex. Seeds small, orbicular, flattened, narrowly winged, with the margins erose. Venezuela; 1 species. Duidania is similar to and generally sympatric with Chalepophyllum and Maguireocharis. Their distinctions are discussed under Maguireocharis. Steyermark’s emendation of this genus primarily amplified the description of the gynoecium and fruits. Standley described the ovules as solitary in each locule, the capsules as septicidal, and the seeds as smooth, all of which were found by Steyermark to be incorrect when more material was available. Based on his corrections, Steyermark also noted that Standley’s original classification of Duidania in the tribe Chiococceae could no longer be supported, and that this genus seems most closely related to Hindsia Benth. ex Lindl. of Brazil. Hindsia itself is of controversial tribal position, but these genera seem likely to belong to the tribe Hedyotideae. Duidania montana Standl., Publ. Field Columbian Mus., Bot. Ser. 7: 409. 1931, emend. Steyerm., Mem. New York Bot. Gard. 17(7): 230, fig. 29A–F. 1967. Shrub or small tree to 3 m tall; leaves of variable size even on a single stem, 3–14 × 2.5–8.5 cm; stipules 4–7 mm long; inflorescences 3–12 × 3–12 cm; calyx lobes 5–16.5 mm long; corolla tube 10–11 mm long, lobes 1.5–2.5 mm long; capsules 7–8 × 8–9 mm. Tepui slope and summit scrub on low forests, 1200–2600 m; Amazonas (Cerro Duida, Cerro Huachamacari, Cerro Marahuaka). Endemic. ◆Fig. 458.

Fig. 458. Duidania montana

Duroia 579

30. DUROIA L. f., Suppl. 30, 209. 1782, nom. cons. by Charlotte M. Taylor and Julian A. Steyermark Trees or shrubs, unarmed, terrestrial, dioecious, often inhabited by ants; stems often myrmecophilous and swollen, often with some internodes markedly prolonged. Leaves opposite or 3–5-verticillate, sometimes swollen and myrmecophilous at base, sessile to petiolate, venation not lineolate; stipules calyptrate (i.e., united into a conical cap), densely sericeous to hirsute, caducous, circumscissile or sometimes splitting along one side and appearing spathaceous. Inflorescence terminal, bracteate, the staminate 3–many-flowered, subcapitate to fasciculate or cymose, the pistillate 1–several-flowered, capitate or similar to the staminate. Flowers medium-sized to large, unisexual, sessile to pedicellate. Staminate flowers: hypanthium reduced; calyx limb subtruncate to dentate, lobes 5–8, without calycophylls; corolla salverform, white to cream, externally densely sericeous to hirsute, internally glabrous or variously pubescent, lobes 5–8, convolute in bud; stamens 5–8, inserted in corolla tube; anthers subsessile, narrowly oblong, dorsifixed, included; pistillode present, similar to style and stigma. Pistillate flowers: hypanthium turbinate to hemispherical; calyx and corolla similar to staminate or sometimes larger, the corolla with up to 12 lobes; staminodes present, similar to anthers; ovary 1-locular; ovules numerous in each locule, on parietal placentas. Fruit baccate, subglobose, fleshy, yellow-green to brown. Seeds medium-sized, compressed, suborbicular, smooth, embedded in pulp. Central America, Colombia, Venezuela, Trinidad-Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; ca. 30 species, 14 in Venezuela, all in the flora area. Steyermark (1974) described the ovary of Duroia as “commonly 1-celled with parietal placentation; 5 or 6 placentas almost together in the center of the ovary.” Examination of specimens of Duroia shows the ovaries to be 1-locular with a central column apparently extending from bottom to top, and very possibly axile placentation. Duroia is similar to Amaioua, and these genera are frequently confused. They are distinguished primarily by the number of pistillate flowers and fruit; plants with staminate flowers are difficult to assign to genus. Duroia fusifera and D. geniopoides are the most commonly collected species in the flora area. Key to the Species of Duroia 1. 1. 2(1). 2. 3(2). 3. 4(2). 4.

Leaves sessile or subsessile, with inflated vesicles or pouches at their base ............................................................................................ D. saccifera Leaves petiolate, without inflated vesicles or pouches at their base ...... 2 Lower surface of the leaves and external surface of the calyx limb densely ferruginous-velutinous or ferruginous-sericeous ................................. 3 Lower surface of the leaves and external surface of the calyx limb glabrous, strigillose, strigose, pilosulous, or sericeous ............................. 4 Leaves 3- or 4-verticillate; petioles 3–7 cm long; staminate corollas with lobes ca. 13 mm long ................................................................. D. maguirei Leaves opposite; petioles 1–3 cm long; staminate corollas with lobes ca. 8 mm long ................................................................................... D. velutina Leaves opposite .......................................................................................... 5 Leaves 3- or 4-verticillate ........................................................................ 10

580

R UBIACEAE

5(4). 5.

6(5). 6. 7(6). 7. 8(7). 8. 9(8). 9. 10(4). 10. 11(10). 11. 12(11).

12.

13(10).

13.

14(13). 14. 15(14). 15.

Youngest stem internodes, pedicels, calyx limbs, and fruits hirsute to pilose with long, usually spreading trichomes .......................... D. eriopila Youngest stem internodes, pedicels, calyx limbs, and fruits glabrous to strigillose or strigillose with short to rather long, appressed trichomes ................................................................................................................ 6 Veins on abaxial leaf surface and youngest stem internodes strigillose to strigose, sometimes becoming glabrescent ................................D. strigosa Veins on abaxial leaf surface and youngest stem nodes glabrous ........... 7 Staminate corolla with tube glabrous internally, lobes 5 ..................... ........................................................................................ D. gransabanensis Staminate corolla with tube puberulous near base internally, lobes 6– 8 .............................................................................................................. 8 Leaves 2–4.5 cm wide ....................................................................... D. nitida Leaves 6–23 cm wide ................................................................................. 9 Staminate calyx limb 8–9 mm long, staminate corolla with tube 15– 20 mm long, lobes 7 or 8 ...................................................... D. bolivarensis Staminate calyx limb 4–4.5 mm long; staminate corolla with tube 10– 11 mm long, lobes 6 .................................................................D. paruensis Leaves pubescent, at least abaxially on principal veins ........................ 11 Leaves glabrous throughout .................................................................... 13 Pedicels of staminate flowers 20–30 mm long ................... D. kotchubaeoides Pedicels of staminate flowers 0–13 mm long .......................................... 12 Staminate calyx limb 5–8 mm long, externally hirsutulous to strigose sometimes becoming glabrescent; developing fruit with peduncle or stipe 6– 17 mm long, at maturity fruit stipitate to subsessile ............... D. fusifera Staminate calyx limb 4–5.5 mm long, externally puberulous to strigose, sometimes becoming glabrescent; developing fruit sessile or subsessile, at maturity fruit sessile to pedunculate or stipitate for up to 5 mm ................................................................................................ D. micrantha Staminate inflorescences with 3–5 fasciculate peduncles 10–50 mm long, each terminating in a cyme of flowers; pistillate flowers 3–5, fasciculate on peduncles 10–35 mm long ......................................... D. genipoides Staminate inflorescences congested-cymose to subcapitate, sessile to subsessile with peduncles to 4 mm long; pistillate flowers solitary, sessile or subsessile ............................................................................. 14 Staminate corollas with the lobes 5 and ca. 9 mm long ... D. gransabanensis Staminate corollas with the lobes 6 and at least 10–13 mm long ......... 15 Leaves elliptic to broadly so, 8.5–23 cm wide, cuneate to truncate, rounded, or cordate at base ...................................................... D. maguirei Leaves elliptic to obovate, 6–10 cm wide, acute to cuneate at base ............................................................................................... D. retrorsipila

Duroia bolivarensis Steyerm., Mem. New York Bot. Gard. 12(3): 206. 1965. Tree to 9 m tall; stems glabrous; leaves opposite, 13–31 × 6–20 cm, petiolate; staminate inflorescence subsessile, congested-cymose; pistillate flowers solitary, sessile; calyx limb externally glabrous, 8–9 mm long, truncate to denticulate; corolla tube 15–20 mm

long, lobes 7 or 8, 11–15 mm long; fruit glabrous, 5–7.5 × 3.5–7 cm. Montane or lower montane forests, 300–1600 m; Bolívar (Cerro Uroi, Macizo del Chimantá, Ptari-tepui). Endemic. Duroia eriopila L. f., Suppl. Pl. 209. 1781 [1782]. —Conserva.

Duroia 581

Tree to 20 m tall; stems hirsute; leaves opposite, 15–35 × 6–17 cm, petiolate; staminate inflorescence subcapitate to congested-cymose, subsessile; pistillate flowers solitary, sessile; calyx limb externally densely sericeous and hirsute, 3–6.5 mm long, truncate to denticulate; corolla tube 10–20 mm long, lobes 6, 8–18 mm long; fruit densely hirsute, 4–6 × 4–5 cm. Venezuela, Guyana, Suriname, French Guiana, Amazonian Brazil (Amazonas); 3 varieties, 1 in Venezuela. The other two varieties of Duroia eriopila recognized by Steyermark were reported from more restricted distributions: var. brevidentata Steyerm. was reported only from central Amazonian Brazil, and var. tafelbergensis Steyerm. from Suriname. D. eriopila var. eriopila Duroia eriopila var. eriopila f. glabra Steyerm., Mem. New York Bot. Gard. 12: 203. 1965. Evergreen forests, riparian forests, 50– 600 m; Delta Amacuro (Caño Güiniquina, northwest of Epaña), Bolívar (eastern), Amazonas (base of Cerro Duida, upper Río Orinoco). Anzoátegui; Guyana, Suriname, French Guiana, Amazonian Brazil (Amazonas). ◆Fig. 460. Steyermark’s f. glabra was described based on a specimen from Guyana with glabrous leaves; occasional specimens of D. eriopila from the Guianas do have sparsely pubescent to glabrous leaves, but the leaves intergrade with other plants in this respect and are recognizable as D. eriopila by the pilose to hirsute stems. Duroia fusifera Hook. f. ex K. Schum. in Mart., Fl. Bras. 6(6): 363. 1889. —Amaioua fusifera Spruce ex Benth. & Hook. f., Gen. Pl. 2: 82. 1873, nom. nud. —Carutillo, Carutillo rebalsero. Tree or shrub to 14 m tall; stems strigose to glabrescent; leaves 3- or 4-verticillate, 8– 15 × 3–7 cm, petiolate; staminate inflorescences congested-cymose or with 2 or 3 fasciculate peduncles each terminating in a cyme; pistillate flowers solitary (3), shortly pedunculate; calyx limb externally hirsutulous to strigose sometimes becoming glabrescent, 5–8 mm long, truncate to denticulate; corolla tube 8–13 mm long, lobes 5–6, 8.5–13 mm long; fruit glabrous, 3.5–5 × 3–5.4 cm. Evergreen lowland to lower montane forests,

flooded margins of streams, 50–500 m; Bolívar (Río Orinoco, Serranía Kako), Amazonas (basin of upper Río Guianía, Río Negro, and Río Orinoco). Apure; adjacent Colombia. Duroia genipoides Hook. f. ex K. Schum. in Mart., Fl. Bras. 6(6): 364. 1889. —Duroia genipoides Hook. f. in Benth. & Hook. f., Gen. Pl. 2: 82. 1873, nom. nud. —Amaioua genipoides Spruce ex K. Schum. in Mart., Fl. Bras. 6(6): 364. 1889, nom. nud. —Yayabare. Tree to 12 m tall; stems strigose to glabrescent, frequently with long myrmecophilous swellings; leaves 3- or 4-verticillate, 10– 20 × 4.5–9.5 cm, petiolate; staminate inflorescences with 3–5 fasciculate peduncles 1–5 cm long each terminating in a cyme; pistillate flowers 3–5, on peduncles 1–3.5 cm long; calyx limb externally sericeous to glabrescent, 5–10.5 mm long, truncate; corolla externally densely strigose, tube 8–10 mm long, lobes 6, 9–16 mm long; fruit glabrous, 3–4 × 2–2.5 cm. Evergreen forests, riparian forests, savanna borders, 100–400 m; Bolívar (upper Río Suapure, Sabana Cardona), Amazonas (Río Orinoco and tributaries). Guyana, Amazonian Brazil (Rio Castanho of Rio Padauari at Brazil-Venezuelan border). ◆Fig. 461. Duroia genipoides is similar in general aspect to D. fusifera, D. micrantha, and D. kotchubaeoides. As noted by Steyermark (1965), D. paraensis Ducke of Amazonian Brazil is similar to D. genipoides, and possibly these may not be distinct. Duroia gransabanensis Steyerm., Mem. New York Bot. Gard. 12(3): 205. 1965. Tree to 8 m tall; stems glabrous; leaves opposite or 3-verticillate, 15–25 × 7.5–25 cm, petiolate; staminate inflorescences congested-cymose to subcapitate, subsessile; pistillate flowers solitary, sessile; calyx limb externally sericeous to glabrous, 6–7 mm long, truncate; corolla tube 10–15 mm long, lobes 5, ca. 9 mm long; fruit glabrous, 6–7.5 × 4–5.5 cm. Upland gallery forests, forest borders, 1000–1400 m; Bolívar (Gran Sabana). Endemic. Duroia kotchubaeoides Steyerm., Mem. New York Bot. Gard. 12(3): 201, fig. 30E– G. 1965. Tree to 10 m tall; stems pilosulous to gla-

582

R UBIACEAE

brescent; leaves 4-verticillate, 6.5–14 × 3–6.5 cm, petiolate; staminate inflorescences with 3–6 fasciculate peduncles, each terminating in a single flower; pistillate inflorescences unknown; calyx limb externally strigose to glabrescent, 5–6 mm long, truncate; corolla tube 8–10 mm long, lobes 6–8, 17–18 mm long; fruit not known. Evergreen forests, riparian forests, 100–200 m; Amazonas (Caño Tama Tama). Colombia (Vaupés). ◆Fig. 462. Duroia maguirei Steyerm., Mem. New York Bot. Gard. 23: 343. 1972. Tree to 10 m tall; stems densely pilosulous or velutinous; leaves 3- or 4-verticillate, 16– 28 × 8.5–23 cm, petiolate; staminate inflorescences congested-cymose to subcapitate, subsessile; pistillate flowers solitary, sessile; calyx limb externally densely sericeous to velutinous, 4–5 mm long, truncate; corolla tube 12–13 mm long, lobes 6, ca. 13 mm long; fruit densely pilosulous, 5–6.5 × 5–5.5 cm. Evergreen forests, 100–400 m; Venezuela, Brazil, Colombia; 2 varieties, both in the flora area. Steyermark (1972) distinguished Duroia maguirei by its rounded to cordate leaf bases, but with more collections now available this character can be seen to vary to obtuse or cuneate (e.g., Zent 186-42, MO). Key to the Varieties of D. maguirei 1. Lower leaf surfaces pilosulous to velutinous ............................ var. maguirei 1. Lower leaf surfaces glabrous ..................... ............................... var. patentinervia D. maguirei var. maguirei 100–400 m; Amazonas (Río Cuao, 3.5 km from Yavita along road to Maroa). Amazonian Colombia, Amazonian Brazil (base of Serra da Neblina). D. maguirei var. patentinervia Steyerm., Mem. New York Bot. Gard. 23: 345. 1972. 100–200 m; Amazonas (near Yavita). Amazonian Brazil. Duroia micrantha (Ladbr.) Zarucchi & J.H. Kirkbr., Ann. Missouri Bot. Gard. 77: 851. 1990. —Coupoui micrantha Ladbr., J. Bot. 58: 176. 1920. —Barka-makata-

purai (Arekuna), Carutillo, Carutillo rebalsero, Mermo, Ohwe wale ‘empa (Yanomami). Duroia sprucei Rusby, Descr. S. Amer. Pl. 133. 1920. Tree to 18 m tall; stems strigose to glabrescent; leaves 3- or 4-verticillate, 8–18 × 3.5–8.5 cm, petiolate; staminate inflorescences congested-cymose or with 2 or 3 fasciculate peduncules each terminating in a cyme; pistillate flowers solitary, subsessile; calyx limb externally puberulous to strigose sometimes becoming glabrescent, 4–5.5 mm long, subtruncate to denticulate; corolla tube 10–11.5 mm long, lobes 6 or 7, 6.5–10 mm long; fruit glabrous, 3.5–6 × 2.5–4 cm. Evergreen forests, riparian forests, 50–200 m; Delta Amacuro (Sacupana), Bolívar (Río Caroní, Río Caura, Río Cuchivero, Río Paragua), Amazonas (basins of Río Orinoco and Río Casiquiare at La Esmeralda and Río Pasiba). Anzoátegui, Apure, Guárico; Colombia, Peru, Brazil. The fruits of Duroia micrantha are edible. It is similar in aspect to D. genipoides and D. fusifera. As noted by Steyermark (1965, 205) D. micrantha and D. fusifera may not be distinct. Duroia nitida Steyerm., Mem. New York Bot. Gard. 12: 208. 1965. Tree to 12 m tall; stems glabrous; leaves opposite, 5.5–16 × 2–5 cm, petiolate; staminate inflorescences subcapitate to congestedcymose, subsessile; pistillate flowers solitary, sessile; calyx limb externally sericeous, ca. 5 mm long, truncate; corolla in bud with tube to 9 mm long, lobes 6, to 10 mm long; fruit glabrous, ca. 3.5 × 2.5 cm. Savannas, rock outcrops, ca. 100 m; Amazonas (Río Baría). Amazonian Brazil (Amazonas). Duroia paruensis Steyerm., Mem. New York Bot. Gard. 12(3): 206. 1965. Tree to 6 m tall; stems glabrous; leaves opposite, 20–26 × 10–16 cm, petiolate; staminate inflorescences congested-cymose to subcapitate, subsessile; pistillate flowers solitary, sessile; calyx limb externally sericeous, 4–4.5 mm long, truncate to denticulate; corolla tube 10–11 mm long, lobes 6, 10– 11.5 mm long; fruit glabrous, ca. 5 × 4.5 cm. Montane slope forests, ca. 500 m; Amazonas (Cerro Parú). Apure; Colombia (Vaupés).

Duroia 583

Duroia retrorsipila Steyerm., Mem. New York Bot. Gard. 12(3): 208. 1965. Tree to 20 m tall; stems glabrous; leaves 3–5-verticillate, 15–24 × 6–10 cm, petiolate; staminate inflorescences congested-cymose, subsessile; pistillate flowers not seen; calyx limb externally puberulous, 5–6 mm long, truncate; corolla tube 7–8 mm long, lobes 6, ca. 11 mm long; fruits unknown. Tepui slope forests, ca. 1100 m; Bolívar (near Cerro Venamo). Endemic. Duroia saccifera (Mart. ex Roem. & Schult.) Hook. f. ex K. Schum. in Mart., Fl. Bras. 6(6): 362, pl. 146, fig. 1. 1889. —Amaioua saccifera Mart. ex Roem. & Schult., Syst. Veg. 7: 91. 1829. —Palo de baba. Tree or shrub to 10 m tall; stems hirsute to glabrescent; leaves ternate, 15–35 × 7–17 cm, sessile, the base with 2 sacs 10–12 × 5–8 mm; staminate inflorescences cymose; pistil-

late flowers solitary, sessile; calyx limb externally hirsute, tube 10–20 mm long, 5–7lobed, lobes 1–4 mm long; corolla tube 8–20 mm long, lobes 6, 7–17 mm long; fruits hirsute, 5–6 × 4–5 cm. Evergreen lowland and riparian forests, 100–200 m; Amazonas (basin of upper Río Negro and Río Guainía, near San Carlos de Río Negro, Yavita to Maroa). Amazonian Colombia, Peru, and Brazil. ◆Fig. 463. Duroia strigosa Steyerm., Mem. New York Bot. Gard. 12(3): 204. 1965. Small tree; stems strigose to glabrescent; leaves opposite, 18–26 × 9–14 cm, petiolate; staminate inflorescences congested-cymose to subcapitate, subsessile; pistillate flowers unknown; calyx limb externally sericeous, 6– 8 mm long, truncate; corolla tube 15.5–17.5 mm long, lobes 6, 11.5–14 mm long; fruit unknown. Slope forests, ca. 400 m; Bolívar (Cerro Pitón in basin of Río Chicanán). Endemic.

Fig. 459. Duroia velutina

584

R UBIACEAE

Fig. 460. Duroia eriopila var. eriopila

Duroia 585

Fig. 461. Duroia genipoides

586

R UBIACEAE

Fig. 462. Duroia kotchubaeoides

Fig. 463. Duroia saccifera

Elaeagia 587

Duroia velutina Hook. f. ex K. Schum. in Mart., Fl. Bras. 6(6): 366. 1889. —Amaioua velutina Spruce ex Benth. & Hook., Gen. 2: 82. 1873, nom. nud. Small tree to 5 m tall; stems densely velutinous; leaves opposite, 9–21 × 5–11 cm, petiolate; staminate inflorescences subcapitate to congested-cymose, subsessile; pistillate flowers solitary, sessile; calyx limb externally

densely velutinous, 4–7 mm long, denticulate; corolla tube ca. 8 mm long, lobes 6, ca. 8 mm long; immature fruit densely velutinous, to 5 × 4 cm. Evergreen lowland forests, 100– 200 m; not yet known from the flora area but expected in Amazonas. Amazonian Colombia, Brazil (Brazilian-Venezuelan border on Isla Macará, near confluence of Río Paduairi, Río Negro). ◆Fig. 459.

31. ELAEAGIA Wedd., Monogr. Cinch. 94. 1849. by Charlotte M. Taylor and Julian A. Steyermark Trees, unarmed, terrestrial, often with copious resin on young growth and inflorescences. Leaves opposite, sessile to petiolate, venation not lineolate; stipules united around the stem into a tubular sheath, in bud usually fused completely into a cap, this fused structure flattened perpendicularly to the subtending leaves, later splitting into 2 intrapetiolar sections, these persistent or the apical portions sometimes caducous but the truncate intrapetiolar bases persisting. Inflorescence terminal and sometimes in uppermost leaf axils, pedunculate, multiflowered, paniculate, ebracteate or bracts reduced. Flowers subsessile to pedicellate, homostylous, protandrous, small to medium-sized, fragrant. Hypanthium turbinate to cupular or subglobose. Calyx limb truncate to denticulate or lobed, lobes 5, without calycophylls; corolla tubular to campanulate or rotate, white or cream to yellow, internally densely pubescent at stamen insertion, lobes 5, convolute or imbricate in bud. Stamens 5, inserted in upper part of corolla tube; anthers elliptic to narrowly oblong, dorsifixed, exserted; filaments villous on their basal portions. Ovary 2-locular; ovules numerous in each locule, on axile placentas. Fruits capsular, subglobose, woody, the apical portion (i.e., the part distal to the calyx limb) often elongating so the fruits appear semi-inferior, loculicidal and often opening only through the apical portion. Seeds small, angled. Central America, Cuba, Colombia, Venezuela, Guyana, Suriname, Ecuador, Peru, Brazil, Bolivia; ca. 15 species, at least 6 in Venezuela, 1 of these in the flora area. In fruit Elaegia is sometimes confused with Chimarrhis, but its distinctive stipules separate it. The resin of some Elaeagia species, notably E. utilis Wedd. and E. pastoense Mora of the Andes, is produced in rather large amounts at vegetative stem apices. In western Colombia this resin is collected and used as a lacquer for wood carvings. Elaeagia maguirei Standl., Bull. Torrey Bot. Club 75: 568. 1948. Tree to 20 m tall; leaves 9–23 × 3–13 cm; stipules 7–20 mm long, the apical part caducous leaving persistent, truncate, interpetiolar bases 2–4 mm long; inflorescence 10–20 × 10–18 cm; flowers subsessile; calyx limb ca. 1 mm long, subtruncate; corolla with tube ca. 2 mm long, lobes ca. 2 mm long; capsules ca. 2 × 2.5–3 mm. Tepui slope forests, 1100–1800 m. Venezuela, Guyana, Suriname, Brazil; 2 varieties, both in the flora area.

Key to the Varieties of E. maguirei 1. Lower leaf surfaces glabrous or nearly so; corolla 4–4.5 mm long, the tube 2–2.5 mm long; peduncle appressed-puberulous ............................... var. maguirei 1. Lower leaf surfaces markedly pubescent; corolla ca. 3.8 mm long, the tube ca. 1.8 mm long; peduncle densely hirtellous with spreading hairs ....... var. pubens E. maguirei var. maguirei Bolívar (Amaruay-tepui, Cerro Venamo,

588

R UBIACEAE

Macizo del Chimantá, Uaipán-tepui), Amazonas (Cerro Duida, Cerro Huachamacari, Cerro Parú, Cerro Yutajé). Suriname, Brazil. ◆Fig. 464. E. maguirei var. pubens Steyerm., Mem. New York Bot. Gard. 12(6): 190. 1965. Bolívar (Gran Sabana), Amazonas (Sierra Parima). Endemic.

Fig. 464. Elaeagia maguirei var. maguirei

32. EMMEORHIZA Pohl, Flora 8: 183. 1825, nom. nud.; Pohl ex Endl., Gen. Pl. 1: 565. 1838. Endlichera C. Presl, Symb. Bot. 1: 73, pl. 49. 1832, non Endlicheria Nees 1833. by Charlotte M. Taylor and Julian A. Steyermark Climbing or sprawling herbs, terrestrial, unarmed. Leaves opposite, subsessile to petiolate, venation not lineolate; stipules interpetiolar and fused to the petiole bases, persistent, generally erect and appressed to valvate in bud, the sheath truncate to broadly rounded, setose. Inflorescence terminal and in uppermost leaf axils, pedunculate, multiflowered, paniculate, bracteate. Flowers small, homostylous, protandrous, pedicellate in umbelliform cymules. Hypanthium obconic to turbinate. Calyx limb deeply lobed, lobes 4, without calycophylls; corolla rotate to shortly funnelform, white, internally villous, lobes 4, valvate in bud. Stamens 4, inserted in the corolla throat; anthers dorsifixed near base, ellipsoid, exserted. Ovary 2-locular;

Faramea 589

ovules solitary in each locule, axile. Fruit capsular, obconic-cylindrical, chartaceous to coriaceous, becoming prolonged above insertion of persistent calyx limb and thus appearing semi-inferior, septicidal from this prolonged apical portion. Seeds small, oblong, flattened, papery, falsely appearing winged due to the spongy placenta to which the seeds adhere. Colombia, Venezuela, Trinidad, Guyana, Ecuador, Peru, Brazil, Bolivia; 1 species. Emmeorhiza umbellata (Spreng.) K. Schum. in Mart., Fl. Bras. 6(6): 408. 1889. —Borreria umbellata Spreng., Neue Entdeck. Pflanzenk. 2: 144. 1821. Stems to several m long; leaves 2–10 × 1–3 cm; stipule sheath 4–8 mm long, setae ca. 5, 0.5–6 mm long and unequal; inflorescences 3.5–13 × 3–11 cm; pedicels 1–6 mm long; calyx lobes 0.5–2 mm long; corolla tube ca. 0.8 mm long, lobes 1–2 mm long; capsules 3–4.5 × 1–2 mm; seeds and placenta together 2–4 × 1 mm. Distribution as in genus; 2 subspecies and 4 varieties, 1 subspecies and 2 varieties in Venezuela, 1 in the flora area. Steyermark distinguished var. umbellata of the southern and southeastern parts of the range of Emmeorhiza umbellata by its capsules 2–3 mm long and calyx lobes 0.3–1 mm long. He separated the plants of northeastern South America as subsp. septentrionalis. With more collections now available, these subspecies or varieties need review, and will likely be shown to represent end points of continuous variation rather than distinct taxa. Steyermark distinguished two varieties within subsp. septentrionalis based on the degree and distribution of pubescence, with var. pubens restricted to the Coastal Cordillera of Venezuela and sympatric with var. septentrionalis. E. umbellata subsp. septentrionalis Steyerm., Mem. New York Bot. Gard. 23: 785. 1972.

Tepui slope and summits, (700–)1300– 2300 m; Bolívar (Auyán-tepui, Cerro Bolívar, Ilú-tepui, Macizo del Chimantá). Venezuelan Andes and Coastal Cordillera; Colombia, Trinidad, Guyana. ◆Fig. 465.

Fig. 465. Emmeorhiza umbellata subsp. septentrionalis

33. FARAMEA Aubl., Hist. Pl. Guiane 102, pl. 40. 1775. by Charlotte M. Taylor and Julian A. Steyermark Trees or shrubs, terrestrial, unarmed, usually glabrous; stems often with the internodes rather flattened. Leaves opposite, sessile or petiolate, often distichous, venation not lineolate, the secondary veins sometimes united in a well-developed submarginal vein, sometimes with foveolate domatia in their abaxial axils; stipules persistent or caducous, interpetiolar to united around the stem and generally valvate to imbricate in bud or occasionally calyptrate (i.e., united into a conical cap), generally triangular, usually aristate, the aristas frequently crossed in bud. Inflo-

590

R UBIACEAE

rescences terminal and/or sometimes axillary, sessile to pedunculate, ebracteate or sometimes bracteate, capitate to cymose, paniculate, or fasciculate and few-flowered to multiflowered or sometimes reduced to a solitary flower. Flowers sessile or pedicellate, small to medium-sized, fragrant, distylous, protandrous. Hypanthium turbinate to ellipsoid. Calyx limb truncate to denticulate or lobed, the lobes 4, without calycophylls; corolla salverform to funnelform, white to blue or violet, internally usually glabrous, lobes 4, valvate in bud. Stamens 4, inserted in lower to upper part of corolla tube; anthers narrowly oblong, dorsifixed, included, subsessile or with short filaments. Ovary 1-locular or incompletely 2-locular; ovules basal, solitary in the ovary or 2 and fused. Fruit baccate, subglobose to oblate (“depressed-globose”), sometimes laterally compressed, blue, black, or rarely red, coriaceous to carnose, endocarp papery to chartaceous. Seeds solitary, rather large, subglobose to oblate, smooth. Mexico, Central America, West Indies, Colombia, Venezuela, Trinidad-Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay; ca. 150 species, 27 in Venezuela, 21 of these in the flora area. Faramea is similar and often confused with Coussarea; their separation is discussed under Coussarea. Some species of Faramea with well-developed submarginal leaf veins are occasionally confused with Melastomataceae, but that family lacks stipules. Faramea can often be recognized by the crossed stipule aristas at the stem apex. Key to the Species of Faramea 1. 1. 2(1).

2.

3(2). 3. 4(3). 4. 5(4). 5. 6(1). 6. 7(6).

7.

Inflorescences all axillary .......................................................................... 2 Inflorescences terminal, or terminal and axillary .................................... 6 Inflorescences sessile to subsessile, with the flowers sessile to subsessile; leaves with the submarginal vein well developed, about as well marked as the midrib, nearly straight ..................................... F. maguirei Inflorescences pedunculate, with flowers sessile to pedicellate; leaves with the submarginal vein weakly to rather well developed, less well marked than the midrib, looping .......................................................... 3 Flowers subtended by a pair of narrowly lanceolate bracts, these 1–3 mm wide ............................................................................................. F. morilloi Flowers ebracteate, with bracts reduced, or with bracts 6–8 mm wide ................................................................................................................ 4 Flowers in cymes of 3–15 .............................................................. F. capillipes Flowers solitary in each axil ..................................................................... 5 Peduncles 20–45 mm long; flowers not subtended by bracts; calyx limb 4– 6 mm long ...................................................................................... F. egregia Peduncles 2–15 mm long; flowers subtended by a pair of bracts 13–23 × 6–8 mm; calyx limb ca. 2 mm long .......................................... F. yutajensis Leaves sessile or subsessile, cordulate at base .......................... F. sessilifolia Leaves petiolate, truncate to cuneate or acute at base ............................ 7 Leaves with secondary veins extending to unite with the margins, the margins cartilaginous, thickened, about as strongly developed at the midrib ..................................................................................................... 8 Leaves with secondary veins branching before reaching the margins, the margins papery to cartilaginous, not or a little thickened but not as well developed as the midrib ................................................................. 9

Faramea 591

8(7). 8. 9(7).

9.

10(9). 10. 11(10). 11. 12(11). 12. 13(12). 13. 14(9). 14. 15(14). 15. 16(14). 16. 17(16). 17. 18(17). 18. 19(17).

19.

Inflorescences ebracteate or the bracts reduced; leaves 10–18 cm long ....................................................................................................... F. boomii Inflorescences bracteate; leaves 6.5–9.5 cm long ...................... F. crassifolia Inflorescences with the flowers solitary, the flowers borne in 1 fascicle or umbelliform cyme, or the flowers borne on 3–19 peduncles that terminate in an umbelliform or fasciculate cyme, this cyme often enclosed by a pair of lanceolate to ovate bracts ..................................................... 10 Inflorescences with flowers several to numerous, borne in cymose or paniculate inflorescences branched to several orders, without bracts or bracts reduced ...................................................................................... 14 Flowers solitary............................................................................ F. yutajensis Flowers 2–numerous ................................................................................ 11 Calyx limb 2–3.5 mm long; corolla with tube 12–27 mm long, lobes 9– 10 mm long ................................................................................ F. cardonae Calyx limb 0.5–1.5 mm long; corolla with tube 3–10 mm long, lobes 2– 9.5 mm long .......................................................................................... 12 Corolla tube ca. 10 mm long; peduncles each with a pair of white petaloid bracts 15–25 × 7–15 mm ........................................................ F. anisocalyx Corolla tube 3–7.5 mm long; peduncles ebracteate or with a pair of green to white bracts 3–10 × 1–7 mm ........................................................... 13 Leaves 4.5–11.5 cm long; inflorescences with 3–7 peduncles; corolla with tube 3–3.5 mm long, lobes ca. 2 mm long ...................................... F. berryi Leaves 8.5–14 cm long; inflorescences with 7–19 peduncles; corolla with tube 6–7.5 mm long, lobes 6–9.5 mm long .......................... F. parvibractea Stipules rounded to obtuse and not aristate; peduncles jointed near base .............................................................................................................. 15 Stipules rounded to acute and with an arista 0.5–10 mm long; peduncles jointed near base or not ....................................................................... 16 Calyx limb ca. 3.5 mm long; leaves with tertiary venation plane and usually not visible; Sierra de la Neblina ........................................ F. neblinae Calyx limb 2–3 mm long; leaves with tertiary venation visible and usually prominulous; widespread ................................................... F. torquata Calyx limb 2–3.5 mm long, lobed for more than 1/ 2 its length .................................................................................................. F. yavitensis Calyx limb 0.2–3 mm long, truncate to lobed for up to 1/2 its length ..... 17 Stipules tubular, united around the stem for 1/2 or more of their length, generally persistent ............................................................................. 18 Stipules interpetiolar or only shortly united intrapetiolarly, for 1/4 or less of their length, persistent or caducous ............................................... 19 Leaves linear-lanceolate to lanceolate, 0.6–2.5 cm wide, 5.5–7 times as long as wide ........................................................................... F. angustifolia Leaves elliptic-oblong, oblong, or lance-oblong, 1.5–6 cm wide, 2.5– 5 times as long as wide ............................................................ F. multiflora Peduncles, inflorescence axes, and pedicels relatively slender and flexuous; corolla with lobes longer than tube; fruit 5–9 mm diameter ................................................................................................... F. capillipes Peduncles, inflorescence axes, and pedicels not particularly slender nor flexuous; corolla with tube equal to or shorter than lobes; fruit 5–13 × 7–15 mm ............................................................................................... 20

592

R UBIACEAE

20(19). Calyx limb ca. 0.2 mm long; corolla violet, with tube ca. 6 mm long ........................................................................................... F. tamberlikiana 20. Calyx limb 1.5–3 mm long; corolla cream to white, with tube 8–23 mm long ....................................................................................................... 21 21(20). Leaves 4–10 cm long, elliptic, chartaceous; inflorescence 3–4 cm wide, with 1 pair or 1 whorl of 4 secondary axes ........................... F. orinocensis 21. Leaves 8.5–25 cm long, narrowly elliptic, elliptic, lance-elliptic, or oblanceolate, papyraceous; inflorescences 4–10 cm wide, with 1–several pairs of secondary axes ........................................................................ 22 22(21). Leaves with tertiary venation usually prominulous on both surfaces; corolla with tube 8–12 mm long, lobes 6–8 mm long ............... F. torquata 22. Leaves with tertiary venation plane to a little raised on both surfaces; corolla with tube 15–23 mm long, lobes 10–25 mm long ................... 23 23(22). Corolla lobes 17–25 mm long ................................................... F. occidentalis 23. Corolla lobes 10–12 mm long .................................................... F. stenopetala Faramea angustifolia Spruce ex Müll. Arg., Flora 58: 474. 1875. Shrub or tree to 3 m tall; leaves 8–12 × 0.5–2.5 cm, petiolate; stipules generally persistent, tubular, 2–3 mm long, aristas 1–2 mm long; inflorescence terminal, cymose, pedunculate; calyx limb ca. 0.5 mm long, denticulate; corolla white to blue, tube ca. 4 mm long, lobes ca. 3 mm long; fruit oblate, 5–7 × 7–10 mm. Evergreen lowland forests, 100– 200 m; Amazonas (Río Cunucunuma, basin of Río Negro). Amazonian Colombia and Brazil. As noted by Taylor (1999, 298), this species is similar to Faramea multiflora. Faramea multiflora is morphologically variable, and occasional plants from throughout its range have relatively narrow leaves similar to those that distinguish F. angustifolia. Thus, F. angustifolia is here recognized provisionally but may eventually prove to be a synonym of F. multiflora. Faramea anisocalyx Poepp. & Endl., Nov. Gen. Sp. Pl. 3: 28. 1845. Faramea pulchella Müll. Arg., Flora 58: 475. 1875. —Faramea anisocalyx var. pulchella (Müll. Arg.) Steyerm., Mem. New York Bot. Gard. 17(1): 375. 1967. Treelet to 3 m tall; leaves 4–14 × 1–1.5 cm, petiolate; stipules caducous, interpetiolar, 1– 1.5 mm long, aristas 1–3 mm long; inflorescences terminal, with 3 fasciculate peduncles, each terminating in a flower or fascicle, this enclosed by a pair of bracts 15–25 × 7–15 mm; calyx limb ca. 1 mm long, dentate; corolla blue, tube ca. 10 mm long, lobes ca. 5 mm long; fruit subglobose to oblate, 5–6 mm

diameter. Lower montane forests, 500–600 m; Bolívar (Cerro Abismo, south of El Paují, Río Samay), Amazonas (upper Río Orinoco). Colombia, Ecuador, Peru, Brazil, Bolivia. The distinctive petaloid bracts of the inflorescence of Faramea anisocalyx are caducous after anthesis, so conclusive determination of this species in fruit is sometimes difficult. The leaves of Faramea anisocalyx are generally elliptic-oblong to oblanceolate, papyraceous, and matte or dull on the upper surface; those of the similar species F. berryi are generally elliptic to oblanceolate, papyraceous, and shiny on the upper surface; those of another similar species, F. parvibractea, are generally chartaceous to subcoriaceous, elliptic-oblong, and quite shiny on the upper surface. Steyermark recognized three varieties of Faramea anisocalyx based on leaf size, degree of visibility of the leaf venation, number of flowers per peduncle, and corolla size. This variation does appear to be partly clinal, but with more specimens now available his varieties cannot be clearly separated and are not recognized here. Steyermark reported F. corymbosa Aubl. from Lara and the Río Negro basin of the flora area, but the specimens from there he treated as F. corymbosa are here treated in, variously, F. larensis Steyerm., F. berryi, and F. anisocalyx. H. Clark et al. (2000) also reported F. corymbosa from the region of San Carlos de Río Negro; the source of their report is unknown. Faramea berryi Steyerm., Pittieria 9: 5. 1981.

Faramea 593

Tree to 15 m tall; leaves 4.5–11.5 × 1.5–4.5 cm, petiolate; stipules caducous, interpetiolar, 1–5 mm long, aristas 1–1.5 mm long; inflorescence terminal, with 3–7 fasciculate peduncles, each terminating in a flower or fascicle, this enclosed by a pair of bracts 3–5 × 1–2 mm long; calyx limb ca. 0.5 mm long, denticulate; corolla white, tube 3–3.5 mm long, lobe ca. 2 mm long; fruit oblate, ca. 4 × 6 mm. Evergreen lowland to lower montane forests, 100–800 m; Bolívar (Río Caura), Amazonas (San Carlos de Río Negro). Brazil, Colombia. Steyermark reported Faramea corymbosa Aubl. from the Río Negro basin of the flora area, but the specimens from the flora area he treated under this name are here identified as F. berryi and F. anisocalyx. Faramea berryi is similar to and easily confused with F. anisocalyx and F. parvibractea; comments regarding their separation are under F. anisocalyx. Faramea boomii Steyerm., Ann. Missouri Bot. Gard. 75: 335. 1988. Tree to 12 m tall; leaves 10–18 × 3–7 cm, petiolate; stipules generally persistent, shortly united around the stem, 2–2.5 mm long, aristas 1–3 mm long; inflorescence terminal, cymose, pedunculate; calyx limb ca. 1 mm long, subtruncate; corolla white, tube ca. 17 mm long, lobes 8–9 mm long; fruit subglobose, 15–16 mm diameter. Evergreen lowland forests, low forests on igneous outcrops, 100–200 m; Amazonas (slopes of Cerro Aracamuni, Río Cunucunuma, base of Sierra de la Neblina). Adjacent Brazil. Faramea boomii is similar to and perhaps not distinct from F. juruana K. Krause of the central and western Amazon basin. Plants of the Orinoco River and Río Negro basins are here treated in F. boomii, while those from Peru, Brazil, and French Guiana are included in F. juruana. These species appear to differ in corolla size; F. juruana has corollas with the tube ca. 12 mm long and the lobes ca. 6 m long. However, only one collection with mature flowers has been seen for each species (F. juruana, W. Thomas et al. 4747, MO; F. boomii, Boom & Weitzman 5278, MO), and the flowers are nocturnal and thus may vary widely in size as in other nocturnalflowered Rubiaceae. Faramea capillipes Müll. Arg., Flora 58: 470. 1875.

Shrub or tree to 6 m tall; leaves 4–15 × 1– 5.5 cm, petiolate; stipules usually persistent, interpetiolar, ca. 1 mm long, aristas 2–4 mm long; inflorescence terminal and/or axillary, cymose, pedunculate; calyx limb ca. 0.5 mm long, denticulate; corolla cream, tube 3–5 mm long, lobes 5–6 mm long; fruit subglobose to oblate, 5–9 mm diameter. Rain forests, evergreen seasonally dry forests, 100–700 m; western Bolívar (widespread), Amazonas (Cerro Yapacana, near Puerto Ayacucho, Yavita). Apure, Barinas, Mérida, Táchira, Zulia; Colombia, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 467. Faramea capillipes is quite variable in the size of the leaves and inflorescences. Faramea cardonae Standl. & Steyerm., Fieldiana, Bot. 28: 574, fig. 124. 1953, emend. Steyerm., Mem. New York Bot. Gard. 17: 377. 1967. Shrub or small tree to 7 m tall; leaves 7– 16 × 2–8 cm, petiolate; stipules apparently caducous, interpetiolar, 3–3.5 mm long, aristas 2.5–3.8 mm long; inflorescence terminal, with 5–8 fasciculate peduncles, each terminating in a flower or fascicle, this enclosed by a pair of bracts 7–13 mm long; calyx limb 2– 3.6 mm long, dentate; corolla white sometimes flushed with lavender, tube 12–27 mm long, lobes 9–10 mm long; fruit subglobose, 7–12 mm diameter. Tepui slope forests, 1000–1400 m; southeastern Bolívar (Cerro Guaiquinima, Cerro Venamo, La Escalera, Macizo del Chimantá, basin of Río Cuyuní west to Río Paragua). Endemic. ◆Fig. 468. Faramea crassifolia Benth., J. Bot. (Hooker) 3: 233. 1841. —Cafecito, Pata de grulla. Tree to 6 m tall; leaves 6.5–9.5 × 2.5–9.5 cm, petiolate; stipules caducous, interpetiolar, 1.5–2.5 mm long, aristas 1.5–4 mm long; inflorescence terminal, cymose, pedunculate; calyx limb 0.5–2 mm long, dentate; corolla white, tube ca. 5 mm long, lobes ca. 4.5 mm long; fruit oblate, 4–5 × 8 mm. Semideciduous forests, savanna borders, 100–300 m; northwestern Bolívar (base of Cerro El Tigre south of Caicara del Orinoco). Apure; Guyana, northern Brazil. Dried specimens of Faramea crassifolia often have a distinctive yellow cast. The thickly coriaceous leaves are distinctive.

594

R UBIACEAE

Faramea egregia Sandwith, Kew Bull. 1939: 12. 1939. Shrub or treelet to 5 m tall; leaves 5.5– 17.5 × 1.5–5.5 cm, petiolate; stipules caducous, interpetiolar, ca. 1 mm long, aristas 2–9 mm long; flowers axillary, solitary, pedunculate; calyx limb 4–6 mm long, truncate; corolla white, tube 15–22 mm long, lobes 16–30 mm long; fruit subglobose, 9–12 mm diameter. Lower montane forests, 500–600 m; eastern Bolívar (basin of Río Cuyuní). Guyana. ◆Fig. 470. Faramea egregia is similar to F. yutajensis, and these species can be confused when the distinctive floral bracts of the latter species have fallen. Faramea maguirei Steyerm., Mem. New York Bot. Gard. 17(1): 398. 1967. Shrub or small tree to 5 m tall; leaves 15– 24 × 5.5–10 cm, petiolate; stipules caducous, shortly tubular, 3–4 mm long, aristas 2–7 mm long; inflorescence axillary, subcapitate, subsessile; calyx limb ca. 1 mm long, dentate; corolla white, tube 15–16 mm long, lobes 9– 10 mm long; fruit markedly oblate, 5 × 9–11 mm, ridged. Lower montane forests, 500–600 m; eastern Bolívar (basin of Río Cuyuní). Guyana. Faramea morilloi Steyerm., Ann. Missouri Bot. Gard. 75: 337. 1988. Shrub or subshrub to 2 m tall; leaves 8– 12.5 × 1.5–4.5 cm, petiolate; stipules caducous, interpetiolar, ca. 2 mm long, aristas 2–5 mm long; inflorescence axillary, pedunculate, flowers 1–3, subsessile, subtended by 1–3 pairs of bracts; calyx limb ca. 2 mm long, lobed; corolla cream, tube ca. 17 mm long, lobes 7–10 mm long; fruit subglobose to oblate, 8–9 × 10–12 mm, ridged. Semideciduous forests, ca. 100 m; Bolívar (Las Claritas, upper Río Caura, Río Karún, 64 km southeast of Los Pijiguaos), Amazonas (between Puerto Ayacucho and Gavilán). Guyana, northern Brazil. Faramea multiflora A. Rich. ex DC., Prodr. 4: 497. 1830. —Yaerute ukwae (Piaroa). Faramea multiflora var. epedunculata Steyerm., Mem. New York Bot. Gard. 17(1): 392. 1967. Shrub or tree to 10 m tall; leaves 7.5–17 × 1.5–6 cm, petiolate; stipules generally persis-

tent, tubular, 2–10 m long, aristas 1.5–4 mm long; inflorescence terminal, cymose, pedunculate; calyx limb 1–1.5 mm long, denticulate to shortly lobed; corolla very pale to deep blue, tube 5–7 mm long, lobes 4–6.5 mm long; fruit oblate, 5–6 × 9–11 mm. Evergreen lowland to lower montane forests, 100–800 m; Bolívar (widespread), Amazonas (Río Atabapo near Caño Caname, upper Río Cuao, base of Sierra de la Neblina). Central America, Colombia, Guyana, French Guiana, Suriname, Ecuador, Peru, Brazil, Bolivia. Faramea multiflora is widespread, frequently collected, especially in western Amazonia, and morphologically quite variable. Steyermark (1967, 390–395) recognized six varieties of Faramea multiflora based largely on overlapping ranges of flower, fruit, and leaf size. He reported two of these, var. epedunculata Steyerm. and var. multiflora, from the flora area. However, none of these varieties were recognized by Taylor (1999, 294–298), and as discussed there appear to represent only distinctive but not completely separable combinations of continuously varying characters. Faramea quinqueflora Poepp. & Endl. is similar to F. multiflora, but can be distinguished by its inflorescences with 3–9 flowers borne in a single, pedunculate, terminal, umbelliform fascicle. Faramea quinqueflora is known from Amazonian southern Colombia, Ecuador, Peru, west-central Brazil, and east to French Guiana, and thus might be expected in Amazonas. Faramea neblinae Steyerm., Mem. New York Bot. Gard. 17(1): 395. 1967. Shrub or tree to 4 m tall; leaves 9–16 × 3– 8.5 cm, petiolate; stipules caducous, interpetiolar, 2–4 mm long; inflorescence terminal and sometimes also axillary, cymose, pedunculate; calyx limb ca. 3.5 mm long, truncate; corolla white, tube 17–18 mm long, lobes 14– 18 mm long; fruit subglobose, ca. 9 mm diameter. Upper tepui slope forests, 1100–1400 m; Amazonas (Sierra de la Neblina). Endemic. Faramea occidentalis (L.) A. Rich., Mém. Rubiaceae 96. 1830. —Ixora occidentalis L., Syst. Nat. ed. 10, 2: 893. 1759. —Cafecillo, Cafecillo de danta, Cafecito, Cruceto, Guaricha, Jazmín montañero, Pata de grulla.

Faramea 595

Faramea occidentalis subsp. occidentalis var. meridionalis Steyerm., Mem. New York Bot. Gard. 17(1): 385. 1967. Tree or shrub to 10(–18) m tall; leaves 8.5– 16 × 3–8 cm, petiolate; stipules caducous, interpetiolar, 2.5–3.5 mm long, aristas 2–6 mm long; inflorescence terminal and sometimes also axillary, cymose, pedunculate; calyx limb 1.5–3 mm long, truncate to denticulate; corolla white, tube 15–20 mm long, lobes 17–25 mm long; fruit subglobose to oblate, 5–10 × 8–15 mm. Evergreen lowland to lower montane forests, 50–800 m; Bolívar (Río Caura), Amazonas (Sierra Parima). Widespread elsewhere in Venezuela; Mexico, Central America, Antilles, Colombia, Guyana, Suriname, Ecuador, Peru, Brazil, Bolivia. Faramea occidentalis is widespread and morphologically variable in a number of features. The variation in corolla size in the South American plants appears to be partly clinal, with the flowers larger in the northeastern part of the continent, including the flora area. Several rather distinctive forms of F. occidentalis have been named. Steyermark (1967, 384–385) recognized two subspecies of F. occidentalis, but his subsp. lonchocalyx Steyerm. of western Ecuador has more recently been treated as a synonym of F. spathacea Müll. Arg. ex Standl. (Taylor 1999, 305–306.). Within subsp. occidentalis Steyerm., he recognized three varieties based on overlapping size ranges of the calyx limb and corolla. Plants of the flora area mostly fall into his var. meridionalis Steyerm., but because the variation in these features is continuous and these varieties cannot clearly be delimited, they do not seem taxonomically informative and are not recognized here. Faramea stenopetala is very similar to F. occidentalis, and they may not be completely distinct species but at present appear separable. Faramea cyanea Müll. Arg. is very similar to F. occidentalis, and replaces it from south-central Brazil southward; the distinctions between these species are not completely clear. Faramea orinocensis Standl., Publ. Field Columbian Mus., Bot. Ser. 7: 418. 1931. Shrub or tree to 4 m tall; leaves 4–10 × 2– 5 cm, shortly petiolate; stipules usually caducous, interpetiolar, 1–1.5 mm long, aristas 3–

4 mm long; inflorescence terminal, cymose, pedunculate; calyx limb 1.5–2 mm long, denticulate; corolla white, tube 12.5–14 mm long, lobes 12.5–13 mm long; fruit ca. 6 × 7 mm. Riparian forests, evergreen seasonally dry forests, 100–200 m; Bolívar (tributaries of Río Orinoco, Río Paragua), Amazonas (near Puerto Ayacucho). Anzoátegui, Apure, Guárico; eastern Colombia. Faramea parvibractea Steyerm., Mem. New York Bot. Gard. 17(1): 376. 1967. Shrub or small tree to 4 m tall; leaves 8.5– 14 × 1.5–5.5 cm, petiolate; stipules caducous, calyptrate, 1–3 mm long, without aristas; inflorescence terminal, with 7–19 fasciculate peduncles, each terminating in a small fascicle, this often enclosed in a pair of bracts 5– 12 × 3–7 mm; calyx limb 1–1.5 mm long, denticulate; corolla white, tube 6–7.5 mm long, lobes 6–9.5 mm long; fruit oblate, 5–6 × 8 mm. Evergreen lowland to montane forests, 100–1100 m; Bolívar (Río Caura) Amazonas (Cerro Huachamacari, Río Casiquiare, Río Cuao, Río Orinoco). Central America, Colombia, Ecuador, Brazil. Faramea sessilifolia (Kunth) DC., Prodr. 4: 497. 1830. —Tetramerium sessifolium Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 3: 374. 1818 [1819]. —Cafecillo, Guarataro. Shrub or small tree to 7 m tall; leaves 10.5–26 × 3.5–8 cm, subsessile; stipules caducous, interpetiolar, 2–7 mm long, aristas 5–13 mm long; inflorescence terminal, cymose, pedunculate; calyx limb ca. 0.5 mm long, denticulate; corolla pale to deep blue, tube 4.5–6.5 mm long, lobes 3.5–5 mm long; fruit oblate, 5–6 × 6–7 mm. Evergreen lowland forests, granitic outcrops, 50–200 m; Amazonas (common in the basins of the upper Río Orinoco, Río Casiquiare, Río Yatúa, and the Río Negro northeast to Río Ventuari). Colombia, Guyana, Peru, Bolivia, Brazil. ◆Fig. 469. The size and especially the shape of the leaves is notably variable in Faramea sessilifolia. This name is often misspelled as “Faramea sessifolia.” Faramea sessifolia P.H. Allen is based on a type from Costa Rica and is a synonym of F. tamberlikiana. Faramea sessilifolia is also related to F. tamberlikiana; for further discussion of the relationships among these species see Taylor (2002a).

596

R UBIACEAE

Faramea stenopetala Mart., Flora 24(Beibl. 2): 73. 1841. —Jasmín montañero. Shrub or small tree to 10 m tall; leaves 12–22 × 3–7 cm, petiolate; stipules caducous, interpetiolar, ca. 2 mm long, aristas 6–9 mm long; inflorescence terminal and sometimes also axillary, cymose, pedunculate; calyx limb ca. 3 mm long, denticulate; corolla white, tube 15–23 mm long, lobes 10–12 mm long; fruit oblate, ca. 8 × 10 mm. Riparian forests, 50–200 m; Amazonas (La Esmeralda, San Fernando de Atabapo, Tama Tama). Amazonian Brazil. Faramea tamberlikiana Müll. Arg. in Mart., Fl. Bras. 6(5): 133. 1881. Southern Central America, Colombia, Ecuador, Peru, Brazil, Bolivia; 2 subspecies, 1 in Venezuela. The second subspecies of this species, subsp. sessifolia (P.H. Allen) C.M. Taylor, is found in southern Central America and coastal western Colombia (Taylor 2002a). F. tamberlikiana subsp. tamberlikiana Shrub or small tree to 4 m tall; leaves 8– 25 × 3.5–11 cm, shortly petiolate; stipules caducous, interpetiolar, 4–6 mm long, aristas 3.5–5 mm long; inflorescence terminal, cymose, pedunculate; calyx limb ca. 0.2 mm long; corolla pale to dark blue-violet, tube ca. 6 mm long, lobes ca. 3 mm long; fruit oblate, ca. 5 × 7 mm. Evergreen lowland forests, 100–200 m; Amazonas (La Esmeralda). Colombia, Ecuador, Peru, Brazil, Bolivia. Faramea acuminatissima Müll. Arg. may be an older name for this species, but the information available about that name is inadequate to confirm this. If so, the type collection of F. acuminatissima has atypically very narrow leaves for this species. Faramea torquata Müll. Arg., Flora 58: 471. 1875. —Cafecillo, Carrisillo, Palo de tema, Pata de grulla. Faramea pachydictyon Müll. Arg., Flora 58: 471. 1875. Faramea paludicola Steyerm., Ann. Missouri Bot. Gard. 75: 338. 1988. Shrub or tree to 3 m tall; leaves 15–25 × 3.5–9 cm, petiolate; stipules interpetiolar,

sometimes persistent, 2–9 mm long, aristas absent or 0.5–1 mm long; inflorescences terminal and sometimes also axillary, cymose, pedunculate; calyx limb 2–3 mm long, subtruncate; corolla white, tube 8–12 mm long, lobes 6–8 mm long; fruit subglobose to oblate, 8–13 mm diameter. Evergreen lowland forests, 100–400 m; Bolívar (near Upata), Amazonas (upper Río Baría, Río Casiquiare, Río Pasimoni, Río Vasiva, San Carlos de Río Negro, Solano). Amazonian Colombia, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 466. Steyermark separated Faramea paludicola from F. torquata only by its leaves with the “marginal nerve prominent,” but this feature varies continuously in the ample material of F. torquata available from individual study sites in Ecuador and Peru, and thus does not seem taxonomically informative and thus does not support the separation of these as distinct species. Faramea yavitensis Steyerm., Ann. Missouri Bot. Gard. 75: 338. 1988. Shrub or small tree to 2 m tall; leaves 11– 21 × 3.5–9 cm, petiolate; stipules caducous, interpetiolar, 2–3 mm long, aristas 2–4 mm long; inflorescence terminal, cymose, pedunculate; calyx limb 2–3.5 mm long, deeply lobed; corolla white, tube ca. 7 mm long, lobes ca. 5 mm long; fruit oblate, 4–6 × 6–8 mm. Evergreen lowland forests, 100–200 m; Amazonas (Río Atacavi, Río Caname, Río Sipapo, near Yavita). Endemic. Faramea yutajensis Steyerm., Ann. Missouri Bot. Gard. 75: 339. 1988. Shrub to 2 m tall; leaves 8–14 × 2–4.5 cm, petiolate; stipules caducous, interpetiolar, ca. 1 mm long, aristas 1–2 mm long; flowers terminal and/or axillary, solitary, subsessile to pedunculate, subtended by a pair of bracts 13–23 × 6–8 mm; flowers unknown; fruit subglobose, 9–11 mm diameter, with persistent calyx limb ca. 2 mm long. Tepui slope forests, 1700–1800 m; Amazonas (west of Cerro Yutajé). Endemic. Faramea yutajensis is similar to F. egregia, and when the floral bracts of F. yutajensis have fallen after anthesis these two species can easily be confused.

Faramea 597

Fig. 466. Faramea torquata

Fig. 467. Faramea capillipes

598

R UBIACEAE

Fig. 468. Faramea cardonae

Faramea 599

Fig. 469. Faramea sessilifolia

Fig. 470. Faramea egregia

600

R UBIACEAE

34. FERDINANDUSA Pohl, Pl. Bras. Icon. Descr. 2: 8, t. 106–108. 1831. —Ferdinandea Pohl, Flora 10: 153. 1827, hom. illeg., non Ferdinanda Lag. 1816. Aspidanthera Benth., J. Bot. (Hooker) 3: 217. 1841. Gomphosia Wedd., Ann. Sci. Nat. Bot. sér. 3, 10: 14. 1848. by Charlotte M. Taylor, Elisete Araujo da Anunciação, and Julian A. Steyermark Shrubs to large trees, unarmed, terrestrial, with the branches sometimes clambering. Leaves opposite, petiolate, venation not lineolate; stipules interpetiolar, caducous, triangular, twisted in bud. Inflorescences terminal and in the uppermost leaf axils, sessile to pedunculate, cymose, multiflowered, bracteate. Flowers medium-sized to large, subsessile to pedicellate, homostylous, protandrous. Hypanthium ellipsoid to turbinate. Calyx limb shortly lobed, lobes 4 or 5, without calycophylls; corolla funnelform to salverform, white to pale green or red, internally glabrous, lobes 4 or 5, emarginate in bud becoming bifid at anthesis, convolute in bud. Stamens 4 or 5, inserted in upper part of corolla tube; anthers ellipsoid, exserted or included, dorsifixed near middle or sometimes versatile, exserted or included. Ovary 2-locular; ovules numerous in each locule, on axile placentas. Fruit capsular, cylindrical to narrowly oblong or obovoid, woody, septicidal from the apex. Seeds small, elliptic to obovate or oblong, flattened, papery, with margins winged, entire to lacerate or rarely fimbriate. Central America, Colombia, Venezuela, Guyana, French Guiana, Ecuador, Peru, Brazil, Bolivia; ca. 26 species, 6 in Venezuela, all in the flora area. Ferdinandusa is similar to and sometimes confused with Ladenbergia, especially in fruit; their separation is discussed under Ladenbergia. Steyermark (e.g., 1974) has described the capsules of Ferdinandusa as loculicidal, but as noted by Andersson and Taylor (1994, 7–11) they actually are septicidal. Key to the Species of Ferdinandusa 1. 1. 2(1).

2.

3(1). 3. 4(3). 4. 5(4).

5.

Inflorescence axes glabrous ....................................................................... 2 Inflorescence axes pubescent..................................................................... 3 Flowers few (4–8) in each inflorescence, these subumbellate or fasciculate; pedicels 9–11.5 long; peduncles 0.5–1.2 cm long; calyx and corolla lobes 4; corolla 35–45 mm long; stigma included ..................... F. guainiae Flowers numerous in each inflorescence, these pyramidal to corymbiform cymes; pedicels at most 2.5 mm long; peduncles 3.5–6 cm long; calyx and corolla lobes 5; corolla 15–20 mm long; stigma exserted .................................................................................................. F. uaupensis Lower surface of leaf hirsute, the trichomes to 1 mm long ...... F. neblinensis Lower surface of leaf puberulous, tomentulose, or strigillose, the trichomes < 0.5 mm long ........................................................................... 4 Corolla tube 35–36 mm long ............................................................ F. boomii Corolla tube 11–31 mm long ...................................................................... 5 Leaves rounded to obtuse or acute at base, with secondary veins 7– 12 pairs; petioles glabrous or strigillose throughout or only on upper side; seeds lacerate ................................................................ F. goudotiana Leaves subcordate to rounded at base, with secondary veins 12–19 pairs; petioles tomentulose throughout; corolla tube 21–31 mm long; seeds fimbriate ........................................................................................ F. sprucei

Ferdinandusa 601

Ferdinandusa boomii Steyerm., Ann. Missouri Bot. Gard. 75: 339. 1988. Tree to 10 m tall; stems puberulous; leaves 7.3–14 × 5–9 cm; petioles 0.8–1.5 cm long; stipules unknown; calyx limb and hypanthium together 2–2.5 mm long (length of the calyx limb along unknown); corolla white, tube 35–36 mm long, lobes 4, 4.5–5 mm long; fruit unknown. Evergreen lowland forests, 100–200 m; Amazonas (Río Atabapo, Sierra de la Neblina). Endemic. This species is known from only two collections. It is similar to Ferdinandusa schultesii Steyerm. of Amazonian Colombia. Ferdinandusa goudotiana K. Schum. in Mart., Fl. Bras. 6(6): 211. 1889. Tree to 18 m tall; stems strigillose; leaves 12–20 × 6.5–9 cm; petioles 1–1.7 cm long; stipules 1.7–2.5 cm long; calyx limb ca. 1.5 mm long; corolla white to pale green, tube 11–20 mm long, lobes 4, 4–6 mm long; fruit 2–5 × 0.8 cm. Evergreen forests, riparian forests, gallery forests, 100–1400 m. Venezuela, Guyana, Brazil; 3 varieties, all in the flora area. Key to the Varieties of F. goudotiana 1. Hypanthium pubescent; lower surface of leaf with areoles pilose .......................... ................................... var. goudotiana 1. Hypanthium glabrous or sparsely pubescent; lower surface of leaf with areoles glabrous to glabrescent .................... 2 2. Calyx lobes glabrous throughout .............. ......................................... var. eciliata 2. Calyx lobes ciliolate at apex ...................... .................................... var. psilocarpa F. goudotiana var. eciliata Steyerm., Mem. New York Bot. Gard. 23: 281. 1972. Bolívar (common in the Gran Sabana, Macizo del Chimantá, Ptari-tepui, Río Caroní, Uaipán-tepui), Amazonas (upper Río Orinoco). Guyana, Brazil (Rio Branco). ◆Fig. 472. This is the most common variety in the flora area. F. goudotiana var. goudotiana Bolívar (Cerro Ichún, Roraima-tepui), Amazonas (Cerro Yutajé, San Carlos de Río Negro, Sierra Parima). Brazil. F. goudotiana var. psilocarpa Steyerm., Mem. New York Bot. Gard. 23: 281. 1972. Bolívar (Gran Sabana, near Luepa). Guyana.

Ferdinandusa guainiae Spruce ex K. Schum. in Mart., Fl. Bras. 6(6): 208. 1889. Shrub or tree to 18 m tall; stems glabrous; leaves 4–7.5 × 2–4.5 cm; petioles 0.3–0.6 mm long; stipules ca. 1 cm long; calyx limb ca. 1 mm long; corolla green to white, tube 29–39 mm long, lobes 4, 6–7 mm long; fruit ca. 3 × 1.2 cm. Evergreen dwarf forests, riparian forests, caatinga formations on white sand, 100–200 m; southwestern Amazonas (Río Guainía basin). Colombia, Brazil. ◆Fig. 473. Ferdinandusa guainiae has umbelliform or apparently fasciculate inflorescences due to the highly reduced axes but elongated pedicels. This is the only Ferdinandusa species with included stigmas. Ferdinandusa neblinensis Steyerm., Mem. New York Bot. Gard. 23: 283, fig. 57A–D. 1972. Shrub or tree to 10 m tall; stems hirtellous to hirsute; leaves 6–19 × 2.5–7 cm; petioles 0.4–0.5 mm long; stipules 1–1.6 cm long; calyx limb 1–1.5 mm long; corolla white, tube 20–26 mm long, lobes 4, ca. 6 mm long; fruit 3–4 × 0.5 cm. Riparian and mixed lower montane wet forests, 100–500 m; Amazonas (Cerro Aracamuni, base of Sierra de la Neblina). Endemic. ◆Fig. 471. Ferdinandusa sprucei K. Schum. in Mart., Fl. Bras. 6(6): 210. 1889. Shrub or tree to 5 m tall; stems tomentulose; leaves 20–23 × 7–7.5 cm long; petioles 1–1.2 cm long; stipules 2–2.2 cm long; calyx limb 1–1.5 mm long; corolla white, tube 21–31 mm long, lobes 4, ca. 6 mm long; fruit 10–11.7 × 5–8 cm. Evergreen lowland forests, 100–200 m; southwestern Amazonas (Río Baría, Río Pasimoni, San Fernando de Atabapo, Santa Bárbara, base of Sierra de la Neblina, Yavita). Adjacent Colombia and Brazil. This is the only species of Ferdinandusa with fimbriate seeds. Ferdinandusa uaupensis Spruce ex K. Schum. in Mart., Fl. Bras. 6(6): 212. 1889. Tree to 18 m tall; stems glabrous; leaves 9–16 × 9–19 cm; petiole 1–1.2 cm; stipules unknown; calyx limb 1–1.5 mm long; corolla pale green to white, tube ca. 15 mm long, lobes 5, 3–3.4 mm; fruit unknown. Rain forests, 100–300 m; southwestern Amazonas (San Carlos de Río Negro, Santa Bárbara, Sierra de la Neblina). Amazonian Colombia and Brazil. ◆Fig. 474.

602

R UBIACEAE

Fig. 471. Ferdinandusa neblinensis

Fig. 472. Ferdinandusa goudotiana var. eciliata

Ferdinandusa 603

Fig. 473. Ferdinandusa guainiae

Fig. 474. Ferdinandusa uaupensis

604

R UBIACEAE

35. GALIUM L., Sp. Pl. 105. 1753. Galium sect. Relbunium Endl., Gen. Pl. 523. 1839. —Relbunium (Endl.) Benth. & Hook. f., Gen. Pl. 2: 149. 1873. by Charlotte M. Taylor and Julian A. Steyermark Annual or usually perennial herbs, terrestrial, unarmed, with stems erect to spreading, often elongated to subscandent, usually 4-angled. Leaves small, subsessile, opposite but appearing whorled in groups of 4–10 due to the interpetiolar stipules being foliaceous, simple or deeply divided with the segments similar to the leaves, venation not lineolate; stipules apparently lacking. Inflorescence axillary, subsessile to pedunculate, cymose to paniculate and few-flowered to multiflowered or reduced to a single flower, bracteate. Flowers small, sessile or pedicellate, homostylous and protandrous (or sometimes unisexual on dioecious plants in species from outside the flora area). Hypanthium subglobose. Calyx limb reduced to a minute rim or absent; corolla campanulate to rotate, white to pale green, yellow, dull red, or purple, internally glabrous, lobes 4, valvate in bud. Stamens 4, inserted in the mouth of the corolla tube; anthers elliptic to basifixed, exserted, didymous. Ovary 2-locular; ovules solitary in each locule, axile. Fruit indehiscent, didymous and dry to leathery or occasionally fleshy and white, orange, or red, glabrous or pubescent sometimes with long uncinate trichomes. Seeds small, oblong, the ventral face deeply excavate. Cosmopolitan; ca. 400 species, 5 in Venezuela, 1 of these in the flora area. Galium grows in cool areas, with its species found throughout the temperate zone but only in premontane and montane vegetation in the tropics. The remaining Venezuelan species of Galium are found in the Andes. Galium is often overlooked because it appears to lack stipules between its whorled leaves. In the flora area, one species of Limnosipanea is similar in this regard; however L. sprucei grows in warm savanna habitats. The genus Relbunium was previously separated from Galium based on its flowers that are subtended individually or in cymules by an involucre composed of several bracts together with its fruits that are leathery to succulent. However, recent studies of Galium (e.g., Dempster 1990) have concluded that these features are uncorrelated and have arisen more than once, and thus that Relbunium comprises a polyphyletic assemblage that is derived from and better treated in Galium. Galium hypocarpium (L.) Endl. ex Griseb., Fl. Brit. W. I. 4: 351. 1861. —Valantia hypocarpia L., Syst. Nat. ed. 10, 1304. 1759. —Relbunium hypocarpium (L.) Hemsl., Biol. Cent.-Amer., Bot. 2: 63. 1881. Galium croceum Ruiz & Pav., Fl. Peruv. 1: 59. 1798. —Relbunium croceum (Ruiz & Pav.) K. Schum. in Mart., Fl. Bras. 6(6): 116. 1888. Rubia nitida Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 3: 339. 1818 [1819]. —Relbunium nitidum (Kunth) K. Schum. in Mart., Fl. Bras. 6(6): 114. 1888. —Relbunium hypocarpium subsp. nitidum (Kunth) Ehrend., Bot. Jahrb.

Syst. 76: 536. 1955. Relbunium glaberrimum Standl., Publ. Field Columbian Mus., Bot. Ser. 4: 297. 1929. Anzoátegui, Aragua, Carabobo, Distrito Federal, Mérida, Miranda, Sucre, Táchira, Trujillo, Zulia; Mexico, Central America, Colombia, Guyana, Ecuador, Peru, Brazil, Bolivia, Chile, Paraguay, Argentina, Uruguay; 6 varieties, 1 in Venezuela. Dempster (1990) recognized six varieties of this widely distributed, morphologically variable species. However, most plants of Galium hypocarpium, including those of the northern Andes and the flora area, belong to var. hypocarpium.

Genipa 605

G. hypocarpium var. hypocarpium Scandent to sprawling perennial herb, frequently forming tangled masses; leaves in whorls of 4, 3–22 × 1.5–10 mm; flowers 1–4 per axil, individually involucrate; corolla cream to yellow, ca. 2 mm long; fruits succulent, orange to red, 2–3.5 mm diameter. Open areas on tepui summits, 2000–2100 m; Bolívar (Macizo del Chimantá, Roraimatepui). Other distribution as in species. ◆Fig. 475. Steyermark (1974) treated the plants from the flora area as var. nitidum, but Dempster (1990) showed that the pubescence and ecological characters he used to distinguish var. nitidum from var. hypocarpium vary widely throughout the range of this species and consequently she did not recognize var. nitidum.

Fig. 475. Galium hypocarpium var. hypocarpium

36. GENIPA L., Syst. Nat. ed. 10, 930. 1759. by Julian A. Steyermark and Claes Persson Glabrous or pubescent trees, terrestrial, unarmed, hermaphrodite or pistillate. Leaves opposite, papyraceous to subcoriaceous, petiolate, venation brochidodromous, tertiary venation not lineolate; stipules interpetiolar, persistent or sometimes caducous, generally triangular, slightly connate or free at base. Inflorescence pseudoaxillary or terminal, pedunculate, bracteate, the hermaphroditic a multiflowered to few-flowered cyme, the pistillate with flowers solitary (or 2). Flowers bisexual or pistillate, often subtended by narrowly triangular bracts, rarely widely triangular or subovate. Hypanthium ellipsoid to turbinate. Calyx limb tubular, wavy, jagged, or with (4)5(6) minutely deltoid teeth, sericeous or glabrous except for ring of hairs at base internally; corolla shortly salverform, white, turning yellow after anthesis, densely appressed-pubescent externally, densely pubescent at the base of lobes and throat internally, lobes (4)5(6), contorted in bud. Stamens (4)5(6), inserted near the top of the corolla tube; anthers dorsifixed near base, narrowly oblong, exserted; filaments very short or absent, with 3-colporate pollen. Ovary 2-locular; ovules numerous in each locule, placenta axial and parietal; style branches 2, elliptic, ridged externally, sometimes slightly revolute. Fruit baccate, ellipsoidal or subglobose, usually multi-locular due to formation of false septa, pulp and pericarp fleshy, exocarp yellowish brown, khaki-colored, brownish, or grayish brown. Seeds numerous, large, sublenticular or trapezoid, horizontally arranged in tiers; testa firm, of elongated cells with secondary thickenings in the radial walls and transverse ribs in internal tangential wall. Mexico, Central America, West Indies, Colombia, Venezuela, Trinidad-Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina; 3 species, 2 in Venezuela, both in the flora area. As suggested by a phylogenetic analysis using plastid DNA sequences, the neotropical Genipa, as traditionally circumscribed, is paraphyletic. One group of dioecious species is more closely related to other neotropical Gardenieae taxa, whereas Genipa sensu stricto is more closely related to Old World Gardenieae

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(Persson 2000b). As a result, two former Genipa species were recently transferred to the new genus Agouticarpa C. Persson (Persson 2003). Despite its close relationship to paleotropical genera, it is assumed that Genipa sensu Drake from Madagascar is not congeneric Genipa sensu stricto (Bridson and Robbrecht 1985). The flowers of Genipa americana have been described as either bisexual or unisexual. In the present study each recorded inflorescence of G. americana and G. spruceana consists of either bisexual flowers or pistillate flowers, indicating that both species are gynodioecious. However, according to label information and other studies (e.g., Zappi et al. 1995), staminate flowers are common. If this is correct, G. americana is polygamous rather than gynodioecious. Clearly, further studies are needed in order to elucidate its exact sex distribution. Key to the Species of Genipa 1.

1.

Interior of calyx tube sericeous; interior of corolla tube with conspicuous hirsute upwardly pointing pubescence not markedly differing from the dense, erect brush of hairs in the basal portion either in length or direction of pubescence; base of corolla lobes within conspicuously covered with a mat of elongated, loose, golden brown (in dried material) hirsute pubescence; vegetative parts glabrous to pubescent; upper leaf surface dull or shining; tertiary leaf venation reticulate and prominent or not showing on both surfaces; fruits solitary or rarely two, when dry usually smooth, khaki-colored or rarely black, 5.5–10 × 5.5–7 cm ................................................................................................ G. americana Interior of calyx tube glabrous except for some minute basal pubescence; interior of corolla tube with short, villous, loosely spreading pubescence in upper portion, this markedly contrasting with the dense, erect brush of hairs in the basal portion both as to length and direction of pubescence; base of corolla lobes within shortly villous-pubescent; vegetative parts completely glabrous; upper leaf surface shining; tertiary leaf venation reticulate and prominent on both surfaces; fruits 1–3, when dry usually wrinkled and brownish, 2.5–4 × 3–4.5 cm ......... G. spruceana

Genipa americana L., Syst. Nat. ed. 10, 931. 1759. Tree 4–27 m tall. Forests and forest edges, 50–800 m. Widely distributed in Mexico, Central America, West Indies, Colombia, Venezuela, Trinidad-Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina; 2 varieties, both in the flora area. This treatment follows the traditional division of Genipa americana into two varieties based on the amount of pubescence on the lower leaf surface. However, whether this character is correlated with other traits is not clear. Differences in habit and an ecological separation have been noted between the

two varieties in Costa Rica (C. M. Taylor personal communication). Furthermore, there seems to be a difference in leaf blade shape (short and wide in var. caruto versus long and narrow in var. americana). However, it is unclear if these differences will hold up in a more detailed study; therefore this treatment must be considered provisional. The juicy fruits are edible and are either eaten fresh or used in preserves or drinks. The juice turns dark blue to black when exposed to air and is used extensively by indigenous people as a body paint. Genipa americana is a good timber tree and is used locally for a variety of purposes in carpentry.

Genipa 607

Key to the Varieties of G. americana 1. Lower surface of leaf blade glabrous or nearly so, only the midrib and/or secondary veins sparsely pubescent; upper leaf surface not rugulose .. var. americana 1. Lower surface of leaf blade, including midrib and secondary veins, densely soft-pubescent; upper leaf surface sometimes rugulose ............................. var. caruto G. americana var. americana. ––Caruto. Delta Amacuro (east-northeast of El Palmar, east of Río Grande), Bolívar (between

Upata and Villa Lola), Amazonas (widespread and common). Anzoátegui, Apure, Carabobo, Cojedes, Distrito Federal, Lara, Mérida, Trujillo, Yaracuy, Zulia; Mexico, Costa Rica, Panama, West Indies, Colombia, Suriname, Ecuador, Peru, Brazil, Bolivia, Argentina. G. americana var. caruto (Kunth) K. Schum. in Mart., Fl. Bras. 6(6): 352 1889. —Genipa caruto Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 3: 407. 1818 [1820]. ––Caruto, Caruto montañero, Jagua.

Fig. 476. Genipa spruceana

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Delta Amacuro (Río Acure, Serranía de Imataca), Bolívar (Altaplanicie de Nuria south to Santa Elena de Uairén and west to Río Caura), Amazonas (Puerto Ayacucho). Aragua, Barinas, Carabobo, Guárico, Lara, Monagas, Portuguesa, Trujillo, Yaracuy; Mexico, Central America, West Indies, Colombia, Trinidad-Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina. Genipa spruceana Steyerm., Mem. New York Bot. Gard. 23: 353, fig. 62a–c. 1972. —Huitillo. Tree 3–20 m tall. Evergreen lowland to lower montane and riparian forests, sand

and gravel bars, 50–700 m; Bolívar (common), Amazonas (Río Cuao, Río Guainía, lower Río Mawarinuma, upper Río Orinoco, Río Siapa, Río Yatúa). Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Amazonian Brazil. ◆Fig. 476. Genipa spruceana is a frequent pioneer on newly formed sand beaches and gravel bars along streams. It is sometimes considered merely an ecotype of G. americana. However, in addition to the traditional differentiating flower and vegetative characters, there are also general differences in number of fruits per infructescence, fruit size, and color and texture of the fruit when dry, which supports their separation into two species.

37. GEOPHILA D. Don, Prodr. Fl. Nepalensis 136. 1825, nom. cons., not Geophila Bergeret 1803. Carinta W.F. Wight, Contr. U.S. Nat. Herb. 9: 216. 1905. Geocardia Standl., Contr. U.S. Nat. Herb. 17: 444. 1914. by Charlotte M. Taylor and Julian A. Steyermark Creeping perennial herbs, terrestrial, unarmed, rooting at nodes. Leaves opposite, petiolate, generally ovate with cordate bases, venation not lineolate; stipules interpetiolar, persistent, generally imbricate to valvate in bud, triangular to 2(3)lobed. Inflorescences axillary and terminal, borne on short erect stems, pedunculate, subcapitate, 1–20-flowered, bracteate. Flowers small, sessile to shortly pedicellate, homostylous, protandrous. Hypanthium turbinate to cylindrical. Calyx limb deeply lobed, lobes (4)5(–7), without calycophylls; corolla salverform to funnelform, white, internally densely hirtellous to villous in upper part of tube, lobes (4)5(–7), valvate in bud. Stamens (4)5(–7), inserted in upper part of corolla tube; anthers dorsifixed near middle, narrowly oblong, included. Ovary 2-locular; ovules solitary in each locule, basal. Fruit drupaceous, ellipsoid to subglobose, fleshy, red or sometimes black; pyrenes 2, bony, planoconvex, 1-locular, dorsally smooth to longitudinally ridged, sometimes twisted. Seeds small, planoconvex to ellipsoid. Mexico, Central America, West Indies, Colombia, Venezuela, Trinidad-Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina, Africa, Asia; ca. 20 species, 5 species in Venezuela, 4 of these in the flora area. Key to the Species of Geophila 1. 1. 2(1).

2.

Hypanthium and fruit glabrous; leaf blades acute to usually obtuse or rounded at apex ..................................................................................... 2 Hypanthium and fruit pubescent; leaf blades acute at apex ................... 3 Leaf blades with basal lobes prominently incurved, forming a horseshoeshaped sinus; peduncles 0.4–1 cm long; corolla lobes pubescent with apical trichomes only; trichomes on petioles retrorse, subappressed, 0.2–0.3 mm long ..................................................................... G. orbicularis Leaf blades with basal lobes overlapping or meeting in the center; peduncles 0.5–6.5 cm long; corolla lobes glabrous or pubescent but in this

Geophila 609

3(1). 3.

case more than just apical trichomes; trichomes on petioles ascending, spreading, 0.3-0.5 mm long .......................................................... G. repens Calyx tube externally hispidulous, the lobes ciliate with trichomes 0.8– 1.2 mm long; leaf blades 2.5–10 × 1.2–8.5 cm ........................ G. cordifolia Calyx tube externally glabrous, the lobes glabrous or with a few trichomes to 0.4 mm long; leaf blades 1–4 × 0.8–3 cm .................................. G. tenuis

Geophila cordifolia Miq., Stirp. Surinam. Select. 176. 1850 [1851]. Plants hirsute, at least sparsely; leaves 2.5–10 × 1.2–8.5 cm; stipules 2–6 mm long; peduncles 0.5–3.2 cm long; calyx limb 3–3.5 mm long; corolla tube 2.5–4 mm long, lobes 1.5–3 mm long; fruit ca. 8 mm diameter, red. Central America, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 2 varieties, 1 in Venezuela. G. cordifolia var. cordifolia Evergreen lowland to montane forests, 50–1200 m; Bolívar (vicinity of Canaima, Kavanayén, upper Río Paragua), Amazonas (Cerro Duida, Cerro Sipapo, Piedra Cucuy, Río Cataniapo, Sierra de la Neblina). Tá-

chira; Colombia, Guyana, Suriname, French Guiana, Brazil. ◆Fig. 477. Geophila orbicularis (Müll. Arg.) Steyerm., Mem. New York Bot. Gard. 23: 397. 1972. —Mapouria herbacea var. orbicularis Müll. Arg. in Mart., Fl. Bras. 6(5): 427. 1881. Plant glabrescent; leaves 1.2–3.5 × 1–3.5 cm; stipules 2.5–4.5 mm long; peduncles 0.4– 1 cm long; calyx limb 3–3.5 mm long; corolla tube ca. 5.5 mm long, lobes ca. 3 mm long; mature fruit unknown, apparently red. Forests, 100–1100 m. Venezuela, Brazil; 2 varieties, both of these in the flora area. Although no descriptions of the fruits have been presented, Steyermark (1972; 1974) keyed this species as having red fruits.

Fig. 477. Geophila cordifolia var. cordifolia

Fig. 478. Geophila repens

Fig. 479. Geophila orbicularis var. neblinae

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Key to the Varieties of G. orbicularis 1. Leaf blades ovate or rarely suborbicularovate, 2.5–3.5 × 2.5–3.5 cm, usually longer than broad, the upper surface glabrous, usually acutely obtuse at the apex, with the inner portion of the basal lobes not or only a little prolonged toward the center of the sinus ................... ....................................... var. neblinae 1. Leaf blades reniform or suborbicular to suborbicular-ovate, 1.2–3 × 1–3 cm, generally broader than long or as broad as long, the upper surface subappressedpubescent becoming glabrescent, usually rounded at the mucronate apex, with the inner portion of the basal lobes prolonged toward the center of the sinus ................................... var. orbicularis G. orbicularis var. neblinae Steyerm., Mem. New York Bot. Gard. 23: 398. 1972. Tepui slope forests, 100–700 m; Amazonas (Sierra de la Neblina). Endemic. ◆Fig. 479. G. orbicularis var. orbicularis. —Ají de Morrocoy. Lowland to upland forests, 100–1200 m; Bolívar (Guaiquinima, near Ikabarú and El Paují, Macizo del Chimantá), Amazonas (Maroa, San Carlos de Río Negro, Sierra de la Neblina, Yavita). Amazonian Brazil.

Geophila repens (L.) I.M. Johnst., Sargentia 8: 281. 1949. —Rondeletia repens L., Syst. Nat. ed. 10. 928. 1759. Plant glabrescent; leaves 1.2–5.5 × 1.2–5 cm; stipules 1.5–2 mm long; peduncles 0.5– 6.5 cm long; calyx limb 2–3.5 mm long; corolla tube 7–9 mm long, lobes 2–5 mm long; fruit 9–10 mm diameter, red. Evergreen lowland forests, riparian forests, 50–500 m; Delta Amacuro (Binojoiche, Sacupana), Bolívar (base of Altiplanicie de Nuria, La Guanajuña peak near Mereware), Amazonas (foot of Cerro Duida, near San Fernando). Anzóategui, Apure, Barinas, Miranda, Monagas, Yaracuy, Zulia; Mexico, Central America, West Indies, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina, Africa, Madagascar, Asia. ◆Fig. 478. Geophila tenuis (Müll. Arg.) Standl., Publ. Field Columbian Mus., Bot. Ser. 7: 77. 1930. —Mapouria tenuis Müll. Arg. in Mart., Fl. Bras. 6(5): 425. 1881. —Ají de morrocoy. Plant hirsute to glabrescent; leaves 1–4 × 0.6–3 cm; stipules 1.8–2.5 mm long; peduncles 0.5–2 cm long; calyx lobes 3.5–4 mm long; corolla tube ca. 7 mm long, lobes 2–2.5 mm long; fruit 6–7 × 5 mm, red. Evergreen lowland and riparian forests, 100–200 m; Amazonas (Río Autana, Río Casiquiare, Río Guainía, Río Negro). Guyana, Suriname, French Guiana, northeastern Brazil.

38. GLEASONIA Standl., Publ. Field Columbian Mus., Bot. Ser. 7: 372. 1931. by Julian A. Steyermark and Charlotte M. Taylor Small trees or shrubs, terrestrial, unarmed. Leaves opposite, petiolate, venation not lineolate; stipules persistent to deciduous, calyptrate (i.e., fused into a conical cap) later splitting into two intrapetiolar triangular segments. Inflorescence terminal, paniculate, multiflowered, bracteate. Flowers pedicellate, large, showy, homostylous, protandrous, fragrant. Hypanthium turbinate to subglobose. Calyx deeply lobed, lobes 5, all expanded into petaloid calycophylls, these ligulate-spathulate, white, sometimes flushed roseate basally; corolla funnelform and somewhat zygomorphic, roseate, internally tomentose to villous on tube and lobes, lobes 5, in bud imbricate with one lobe external. Stamens 5, inserted near middle of corolla tube; anthers exserted, narrowly oblong, dorsifixed; filaments somewhat unequal, pubescent. Ovary 2-locular; ovules numerous in each locule, on axile placentas. Fruit capsular, suborbicular to obconic, laterally somewhat compressed parallel to septum, woody, loculicidal generally along the margins, smooth to verrucose. Seeds large, laterally somewhat compressed, angulate. Venezuela, Brazil; 5 species, 1 in Venezuela, restricted to the flora area.

Gleasonia 611

Fig. 480. Gleasonia duidana var. duidana

The five expanded petaloid calyx lobes and stipules of Gleasonia flowers are distinctive. Similar showy, funnelform, slightly zygomorphic corollas are found in Coutarea and Henriquezia; the distinctions among these are outlined under Henriquezia. Gleasonia duidana Standl., Publ. Field Columbian Mus., Bot. Ser. 7: 372. 1931. Shrub or tree to 30 m tall; leaves 4–28 × 4–15 cm; stipules 1.2–3.3 cm long; inflorescences 9–21 × 7–28 cm; calyx lobes 18–60 × 3–15 mm; corolla 2–4.8 cm long; capsules 1.3–3 × 1.5 –3.5 cm. Tepui slopes and summits, 900–1800 m. Venezuela; 2 varieties. Endemic. Key to the Varieties of G. duidana 1. Leaf blades 4–12.5 cm wide, with the apex obtuse to acuminate and the tertiary venation plane to somewhat impressed on each surface; tree to 30 m tall ................. ....................................... var. duidana 1. Leaf blades 12–15 cm wide, with the apex rounded to truncate and the tertiary ve-

nation raised on each surface; shrub or tree to 15 m tall ............. var. latifolia G. duidana var. duidana Gleasonia duidana var. oblanceolata Steyerm., Mem. New York Bot. Gard. 17: 239. 1967. Bolívar (Cerro Jaua, Cerro Sarisariñama, Gran Sabana near Santa Elena, Macizo del Chimantá, Serranía Marutaní, Sierra Pakaraima), Amazonas (Cerro Aracamuni, Cerro Camani, Cerro Duida, Cerro Huachamacari, Cerro Sipapo, Cerro Yapacana, Cerro Yutajé, Sierra de la Neblina). Endemic. ◆Fig. 480. G. duidana var. latifolia Steyerm., Mem. New York Bot. Gard. 17: 239. 1967. Amazonas (Cerro Coro Coro, Cerro Guanay, Cerro Sipapo, Cerro Yutajé). Endemic.

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39. GONZALAGUNIA Ruiz & Pav., Fl. Peruv. Prodr. 12, t. 3. 1794. by Charlotte M. Taylor and Julian A. Steyermark Small trees or shrubs, terrestrial, unarmed. Leaves opposite, subsessile to petiolate, venation not lineolate; stipules interpetiolar and often shortly united intrapetiolarly, persistent, generally imbricate to valvate in bud, triangular to subulate. Inflorescence terminal, pedunculate, multiflowered, spiciform and elongated, bracteate. Flowers sessile to pedicellate, borne separately or in small cymules directly from the primary axis, medium-sized, distylous, protandrous, fragrant. Hypanthium turbinate to subglobose. Calyx limb lobed often deeply, lobes 4, equal or unequal, without calycophylls; corolla salverform, white to pink or red, internally villous in upper part of tube and throat, lobes 4, imbricate in bud. Stamens 4, inserted in the corolla tube; anthers narrowly oblong, included or partially exserted, dorsifixed. Ovary 2- or 4-locular; ovules numerous in each locule, on axile placentas. Fruits subglobose, heteromorphic, either drupaceous, fleshy to spongy, and white or blue to black, or schizocarpous, coriaceous, and green; pyrenes 2 or 4, 1-locular, each containing numerous dry seeds. Seeds small, angled. Mexico, Central America, Antilles, Colombia, Venezuela, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; ca. 35 species, 6 species in Venezuela, 2 of these in the flora area. Gonzalagunia’s elongated narrow inflorescences and infructescences are frequently pendulous at the ends of the branches, giving the plants a distinctive appearance. Steyermark distinguished the Gonzalagunia species of the flora area primarily by patterns of pubescence. This approach is followed here provisionally, although the variation in their reproductive characteristics is notable and apparently overlaps. Thus, the status of these species and the variation in these characters are in need of study in living populations. Gonzalagunia spicata (Lam.) M. Gomez is occasionally found in northern Venezuela, and has been reported by Steyermark (1972) from Guyana and also appears to be represented by a specimen (Delprete et al. 7077, MO) from Suriname. This species is distinguished from the two species treated below by its calyx tubes 10–11 mm long and calyx limbs 1.5–3 mm long. Key to the Species of Gonzalagunia 1.

1.

Lower surfaces of leaves sparsely to moderately pubescent with both persistent strigillose trichomes and persistent to caducous, silky-sericeous or silky-hirtellous trichomes; corolla tube 4.5–8 mm long ...................................................................................................... G. dicocca Lower surface of leaves moderately to densely hirtellous with generally only one type of trichome; corolla tube 6–8 mm long ...... G. surinamensis

Gonzalagunia dicocca Cham. & Schltdl., Linnaea 4: 194. 1829. Gonzalagunia dicocca subsp. dicocca var. guianensis Steyerm., Mem. New York Bot. Gard. 23: 315. 1972. Gonzalagunia dicocca subsp. venezuelensis Steyerm., Mem. New York Bot. Gard. 23: 315. 1972. Shrub or tree to 4 m tall; leaves 3–16.5 ×

1–4.5 cm; stipules 3–6 mm long; inflorescences 10–22 cm long; calyx limbs 0.5–1.5 mm long; corolla tubes 4.5–6(–8) mm long, lobes 2–3 mm long; fruits 3–4 mm diameter. Riparian forests, rain forests, flooded forests, 200–800 m; Bolívar (Gran Sabana near Santa Elena de Uairén, eastern portion west to Río Paragua). Apure, Aragua, Carabobo, Cojedes, Distrito Federal, Falcón, Miranda,

Gonzalagunia 613

Fig. 481. Gonzalagunia surinamensis

Sucre, Táchira, Yaracuy, Zulia; Trinidad, Guyana, Suriname, French Guiana, Ecuador, Brazil. Steyermark recognized two sympatric subspecies of Gonzalagunia dicocca: subsp. venezuelensis Steyerm. was distinguished by its corollas 9–10.5 mm long with their lobes abaxially sparsely strigillose, and was reported from northern to southern Venezuela; subsp. dicocca was distinguished by its corollas 6–8.5 mm long with their lobes abaxially silky-sericeous, and was reported from throughout the range of the species. Steyermark also gave weight to variation in the pubescence of the abaxial leaf surface to distinguish these subspecies, but with more material now available, this character is clearly continuously variable and not informative; corolla length also varies more widely and continuously than noted by Steyermark, and shows a weaker correspondence with pubescence characters than he thought. Consequently, these subspecies are not separable. Within subsp. dicocca Steyermark recognized two varieties: var. dicocca was reported from northern to southern Brazil and distinguished by its corolla tubes that are externally glabrous in their basal 3 mm; var. guianensis was reported from Venezuela, the Guianas, and northern Brazil and distin-

guished by its corolla tubes that are externally glabrous only in their basal 1–1.5 mm. However this character also varies more widely than Steyermark noted; for example, in southern Brazil the corollas are often externally glabrous in only their 1.5 basal mm (e.g., Hatschbach & Barbosa 56985, MO) while in the Guianas they may be glabrous in the basal 2–2.2 mm (e.g., Clarke et al. 5994, MO). Therefore these varieties are not separable. Steyermark also tried to use the length of the calyx lobes to distinguish infraspecific taxa, but these appear to vary markedly, from uniformly rounded and 0.5 mm long to unequally shaped and ranging from 0.8–1.5 mm long on a single flower. Similar variation is seen in calyx limbs of other species of Gonzalagunia [e.g., G. cornifolia (Kunth) Standl. fide Andersson and Ståhl 1999]. Gonzalagunia surinamensis Bremek., Recueil Trav. Bot. Néerl. 31: 264. 1934. Shrub to 2 m tall; leaves 2.5–18 × 1.5–5.5 cm; stipules 4–8 mm long; inflorescences 6– 20 cm long; calyx limbs 0.5–1 mm long; corolla tubes 6–8 mm long, the lobes 2–3 mm long; fruits 3.5–5 mm diameter. Semideciduous forests, 200–500 m; Delta Amacuro (Río Toro, Serranía de Imataca). Guyana, Suriname, French Guiana, Brazil (Roraima). ◆Fig. 481.

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40. GUETTARDA L., Sp. Pl. 991. 1753. by Charlotte M. Taylor and Julian A. Steyermark Shrubs or trees, terrestrial, unarmed or the branches sometimes with short spinescent stems. Leaves opposite or rarely ternate, petiolate, venation sometimes sublineolate; stipules persistent or caducous, interpetiolar or sometimes connate at the base, triangular, twisted in bud. Inflorescence axillary, several-flowered to multiflowered, cymose with the axes frequently markedly dichasial or scorpioid, bracteate or the bracts reduced. Flowers sessile to shortly pedicellate, mediumsized to rather large, fragrant, distylous and protrandrous or perhaps sometimes homostylous (rarely polygamo-dioecious in plants from outside the flora area). Hypanthium turbinate to ellipsoid. Calyx limb truncate, undulate, or (2–)5 or 6(–19)lobed, without calycophylls; corolla salverform to tubular, white to cream, yellow, pink, red, or purple, internally glabrous, the lobes (4)5 or 6(9), in bud imbricate usually quincuncially, the margins frequently crisped, irregularly undulate, or appendaged. Stamens (4)5 or 6(9), inserted in the corolla throat; anthers dorsifixed near middle or base, sessile or subsessile, included or exserted. Ovary 2–9-locular; ovules solitary in each locule, pendulous from its apex. Fruit drupaceous, subglobose to ellipsoid, cylindrical, or quadrangular, fleshy, red to purple or black; pyrene 1, 2–9-locular, smooth or longitudinally ridged. Seeds small, cylindrical. Mexico, Central America, Antilles, Colombia, Venezuela, Trinidad, Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina, Uruguay, Pacific Islands, Madagascar, coastal East Africa; ca. 80 species, 14 in Venezuela (2 of them restricted to the Caribbean islands), 5 of these in the flora area. Guettarda is easily confused with Chomelia; their separation is discussed under Chomelia. The higher-order leaf venation of many species of Guettarda is quite regularly ordered in squares or short rectangles; this is the arrangement here called “sublineolate.” The higher-order leaf venation of other species of Guettarda is irregularly arranged; this form is here called “areolate.” Taylor and Lorence (2001) discussed the characteristics of this genus in more detail and reviewed some of the Guettarda species found in Venezuela, though outside the flora area. Key to the Species of Guettarda 1. 1. 2(1). 2. 3(1). 3. 4(3). 4. 5(3). 5.

Leaves deltoid to obtuse or rounded at apex ............................................ 2 Leaves acute to acuminate at apex ........................................................... 3 Peduncles 20–30 mm long; leaves 7–13 × 3–7.5 cm, with secondary veins 5–7 pairs; floral bracts to 1.5 mm long .................................. G. divaricata Peduncles 4–10 mm long; leaves 3.5–12 × 1.7–7 cm, with secondary veins 7–13 pairs; floral bracts 2–3.5 mm long .......................... G. malacophylla Leaves 7–13 × 3–7.5 cm, with secondary veins 5–8 pairs ........................ 4 Leaves 11–22 × 6–12 cm, with secondary veins 8–10 pairs ..................... 5 Inflorescences with axes well developed and strongly scorpioid; corolla tubes slender; fruits cylindrical ................................................ G. acreana Inflorescences with axes short and usually strongly dichasial; corolla tubes rather stout; fruits subglobose ..................................... G. divaricata Corollas 55–75 mm long ........................................................... G. macrantha Corollas 18–22 mm long ............................................................. G. spruceana

Guettarda 615

Guettarda acreana K. Krause, Notizbl. Königl. Bot. Gart. Berlin 6: 204. 1914. —Cafecillo, Punteral negro. Guettarda ulei K. Krause, Notizbl. Königl. Bot. Gart. Berlin 6: 203. 1914. Antirhea surinamensis Bremek., Acta Bot. Neerl. 8: 479. 1959. Guettarda leiantha Steyerm., Ann. Missouri Bot. Gard. 71: 1175. 1984. Tree to 20 m tall; leaves 7–13 × 3–7.5 cm; secondary veins 5–8 pairs; higher-order venation areolate; petioles 5–20 mm long; stipules 2–5 mm long; peduncles 3–10 cm long; inflorescence principal axes 2 or 3, well developed, dichasial at first node, scorpioid and strongly secund at subsequent nodes; calyx limb ca. 0.5 mm long, subtruncate; corolla greenish yellow to greenish white, externally glabrescent, tube 7–10 mm long, lobes 1.5–2 mm long; fruits cylindrical to ellipsoid, 8–9 × 4–5 mm, black. Semideciduous to evergreen lowland forests, 100–400 m; Delta Amacuro (Cerro La Paloma, Serranía de Imataca), eastern Bolívar, Amazonas (southeast of Puerto Ayacucho). Panama, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil. Guettarda acreana is unusual and distinctive in its general aspect within Guettarda. Bremekamp (1959) considered it to belong to the poorly known neotropical Antirhea Comm. ex Juss. (= Stenostomum C.F. Gaertn.), and this conclusion may eventually be supported. Steyermark provisionally considered G. acreana distinct from G. ulei of Peru and G. fanshawei Steyerm. of Guyana, but with more specimens now available from Peru, G. ulei cannot be separated any more and is here considered synonymous. Guettarda fanshawei has not been adequately documented for a new conclusion about its status. Guettarda divaricata (Humb. & Bonpl. ex Roem. & Schult.) Standl., Publ. Field Columbian Mus., Bot. Ser. 7: 403. 1931. —Dicrobotryum divaricatum Humb. & Bonpl. ex Roem. & Schult., Syst. Veg. 5: 221. 1819. —Cacho de bagre, Jasmín de monte, Punteral negro, Sabañon. Shrub or tree to 7.5 m tall; leaves 7–13 × 3–7.5 cm; secondary veins 5–7 pairs; higherorder venation sublineolate; petioles 2–20 mm long; stipules 2–5 mm long; peduncles 2– 3 cm long; inflorescence principal axes 2, dichasial at first node, relatively short and

dichasial at subsequent nodes; calyx limb 1.5–2 mm long, undulate to bilobed; corolla white to pink, externally densely sericeous, tube 8–17 mm long, lobes 2.5–5 mm long; fruits subglobose, 5–7 mm diameter, blueblack to black. Riparian and gallery forests, dry thickets, savanna borders, 50–400 m; Bolívar (eastern part west to Río Paragua), Amazonas (upper Río Orinoco). Common elsewhere in Venezuela; Colombia, Trinidad, Tobago, Guyana, Brazil (Roraima). ◆Fig. 482. Guettarda divaricata is very similar and probably closely related to the Caribbean species G. odorata (Jacq.) Lam. and G. elliptica Sw., and in general replaces these in continental South America. Not surprisingly, these three species have been confused and sometimes synonymized by some authors. Their distinctions are clarified by Steyermark (1972, 357–371). Guettarda macrantha Benth., J. Bot. (Hooker) 3: 223. 1841. Shrub or tree to 8 m tall; leaves 11–22 × 6– 12 cm; secondary veins 9–12 pairs, higher-order venation sublineolate; petioles 1–4.5 cm long; stipules 3–8 mm long; peduncles 5–13 cm long; inflorescences congested-cymose, the principal axes 2, relatively short, dichasial; calyx limb 2–3 mm long, truncate to undulate; corolla cream-colored to white, externally densely sericeous, tube 30–55 mm long, lobes 4–5 mm long; fruits subglobose, 10–15 mm diameter, perhaps yellow. Rain forests, riparian forests, 100–200 m; Amazonas (Río Ocamo). Guyana, Suriname, French Guiana, northeast Brazil. Guettarda argentea Lam., based on a specimen from “Ile de Cayenne,” French Guiana, may be an older name for this species, but adequate material to confirm this has not been seen. Guettarda malacophylla Standl., Publ. Field Columbian Mus., Bot. Ser. 4: 200. 1929. Small tree or shrub to 8 m tall; leaves 3.5– 12 × 1.7–7 cm; secondary veins 7–13 pairs, higher-order venation sublineolate; petioles 1.5–16 mm long; stipules 3–6 mm long; peduncles 4–10 mm long; inflorescences subcapitate or dichasial with principal axes 2, short; calyx limb 2–2.5 mm long; corollas white, externally densely sericeous, tube 4.5–10 mm long; lobes 1.5–4 mm long; fruits

616

R UBIACEAE

Fig. 482. Guettarda divaricata

subglobose, 5–6 mm diameter, red. Deciduous forests, 100–400 m; Bolívar (vicinity of Upata and Represa Guri). Colombia. Guettarda malacophylla is apparently deciduous and produces the flowers together with the new leaves. Guettarda spruceana Müll. Arg., Flora 58: 449. 1875. Shrub or tree to 15 m tall; leaves 11–20 × 6–12 cm; secondary veins 8 or 9 pairs,

higher-order venation sublineolate; petioles 1–6 cm long; stipules 3–5 mm long; peduncles 5–8 cm long; inflorescence principal axes 2, dichasial or scorpioid at the distal nodes; calyx limb ca. 3 mm long, truncate; corollas cream-yellow, externally densely sericeous, tube 10–15 mm long, lobes ca. 4 mm long; fruits subglobose, 10–12 mm diameter, color unknown. Semideciduous forests, 400– 600 m; northwestern Bolívar (near Cerro Cerbatana). Suriname, Brazil.

41. HAMELIA Jacq., Enum. Syst. Pl. 2: 1760. by Charlotte M. Taylor and Julian A. Steyermark Shrubs or small trees, terrestrial, unarmed. Leaves opposite or verticillate, subsessile to petiolate, venation not lineolate; stipules interpetiolar, triangular, generally imbricate or valvate in bud, deciduous. Inflorescence terminal, multiflowered, cymose, bracteate, the axes usually helicoid and secund. Flowers rather large, sessile, diurnal, homostylous, protandrous. Hypanthium turbinate to ellipsoid. Calyx limb reduced, 5-dentate, without calycophylls; corolla tubular or funnelform and sometimes ventricose, yellow, orange, or red, internally glabrous, lobes 5, imbricate in bud. Stamens 5, inserted at the base of the corolla tube; anthers narrowly oblong, included or partially exserted, basifixed, shortly sagittate at base, shortly appendiculate at apex. Ovary 4- or 5-locular; ovules numerous on axile placentas. Fruit baccate, fleshy, subglobose to ellipsoid, black. Seeds small, irregularly angled to lenticular. Southeastern U.S.A., Mexico, Central America, West Indies, Colombia, Venezuela, Trinidad, Ecuador, Peru, Brazil, Bolivia, Argentina; ca. 16 species, 2 in Venezuela, both in the flora area. Hamelia is sometimes confused with Palicourea, which can be distinguished from Hamelia by its drupaceous fruits with the seeds 1 per locule and its bilobed stipules. Hamelia is also sometimes confused with Bertiera; their separation is outlined under Bertiera.

Henriquezia 617

Fig. 483. Hamelia axillaris

Key to the Species of Hamelia 1. 1.

Corolla yellow, ventricose to funnelform, with tube 9–15 mm long; leaves opposite or infrequently 3- or 4-verticillate ............................. H. axillaris Corolla reddish orange, tubular, with tube 10–20 mm long; leaves 3–5verticillate or occasionally opposite ............................................. H. patens

Hamelia axillaris Sw., Prodr. 46. 1788. Hamelia lutea Rohr ex Sm. in Rees, Cycl. 17: no. 4. 1811. Shrub 1–2 m tall; leaves 5–23 × 2–8 cm; stipules 2–6 mm long; inflorescences 3–8 × 3–8 cm; calyx lobes 1–2 mm long; corolla lobes 1–2 mm long; fruits 4–7 × 3–4 mm. Lowland forests, usually in shade, 50–200 m; Amazonas (Quebrada Chiruire). Apure, Aragua, Carabobo, Zulia; Mexico, Central America, West Indies, Colombia, Ecuador, Peru, Brazil. ◆Fig. 483. Hamelia patens Jacq., Enum. Syst. Pl. 16. 1760. Shrub to tree 1–4 m tall; leaves 5–23 × 2–8 cm; stipules 2–6 mm long; inflorescences 3– 15 × 5–20 cm; calyx lobes ca. 1 mm long; co-

rolla lobes 1–2.5 mm long; fruits 6–8 × 3–4 mm. Riparian and gallery forests, deciduous forests, usually in edges and disturbed sites, 50–700 m; Delta Amacuro (Río Acure), Bolívar (eastern part west to Río Paragua), Amazonas (upper Río Orinoco). Southern U.S.A., Mexico, Central America, Colombia, Venezuela, Trinidad, Guyana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina. Hamelia patens is a widespread, morphologically variable species. Elias (1976) and Steyermark (1974) recognized two varieties: var. patens, with the vegetative and reproductive structures pubescent, and var. glabra Oerst., with the leaves usually glabrous adaxially and the reproductive structures glabrous to pubescent. However, these are not being recognized here.

42. HENRIQUEZIA Spruce ex Benth., Hooker’s J. Bot. Kew Gard. Misc. 6: 338, 1854. by Julian A. Steyermark and Charlotte M. Taylor Shrubs to large trees, unarmed, terrestrial. Leaves 3–6-verticillate, petiolate, venation not lineolate; petioles at abaxial base usually with 1 or 2 orbicular to oblong glands; stipules interpetiolar but deeply lobed and apparently free, generally valvate in bud, persistent, with the segments linear to narrowly deltoid. Inflores-

618

R UBIACEAE

cence terminal, paniculate to cymose, multiflowered, bracteate. Flowers large, showy, homostylous, subsessile to pedicellate, protandrous. Hypanthium turbinate. Calyx limb 4-lobed, the lobes unequal but without calycophylls; corolla broadly funnelform and somewhat zygomorphic, roseate without, white with rose-red stripes and a yellow bearded zone within, the throat with a pubescent ring, lobes 5, in bud imbricate with one lobe external. Stamens 5, unequal, 2 of them inserted lower than the others; anthers narrowly elliptic, dorsifixed, included. Ovary 2-locular, partially inferior; ovules 4 per locule, on axile placentas. Capsule woody, partially inferior, oblong, strongly laterally flattened parallel to the septum, loculicidally dehiscent along the margins, with a circumscissile scar from the deciduous calyx limb across the middle. Seeds flat though unwinged, large, papillose. Colombia, Venezuela, Guyana, Brazil; 3 species, 2 in Venezuela, both in the flora area. The third species recognized in this genus, Henriquezia jenmannii K. Schum., is known only from along the Río Mazaruni in Guyana in a range quite distinct from the other species of Henriquezia. Henriquezia is notable for its ovary position, almost fully inferior in flower but becoming partially inferior in fruit, and its zygomorphic corollas. Platycarpum, Gleasonia, and Coutarea have similar corollas. Gleasonia differs from these genera in its enlarged, spatulate to oblanceolate calyx lobes; Coutarea differs from the remaining genera in its fully inferior fruits and ovaries. Platycarpum and Henriquezia both have partially inferior ovaries and similar capsules, but Platycarpum has calyptrate stipules while those of Henriquezia are deeply bilobed. The stipules of Isertia are similar to those of Henriquezia, but Isertia differs from this genus in its salverform corollas and smaller fleshy fruits. Key to the Species of Henriquezia 1.

1.

Leaf secondary veins plane adaxially and plane to prominulous abaxially; capsules with the calyx scar generally convex-curved across each side ........................................................................................ H. nitida Leaf secondary veins plane to impressed adaxially, prominent abaxially; capsules with the calyx scar generally straight across each side ............................................................................................... H. verticillata

Henriquezia nitida Spruce ex Benth., Trans. Linn. Soc. London 22: 297, t. 54. 1859. Colombia, Venezuela, Brazil; 3 varieties, 1 in Venezuela. A sterile specimen (Liesner 8531, MO, Amazonas, San Carlos de Río Negro) appears to represent H. nitida var. macrophylla (Ducke) Steyerm. This is otherwise known from the lower Rio Negro in Brazil and is distinguished from H. nitida var. nitida by its oblanceolate leaves that are obtuse to acute at the apex, versus elliptic to oblong and rounded at apex in var. nitida. H. nitida var. nitida. —Cachete de vieja, Cartera, Guaco, Pastora.

Henriquezia oblonga Spruce ex Benth., Trans. Linn. Soc. London 22: 297. 1859. —Henriquezia nitida var. oblonga (Spruce ex Benth.) Steyerm., Mem. New York Bot. Gard. 10: 207. 1963. Tree to 20 m tall; leaves 3- or 4-verticillate, 8.5–31 × 2–14 cm; stipules 1–3.5 cm long; inflorescences 8–24 × 6–23 cm; calyx lobes 7–25 mm long; corolla with tube 2.5–5 cm long, lobes 1–2 cm long; capsules 8–10 × 10–11 cm; seeds 3–4 cm long. White-sand savannas, periodically flooded dwarf forests, banks of black-water rivers, 50–100 m; Amazonas (basin of Río Casiquiare, Río Negro, and Río Orinoco). Southeastern Colombia, northwestern Brazil. ◆Fig. 484.

Hillia 619

Fig. 484. Henriquezia nitida var. nitida

Henriquezia verticillata Spruce ex Benth., Hooker’s J. Bot. Kew Gard. Misc. 6: 338. 1854. —Boca de sapo, Guaco. Tree to 40 m tall; leaves 4–6-verticillate, 9–67 × 3–20 cm; stipules 1–8 cm long; inflorescences 11–35 × 9–24 cm; calyx lobes 2–9 mm long; corolla with tube 2–4 cm long, lobes

0.5–2 cm long; capsules 6–8 × 8–11.5 cm; seeds 4.5–5 cm long. Edges of periodically flooded, black-water rivers, 100–200 m; Amazonas (Maroa, along Río Guainía above confluence with Río Casiquiare, San Carlos). Brazil (Amazonas: Rio Negro).

43. HILLIA Jacq., Enum. Syst. Pl. 3. 1760. by Charlotte M. Taylor and Julian A. Steyermark Succulent epiphytic shrubs or small trees, unarmed, glabrous. Leaves opposite, petiolate to subsessile, venation not lineolate and often not visible; stipules interpetiolar, ligulate to oblanceolate, erect and flatly appressed, caducous. Inflorescences terminal, sometimes bracteate, 1(3)-flowered. Flowers large, showy, homostylous, protandrous, diurnal or nocturnal, sometimes fragrant, sessile or pedunculate. Hypanthium cylindrical to ellipsoid. Calyx limb none or deeply 4–10-

620

R UBIACEAE

lobed, without calycophylls; corolla salverform or funnelform to ventricose, red, green, or white, glabrous internally and externally, lobes 4–10, convolute in bud. Stamens 4–7, inserted near top or middle of corolla tube; anthers narrowly oblong, included, dorsifixed near base. Ovary 2-locular; ovules numerous in each locule, on axile placentas. Fruit capsular, cylindric or narrowly oblong, woody, smooth to longitudinally ridged, septicidally dehiscent from the apex. Seeds papery, small, flattened, fusiform, marginally winged and with a tuft of long filaments at the apex. Mexico, Central America, West Indies, Colombia, Venezuela, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 24 species, 7 in Venezuela, 5 of these in the flora area. Hillia is similar to Cosmibuena, which can be distinguished from Hillia by inflorescences with usually several flowers, corolla lobes that are imbricated in bud, and its seeds that lack a tuft of trichomes. Hillia is also similar vegetatively to Schradera, which can be distinguished by its capitate involucrate inflorescences, fleshy fruits, and stems often with regularly produced adventitious roots. Hillia was reviewed by Taylor (1994). Hillia ulei K. Krause has been collected widely but infrequently in Amazonia and the Guianas and may be expected in the flora area. It is distinguished by its subpalmate leaf venation, with all the secondary veins arising below the middle of the blade; all of the other species in the flora area have pinnate venation. Flowers of Hillia ulei are similar to those of H. illustris. Key to the Species of Hillia 1. 1. 2(1). 2. 3(2). 3. 4(3). 4.

Corolla tube funnelform; corolla green, yellow-green, dull red, or gray-roseate; calyx lobes 9–36 mm long ........................................................... 2 Corolla tube narrowly salverform; corolla white; calyx limb absent or deeply lobed, the lobes 3–12 mm long ................................... H. parasitica Leaf blades rounded to obtuse at apex.......................................... H. foldatsii Leaf blades acute to acuminate at apex .................................................... 3 Leaf blades 0.5–1 cm wide; corolla externally dull red; on rocks in or beside rivers .................................................................................. H. rivalis Leaf blades 2–8 cm wide; corolla externally green to yellow-green; epiphytic ...................................................................................................... 4 Leaf blades 8.5–15 cm long ............................................................ H. illustris Leaf blades 4.5–6.5 cm long .................................................. H. psammophila

Hillia foldatsii Steyerm., Mem. New York Bot. Gard. 23: 290. 1972. Fleshy shrub ca. 1 m tall; leaves 3–6.4 × 1.5–3.5 cm; stipules 9–12 mm long; calyx lobes 6, 13–17 mm long; corolla funnelform, green flushed with pink, the tube 35–60 mm long, the lobes 6, 7–9 mm; capsules 6.5–9 × 0.9–1.2 cm. Tepui scrub, ca. 1200 m; Bolívar (Auyán-tepui). Endemic. Hillia illustris (Vell.) K. Schum. in Mart., Fl. Bras. 6(6): 202. 1889. —Saldanha illustris Vell., Fl. Flumin. 3: 141, t. 157. 1825 [1829].

Hillia tubiflora Cham., Linnaea 9: 260. 1834, “tubaeflora.” Fleshy epiphytic shrub or tree to 8 m tall; leaves 8.5–15 × 3–8 cm; stipules 23–33 mm long; calyx lobes 6, 9–35 mm long; corolla funnelform, bright green to yellow-green, the tube 48–61 mm long, the lobes 6, 8–16 mm long; capsules 5–11 × 0.8–1.5 cm. Semideciduous to evergreen lowland forests, 50– 500 m; Delta Amacuro (Río Acure, Río Cuyubini), Bolívar (Altiplanicie de Nuria, Sierra Imataca). Anzoátegui; Colombia, Guyana, Suriname, French Guiana, Ecuador, Amazonian Peru, Brazil, Bolivia. ◆Fig. 486.

Hillia 621

Hillia parasitica Jacq., Enum. Syst. Pl. 18. 1760. Fleshy epiphytic shrub, subshrub, or small tree; leaves 3.4–13 × 1.7–6.5 cm; stipules 9–55 mm long; calyx limb absent or with lobes 6, 3– 12 mm long; corolla salverform, white becoming yellow with age, the tube 6–12.5 cm long, the lobes 6, 1.5–6 cm long; capsules 3–11.5 (–20) × 0.6–1 cm. Tepui slope and summit forests, 1000–2200 m; Bolívar (Auyán-tepui,

Fig. 485. Hillia rivalis

Macizo del Chimantá, Ptari-tepui, basin of Río Cuyuní, Uei-tepui), Amazonas (Cerro Duida, Cerro Sipapo). Aragua, Falcón, Lara, Mérida, Miranda, Monagas, Nueva Esparta, Portuguesa, Yaracuy; widespread in Mexico, Central America, West Indies, Colombia, Trinidad, Guyana, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 487. Steyermark (1972) recognized two varieties, var. nobilis (Vell.) Steyerm. of “southern

Fig. 486. Hillia illustris

622

R UBIACEAE

Fig. 487. Hillia parasitica

Brazil, Bolivia, Colombia, and possibly Peru,” and var. parasitica found in the remaining distribution of this species. However with more collections now available, the differences in corolla size, bract size, and bract shape he used to separate these can be seen to vary continuously and independently; consequently these varieties are not recognized here. Hillia psammophila Steyerm., Mem. New York Bot. Gard. 19(5): 211, fig. 74. 1963. Fleshy climbing epiphyte; leaves 4.5–6.5 × 2–3 cm; stipules not seen; calyx lobes 6, ca. 18 mm long; corolla funnelform, yellow-

green, the tube ca. 6 cm long, the lobes 6, ca. 1 cm long; capsules not seen. Tepui slope and summit forests, 700–1100 m; Bolívar (Cerro Guaiquinima, Cerro Uei). Endemic. Hillia rivalis C.M. Taylor, Ann. Missouri Bot. Gard. 78: 521, fig. 1. 1991. Shrub to 1 m tall; leaves 3.8–10 × 0.5–1 cm; stipules 10–13 mm long; calyx lobes 6, 18–36 mm long; corolla funnelform, dull red, the tube 3.2–5.9 cm long, the lobes 6, 0.8–1 cm long; capsules 5–6.5 × 0.8–1 cm. Rocks along rivers in tepui slope areas, 1200–1300 m; Amazonas (Cerro Marahuaca). Endemic. ◆Fig. 485.

Holstianthus 623

44. HOLSTIANTHUS Steyerm., Ann. Missouri Bot. Gard. 73: 495, fig. 1. 1986. by Charlotte M. Taylor and Julian A. Steyermark Shrubs, unarmed, terrestrial. Leaves opposite, petiolate, with secondary and higher order venation not visible; stipules interpetiolar, persistent, generally imbricate to valvate in bud, narrowly triangular, acuminate. Inflorescence axillary and perhaps sometimes terminal, 1–3-flowered, pedunculate, subtended by a pair of foliaceous bracts. Flowers rather large, pedicellate, floral biology unknown. Hypanthium turbinate. Calyx limb deeply lobed, lobes 5, rather large, equal or a little unequal, without calycophylls; corolla funnelform, dull salmon red, yellow-pilose in throat, lobes 5, convolute in bud. Stamens 5, inserted in the upper part of the corolla tube; anthers narrowly oblong, dorsifixed near middle, included. Ovary 2-locular; ovules numerous in each locule, on axile placentas. Mature fruit not seen; fruit apparently capsular, obconic, cartilaginous with bony inner layer, indehiscent. Seeds angled, small, subcompressed. Venezuela; 1 species. Steyermark described the inflorescences as axillary and terminal, based no doubt on one stem on the holotype that apparently terminates in a single inflorescence. All the rest of the inflorescences seen are axillary, borne one per leaf axil and

Fig. 488. Holstianthus barbigularis

624

R UBIACEAE

thus two per node. The one stem apex with the apparently terminal inflorescence appears to be possibly damaged, however, so this character deserves re-examination when more material is available. Also, Steyermark described the fruits as dry and indehiscent with the large calyx lobes persistent on them, but the fruits he studied may not have been mature. Holstianthus barbigularis Steyerm., Ann. Missouri Bot. Gard. 73: 495, fig. 1. 1986. Shrub to 2 m tall; leaves 4.5–11.5 × 1.5– 3.5 cm; stipules 3–7 mm long; peduncles 7–14 mm long, bracts 8–18 × 4–6 mm; calyx lobes 15–25 × 8–9 mm; corolla tube ca. 23 mm long,

lobes ca. 10 mm long; capsules (possibly immature) 7–10 × 7–8 mm. Steep slopes and bluffs of tepui slopes, 1200–1400 m; Amazonas (Cerro Marahuaka, south-central portion above tributary of Caño Negro). Endemic. ◆Fig. 488.

45. ISERTIA Schreb., Gen. Pl. 1: 234. 1789. Cassupa Bonpl. in Humb. & Bonpl., Pl. Aequinoct. 1: 43, pl. 12. 1806. Yutajea Steyerm., Ann. Missouri Bot. Gard. 74: 576. 1987. by Charlotte M. Taylor and Julian A. Steyermark Trees or shrubs, armed, terrestrial. Leaves opposite or ternate, petiolate, sometimes white-pubescent abaxially, venation not lineolate; stipules persistent with the leaves, generally imbricate to valvate in bud, either interpetiolar and deeply bilobed to form apparently 4 free triangular stipules, or intrapetiolar and shortly to deeply bilobed. Inflorescence terminal, multiflowered, bracteate, pedunculate, thyrsoid-paniculate or thyrsiform-racemose, the secondary axes terminating in dichasial or scorpioid cymes. Flowers large, showy, sessile or pedicellate, bibracteolate, homostylous, protandrous, fragrant. Calyx limb reduced, truncate or 4– 6-lobed, without calycophylls; corolla tubular-funnelform to salverform, white or yellow to red, villous in throat and often on the adaxial faces of the lobes, lobes 5 or 6(7), imbricate or valvate in bud. Stamens 4–7, inserted in upper part of corolla tube; anthers included or partially exserted, narrowly oblong, dorsifixed, with the thecae locellate (i.e., internally divided into several small chambers). Ovary 2–6(7)locular; ovules numerous in each locule, on axile placentas. Fruit baccate or drupaceous, subglobose, fleshy, red to purple or black; pyrenes (when present) 4–6, trigonous, 1-locular, containing numerous dry seeds. Seeds small, irregularly angular, foveolate. Central America, Cuba, Lesser Antilles, Colombia, Venezuela, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 14 species, 6 in Venezuela, 4 of these in the flora area. Boom (1984) revised Isertia and recognized two subgenera: subgenus Isertia with drupaceous fruits that generally have 5 or 6 locules, and subgenus Cassupa with baccate fruits that generally have 2 or 3 locules. These groups are united by their locellate anthers, a unique character in the Rubiaceae. Isertia is frequently confused with Palicourea, which can be separated from Isertia by a single ovule and seed per locule. The stipules of Isertia are variable in form, and sometimes so deeply lobed that they appear free and the plants confused with Vochysiaceae. Henriquezia has similar stipules to those of some species of Isertia; the distinctions between these genera are outlined under Henriquezia. Occasionally, individual plants of Isertia have diseased flowers with short tubular corollas. These are sometimes confused with other genera of Rubiaceae. The

Isertia 625

genus Yutajea Steyerm. appears to also be an example of this confusion; a discussion of its synonymization with Isertia is presented under I. parviflora below. Key to the Species of Isertia 1. 1. 2(1).

2.

3(1). 3.

Lower leaf surface covered with a dense white-canescent pubescence ... 2 Lower leaf surface glabrous to pubescent with a sparse colorless pubescence ....................................................................................................... 3 Ovary 5- or 6-locular; stigmas 5 or 6; corolla tube externally densely tomentellose, smooth or shortly verrucose in distal part; hypanthium, calyx, pedicels, bracts, peduncles, and rachis of inflorescence finely tomentose; inflorescence usually as broad as or broader than long, the lateral axes mainly 3–9-flowered ............................................ I. hypoleuca Ovary 2-locular; stigmas 2; corolla tube externally papillose to glabrous, markedly verrucose in distal part; hypanthium, calyx, pedicels, bracts, peduncle, and rachis of inflorescence glabrous or puberulous to papillose; inflorescence usually longer than broad, the lower lateral axes mainly 3-flowered and the upper ones mainly 1-flowered ...... I. verrucosa Stipules deeply divided, at each node apparently 4 and free or shortly interpetiolar; petiole pubescent .............................................. I. parviflora Stipules at each node 2 and intrapetiolar; petiole glabrous or glabrescent .......................................................................................................... I. rosea

Isertia hypoleuca Benth., J. Bot. (Hooker) 3: 220. 1841. —Chaparrillo rebalsero, Manáse-yek (Arekuna). Tree or shrub to 15 m tall; leaves 18–65 × 9–18 cm, elliptic to obovate, white-canescent abaxially; stipules deeply lobed and apparently 4; corolla reddish orange, the tube 43– 65 mm long, the lobes 6 or 7, 8–13 mm long; fruit drupaceous, ca. 10 mm diameter. Evergreen lowland to montane forests, 100–1400 m; Bolívar (Río Caura in El Paují area, mouth of Río Nichare, common in the eastern part at the base of sandstone tepuis and west to upper Río Paragua), Amazonas (Cerro Parú, base of Cerro Sipapo, upper Río Orinoco, Sierra Parima). Colombia, Guyana, Suriname, French Guiana, Peru, Brazil (Parú), Bolivia. ◆Fig. 489. Isertia parviflora Vahl, Eclog. Amer. 2: 28, t. 15. 1798. —Café negro, Carrutillo morichalero. Isertia parviflora var. hirta Steyerm., Mem. New York Bot. Gard. 17(1): 295. 1967. Yutajea liesneri Steyerm., Ann. Missouri Bot. Gard. 74: 576, fig. 1. 1987. Tree or shrub to 15 m tall; leaves 13–39 × 6–17 cm, ovate; stipules deeply lobed and apparently 4; corolla tube 5–13 mm long, pink,

the lobes 6, 4–7 mm long, white and purple; fruit drupaceous, ca. 5 mm diameter. Gallery forests, riparian forests, savannas, forested igneous outcrops, frequently on white-sand substrates, 50–900 m; Delta Amacuro (Cerro La Paloma, Río Cuyubini), Bolívar (Río Parguaza below El Carmen), Amazonas (near Puerto Ayacucho, Río Ocamo, upper Río Orinoco, Río Parucito, Río Ventuari, Serranía de Yutajé). Trinidad, Guyana, Suriname, French Guiana, Brazil. ◆Fig. 492. Steyermark separated Yutajea from Isertia based on its stamens inserted near the base of the corolla tube and with the anthers basifixed. However Andersson (1996) studied a different specimen than the holotype and found the stamens to be inserted near the corolla throat. Based on a cladistic analysis of the tribe Isertieae using a number of morphological characters, Andersson (1996) considered Yutajea to belong to Isertia and commented that its flowers may actually be a teratological form caused by damage from beetle larvae. As noted by Andersson, occasional flowers are found in various species of Isertia with the corolla tube abnormally wide and short, similarly to those of Yutajea, due to such damage. Yutajea seems to be based on a specimen that is similarly damaged, though more extensively than

626

R UBIACEAE

Fig. 489. Isertia hypoleuca

Fig. 490. Isertia rosea

Isertia 627

usual. The figure presented by Steyermark in his original description of Yutajea regularizes the flowers significantly; on the holotype (Liesner & Holst 21826, MO) there is actually notable variation in the overall length of the corollas and the degree of their lobing, which ranges from subtruncate to lobed for more than 3/4 of their length, and in general the in-

florescence appears to be diseased. Vegetatively this specimen is indistinguishable from Isertia parviflora, and it is accordingly synonymized here. Isertia rosea Spruce ex K. Schum. in Mart., Fl. Bras. 6(6): 284. 1889. —Carisillo, Caruto, Mani. Shrub or tree to 13(–20) m tall; leaves 19– 48 × 6–17 cm, obovate to oblanceolate; stipules intrapetiolar; corolla rose-red to pink externally, yellow or pink in throat, the tube 35–50 mm, the lobes 5 or 6, 6–8 mm long; fruits drupaceous, ca. 8.5 mm diameter. Evergreen lowland forests, riparian forests, borders of forests and savannas, evergreen seasonally dry forests, in poorly drained soils, on white-sand substrates, 50–300 m; Amazonas (basins of upper Río Negro and upper Río Orinoco northeast to Río Cunucunuma and Santa Barbara). Apure; Colombia, Ecuador, Peru, Amazonian Brazil. ◆Fig. 490.

Fig. 491. Isertia verrucosa

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R UBIACEAE

Isertia verrucosa (Bonpl.) Standl., Publ. Field Columbian Mus., Bot. Ser. 8(5): 346. 1931. —Cassupa verrucosa Bonpl. in Humb. & Bonpl., Pl. Aequinoct. 1: 43, pl. 12. 1806. —Caruto. Tree to 16 m tall; leaves 26–80 × 12–35 cm, obovate, white-canescent abaxially; stipules deeply lobed and apparently 4; corolla yellow to orange or rose in the upper half and rose-salmon in the lower half, the tube 40–55 mm, white- or yellow-barbate in the throat, the lobes 6, 9–14 mm long; fruit baccate, ca. 9 mm diameter. Evergreen lowland forests and scrub forests on white sand, 100–200 m; Amazonas (basins of upper Río Negro and Río Casiquiare). Adjacent Brazil. ◆Fig. 491.

Fig. 492. Isertia parviflora

46. IXORA L., Syst. Nat. Sp. Pl. 110. 1753. by Charlotte M. Taylor and Julian A. Steyermark Shrubs or small trees, terrestrial, unarmed. Leaves opposite or ternate, sessile to petiolate, venation not lineolate; stipules triangular, generally valvate in bud, acute to acuminate or aristate, generally persistent. Inflorescences terminal or apparently axillary (i.e., borne on very short axillary short-shoots), multiflowered, congested-cymose or subcapitate to fasciculate, cymose, or paniculate, pedunculate to sessile, bracteate. Flowers sessile to pedicellate, small to rather large, distylous, frequently fragrant. Hypanthium turbinate to narrowly ellipsoid. Calyx limb usually with short tube, lobes 4, without calycophylls; corolla salverform, white to

Ixora 629

yellow or red, internally glabrous, the tube generally slender, the lobes 4, convolute in bud. Stamens 4, inserted in corolla throat; anthers narrowly oblong, exserted at least partially, dorsifixed below middle. Ovary 2-locular; ovules 1 in each locule, axile. Fruits subglobose, drupaceous, carnose or coriaceous, red or black; pyrenes 2, chartaceous to rather hard, 1-locular. Seeds subglobose to hemispherical. Central America, West Indies, Colombia, Venezuela, Trinidad-Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Pacific Islands, Africa, Asia, Australia; ca. 400 species, 10 in Venezuela, 6 of these in the flora area. Ixora can be recognized by its petioles articulated at their bases. The inflorescences often have the bracts fused in pairs and the axes articulated, giving them a distinctive appearance. In fruit Ixora is often confused with Psychotria, but it lacks the articulated petioles of Ixora and usually has bilobed stipules. Several Old World species of Ixora are widely cultivated in tropical regions, including the flora area. Ixora coccinea L. can be recognized by its leaves that are sessile to subsessile with their bases truncate to cordate; the flowers of cultivated forms range from red to yellow. Ixora casei Hance is similar to I. coccinea but has petiolate leaves with obtuse to acute bases. Ixora finlaysoniana Wall. ex G. Don (sometimes incorrectly written as “findlaysoniana”) has petiolate leaves, white flowers, and calyx lobes 3–4 mm long. The distinctions between Ixora schomburgkiana, I. ulei, and I. panurensis (as circumscribed by Steyermark) probably deserve careful reconsideration, but this work is beyond the scope of the present project. Key to the Species of Ixora 1. 1. 2(1). 2. 3(2). 3. 4(2). 4. 5(4). 5.

Inflorescences all apparently axillary, with flowers 1–3 per axil, sessile ................................................................................................... I. yavitensis Inflorescences terminal or both terminal and apparently axillary, with flowers several to numerous in cymes or panicles ............................... 2 Leaves subsessile, rounded to subcordate at base, obtuse, or acute ....... 3 Leaves with petioles 4–15 mm long, obtuse to acute at base .................. 4 Leaves opposite or ternate, 10–25 cm long; plants of lowland forests ........................................................................................ I. acuminatissima Leaves opposite, 3–9 cm long; plants on tepui slopes of tepuis ................. ................................................................................................. I. intropilosa Corolla tubes 18–25 mm long ........................................... I. schomburgkiana Corolla tubes 11–15 mm long .................................................................... 5 Flowers mostly or all sessile ....................................................... I. panurensis At least some flowers borne on pedicels 1.5–2 mm long ....................... I. ulei

Ixora acuminatissima Müll. Arg., Flora 58: 454. 1875, emend Steyerm., Mem. New York Bot. Gard. 17: 345. 1967. —Ciutica, Perro de agua, Simayo. Shrub or tree to 3 m tall; leaves ternate and/or opposite, 10–24 × 2–8 cm; petioles 1–3 mm long; stipules 2–5 mm long, often deciduous; inflorescences terminal, paniculate to corymbiform, 2–8 × 3–10 cm, sessile to pedunculate, with numerous flowers; calyx limb ca. 1 mm long; corolla white to pink, red,

or salmon, tube 6–10 mm long, lobes 3–4 mm long, obtuse; fruits 7–8 mm diameter, red becoming black at maturity. Gallery forests, lowland to lower montane forests, 50–700 m; Bolívar (Río Caroní, Río Caura, Río Paragua), Amazonas (common). Apure; Amazonian Colombia and Brazil. ◆Fig. 493. The leaf bases of Ixora acuminatissima are notably variable in shape, ranging from acute to rounded, truncate, or shortly cordate, and the leaf arrangement often varies

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from opposite to ternate even on the same stem. Ixora intropilosa Steyerm., Mem. New York Bot. Gard. 17: 352. 1967. Shrub to 2 m tall; leaves opposite, 3–9 × 1.5–4.5 cm; petioles 1–6 mm long; stipules 3– 6 mm long; inflorescences terminal, 1.5–4 × 2.5–3.5 cm, cymose, pedunculate, with flowers several; calyx limb ca. 1 mm long; corolla red or often yellow on lobes and throat, tube 8–12 mm long, lobes ca. 3.5 mm long, obtuse; fruit 7–7.5 mm diameter. Tepui shrub formations, ca. 1500 m; Bolívar (Kukenán-tepui). Guyana. Ixora panurensis Müll. Arg., Flora 58: 454, 458. 1875. Shrub or tree to 5 m tall; leaves opposite, 10–20 × 5–10 cm; petioles 4–10 mm long; stipules 3–5 mm long; inflorescences terminal, paniculate to congested-cymose, 2–7 × 4–6 cm, sessile or subsessile, with numerous flowers; calyx limb ca. 1 mm long; corolla red or white tinged with pink or red or the lobes sometimes yellow, tube 10–15 mm long, lobes 4–6 mm long, obtuse; fruit 8–10 mm diameter. Evergreen lowland forests, 100–200 m; Amazonas (Pimichín, Yavita). Amazonian Brazil, Ecuador, Peru, Colombia, Bolivia.

Ixora schomburgkiana Benth., Linnaea 23: 448. 1850. Shrub or small tree to 7 m tall; leaves opposite, 12–20 × 5–8.5 cm; petioles 5–15 mm long; stipules 3–8 mm long; inflorescences terminal, paniculate, 5–10 × 5–17 cm, subsessile, with numerous flowers; calyx limb ca. 1 mm long; corolla red or sometimes yellow on lobes and throat, tube 18–25 mm long, lobes 5–6 mm long, obtuse; fruit not seen. Lower montane forests, 500–600 m; Bolívar (basin of Río Cuyuní). Guyana. Ixora ulei Krause, Notizbl. Bot. Gart. Berlin 6: 205. 1914. Shrub or tree to 12 m tall; leaves opposite, 9–15 × 4–6 cm; petioles 3–8 mm long; stipules 3–6 mm long; inflorescences terminal, paniculate, 3–10 × 3–10 cm, sessile to pedunculate, with flowers numerous; calyx limb ca. 1 mm long; corolla red or lobes sometimes yellow, tube 11–15 mm long, lobes 4–6 mm long, obtuse; fruits 8–12 mm diameter. Lower montane forests, 500–800 m; Bolívar (basin of Río Cuyuní). Colombia, Guyana, Suriname, French Guiana, Amazonian Peru, Brazil (Rio Branco). Ixora yavitensis Steyerm., Mem. New York Bot. Gard. 23: 386. 1972. Fig. 493. Ixora acuminatissima

Kutchubaea 631

Tree to 6 m tall; leaves opposite, 11–18 × 3.5–5 cm; petioles 3–5 mm long; stipules 1.5– 3 mm long; inflorescences axillary, with flowers 1–3, sessile; calyx limb ca. 1 mm long; corolla red, tube ca. 11.5 mm long, lobes 3.5–4.5 mm long, obtuse to acute; fruits not seen. Evergreen lowland forests, 100–200 m; Amazonas (Yavita). Amazonian Colombia.

Plants from Amacayacu National Park, which lies along the Amazon River in southern Colombia, appear to be correctly identified as Ixora yavitensis, but plants from Amazonian Ecuador and Peru treated under this name may belong to a different species. Unfortunately few fertile specimens are available for good evaluation of these populations.

47. KUTCHUBAEA Fisch. ex DC., Prodr. 4: 373. 1830. by Charlotte M. Taylor and Julian A. Steyermark Trees, unarmed, terrestrial, dioecious, frequently resinous on buds and young growth; bark frequently exfoliating in large plates or strips. Leaves opposite, petiolate, venation not lineolate; stipules interpetiolar and sometimes shortly intrapetiolar, generally imbricate to valvate in bud, persistent, triangular to ligulate. Inflorescence terminal, ebracteate, pedunculate to sessile, the staminate severalflowered, corymbose, fasciculate, umbellate-cymose, or capitate, the pistillate 1(–3)flowered. Flowers large, showy, fragrant, unisexual, sessile to pedicellate or pedunculate. Staminate flowers: hypanthium reduced; calyx limb truncate, undulate, or shortly dentate, without calycophylls; corolla salverform, white to yellow, externally glabrous to usually sericeous, internally sericeous to villosulous in throat, the tube well developed, the lobes 6–11, convolute (to left) in bud; stamens 6–11, inserted in upper part of corolla tube; anthers dorsifixed, linear-oblong, cuspidate at apex, bifid at base, included, situated in corolla throat; pistillode present, similar to style and stigma, included or partially exserted. Pistillate flowers: hypanthium ellipsoid; calyx limb and corolla similar to the staminate or the calyx limb larger and the corolla tube shorter; staminodes not seen, presumably present and similar to anthers; ovary at first 2-locular, eventually nearly 1-locular; ovules numerous, on axile placentas. Fruit baccate, subglobose to ellipsoid, mature color not noted, exocarp generally woody, pulp gelatinous. Seeds medium-sized, ellipsoid, flattened to angled. Colombia, Venezuela, Guyana, French Guiana, Ecuador, Peru, Brazil; 11 species, 5 in Venezuela, all in the flora area. This genus has often been incorrectly written as “Kotchubaea” but as noted by Delprete (1999a), it was originally published as “Kutchubaea” and is thus correctly spelled this way. This showy genus is little known and often overlooked or confused with Albertia, which has smaller flowers. The pistillate flowers of most species have not yet been documented. Kutchubaea sericantha is by far the most commonly collected species. Key to the Species of Kutchubaea 1. 1. 2(1). 2. 3(2).

Base and apex of the leaf blade rounded; petioles 1–3 mm long .....K. morilloi Base and apex of the leaf blade obtuse, acute, or acuminate; petioles 3– 30 mm long ............................................................................................. 2 Staminate calyx with teeth 1.5–2 mm long; staminate corolla tube 22– 25 mm long, shorter than the lobes, these 45–60 mm long ... K. longiloba Staminate calyx truncate or with teeth to 1.2 mm long; staminate corolla tube 25–70 mm long, longer than the lobes, these 15–41 mm long .... 3 Leaves with secondary veins 12–18 pairs and the tertiary venation on the lower surface prominulous and manifestly reticulate ......... K. sericantha

632

3.

4(3).

4.

R UBIACEAE

Leaves with secondary veins 4–12 pairs and the tertiary venation on the lower surface not visible or visible but plane and not noticeably reticulate .......................................................................................................... 4 Calyx of the staminate flowers minutely puberulous or granular-puberulent externally; staminate corolla with tube 22–28 mm long, lobes 15– 20 mm long ............................................................................. K. micrantha Calyx of the staminate flowers glabrous externally; staminate corolla with tube 35–45 mm long, lobes 29–31 mm long ................ K. neblinensis

Kutchubaea longiloba Steyerm., Mem. New York Bot. Gard. 10(5): 218, fig 75. 1963. Tree to 15 m tall; leaves 7–13 × 2–6 cm; secondary veins 4 or 5 pairs; petioles 5–10 mm long; stipules 5–7 mm long; staminate flowers 2 or 3; calyx limb 12–17 mm long; corolla tube 22–35 mm long, the lobes 8, 45–60 mm long; pistillate flowers and fruits not

seen. Tepui slope forests, ca. 1000 m; Bolívar (Macizo del Chimantá [Toronó-tepui]). Endemic. ◆Fig. 495. Kutchubaea micrantha Steyerm., Mem. New York Bot. Gard. 10(5): 215.1963. Tree to 15 m tall; leaves 8–14 × 4–9 cm; secondary veins 5–7 pairs; petioles 3–5 mm long; stipules 5–9 mm long; staminate flow-

Fig. 494. Kutchubaea morilloi Fig. 495. Kutchubaea longiloba

Kutchubaea 633

Fig. 496. Kutchubaea neblinensis

Fig. 497. Kutchubaea sericantha

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R UBIACEAE

ers 6–13; calyx limb 7–9.5 mm long; corolla tube 22–28 mm long, the lobes 7, 15–20 mm long; pistillate flowers and fruits not seen. Seasonally flooded riparian forests, 100–200 m; Amazonas (upper Río Atacavi). Amazonian Colombia. Kutchubaea morilloi Steyerm., Act. Bot. Venez. 9(1–4): 289, fig. 1. 1973. Tree to 12 m tall; leaves 5.5–10 × 3–5.5 cm; secondary veins 5–8 pairs; petioles 1–3 mm long; stipules 3–4 mm long; staminate flowers 4–7; calyx limb 8–15 mm long, subtruncate; corolla tube 28–29 mm long, the lobes 7 or 8, 25–26 mm long; pistillate flowers and fruits not seen. Tepui slope and summit forests, 1300–1900 m; Bolívar (Cerro Jaua), Amazonas (Cerro Parú at Laguna Asisa). Endemic. ◆Fig. 494. Kutchubaea neblinensis Steyerm., Mem. New York Bot. Gard. 10(5): 216. 1963.

Tree to 20 m tall; leaves 7–14.5 × 3–7 cm; secondary veins 4–7 pairs; petioles 7–15 mm long; stipules ca. 4 mm long; staminate flowers 4–6; calyx limb 6–10 mm long; corolla tube 35–45 mm long, the lobes 8, 29–31 mm long; pistillate flowers and fruits not seen. Tepui slope forests, 600–700 m; Amazonas (Sierra de la Neblina). Endemic. ◆Fig. 496. Kutchubaea sericantha Standl., Publ. Field Columbian Mus., Bot. Ser. 8(5): 355. 1931. Tree to 20 m tall; leaves 14–35 × 7–15 cm; secondary veins 12–18 pairs; petioles 1–3 cm long; stipules 8–15 mm long; staminate flowers 4–11; calyx limb 12–20 mm long, subtruncate; corolla tube 35–50 mm long, the lobes 8–11, 25–45 mm long; pistillate flowers 1–3; fruits 8–11 × 5–7 mm. Tepui slopes and summits, ca. 1400 m; Amazonas (Cerro Sipapo). Amazonian Ecuador, Peru, and Brazil. ◆Fig. 497.

48. LADENBERGIA Klotzsch in Hayne, Getr. Darst. Gew. 14: ad t. 15. 1846. by Charlotte M. Taylor and Julian A. Steyermark Small to large trees, unarmed, terrestrial. Leaves opposite or rarely ternate, petiolate, venation not lineolate; stipules caducous, variously calyptrate (i.e., fused into a conical cap), partially fused around the stem, or usually interpetiolar, triangular, generally valvate to erect and flatly appressed in bud. Inflorescence terminal and in the uppermost leaf axils, paniculate, multiflowered, pedunculate, the bracts reduced or lacking. Flowers sessile to pedicellate, distylous, medium to large, fragrant, perhaps nocturnal. Hypanthium ellipsoid to turbinate. Calyx limb truncate or with lobes 5–7, without calycophylls; corolla salverform, white, usually glabrous internally, lobes 5–7, valvate in bud, densely papillose along their margins. Stamens 5–7, inserted in corolla tube; anthers narrowly ellipsoid, dorsifixed near base, included to partially exserted. Ovary 2-locular; ovules numerous in each locule, on axile placentas. Fruit capsular, cylindrical to oblong or narrowly cylindrical, septicidally dehiscent usually from the apex, chartaceous to woody. Seeds flattened, small, irregularly elliptic to oblong, marginally winged, often erose. Costa Rica, Panama, Colombia, Venezuela, Ecuador, Peru, Guyana, Brazil, Bolivia; ca. 34 species, 9 in Venezuela, 4 of these in the flora area. Ladenbergia, Cinchona, Cinchonopsis, and Remijia are closely related and have long been confused, but recent work by Andersson (1995) has shed significant light on generic circumscriptions in this group. Ladenbergia was long distinguished from Cinchona by fruit characters, in particular the capsules supposedly dehiscing from the apex in Ladenbergia versus from the base in Cinchona. However, a number of species are known to be variable in this regard, and Andersson showed that these two genera are more usefully distinguished by floral characters. Thus as circumscribed by him and followed here, Ladenbergia has white flowers that are generally glabrous internally and papillose along the margins of the lobes, while Cinchona has white to usually pink or purple flowers that are internally pubescent and villous

Ladenbergia 635

along the margins of the corolla lobes. Remijia is distinguished by its axillary inflorescences. Cinchona is not found in the flora area; our species from the flora area that was formerly placed in this genus differs from Cinchona in its homostylous white flowers and was separated by Andersson as Cinchonopsis. Subsequently, Andersson (1997) reviewed Ladenbergia, including presenting nomenclatural synonymy, a key, and representative exsiccatae records for the 34 species he recognized. Ladenbergia is also similar to Ferdinandusa, especially when in fruit; Ferdinandusa can be recognized by its stipules twisted in bud and its corolla lobes convolute in bud. Steyermark (1972; 1974) considered presence versus absence of pubescence on the style to be taxonomically significant in Ladenbergia, but Andersson (1997) concluded that this character shows continuous variation within species of Ladenbergia, in particular L. lambertiana. Andersson also found details of the pubescence on the leaves and reproductive structures, shape of the leaf base, and degree of lobing of the calyx limb notably variable within most species of Ladenbergia, and certainly less consistent than Steyermark believed. Consequently Andersson (1997) recognized fewer, more clearly circumscribed species than Steyermark; Andersson’s treatment is followed here. In general, the distinctions between even the more conservatively circumscribed species of Ladenbergia are sometimes subtle. Key to the Species of Ladenbergia 1.

1.

2(1).

2.

3(2). 3.

Leaves obtuse to truncate at base, abaxially with costa and secondary veins moderately to densely strigose or strigillose (i.e., the pubescence all appressed), without domatia ........................................ L. amazonensis Leaves obtuse to truncate or cordate at base, abaxially with costa and secondary veins glabrous, sparsely appressed-puberulous with the trichomes short and separated, or sparsely to moderately hirtellous or pilose (i.e., the pubescence at least partially spreading), with or without domatia ............................................................................................ 2 Leaves 3–11 cm wide, lanceolate to lance-oblong, eliptic-oblong, or narrowly elliptic, abaxially glabrous in the axils of the secondary veins or sparsely hirtellous here with trichomes restricted to the vein axils .............................................................................................. L. lambertiana Leaves 4–16 cm wide, narrowly to broadly elliptic or ovate, abaxially densely hirtellous in the axils of the secondary veins with pubescence usually extending out from the axils along both the costa and the secondary veins ........................................................................................... 3 Capsules 25–70 × 6–10 mm ..................................................... L. oblongifolia Capsules 25–120 × 4–5 mm ................................................................. L. sp. A

Ladenbergia amazonensis Ducke, Trop. Woods 31: 21. 1932. Tree to 22 m tall; leaves 11–24 × 5–13 cm; secondary veins 7–11 pairs; petioles 2–6.5 cm long; stipules interpetiolar, 10–28 mm long; inflorescence 6–17 × 5–10 cm; calyx limb 2– 3.5 mm long, lobed for ca. 1/2 its length; corolla tube 13–16 mm long, lobes 7–8.5 mm

long; capsules lanceoloid, 35–60 × ca. 7 mm; seeds 15–18 × 1.5–2 mm long. Lowland white-sand forests, montane forests on granite, 100–1000 m; Amazonas (Cerro Aratitiyope, Río Guainía, Río Mawarinuma, Río Negro, Río Orinoco, San Carlos de Río Negro, Victorino). Amazonian Colombia, Peru, and Brazil. ◆Fig. 498.

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R UBIACEAE

Ladenbergia amazonensis is much more common in the western Amazon basin. Ladenbergia lambertiana (A. Braun ex Mart.) Klotzsch in Hayne, Getr. Darst. Gew. 14: ad t. 15. 1846. —Cinchona lambertiana A. Braun ex Mart., Reise Bras. 1286. 1831. Ladenbergia schomburgkii Klotzsch in Hayne, Getr. Darst. Gew. 14: ad t. 15. 1846. Ladenbergia lucens Standl., Publ. Field Columbian Mus., Bot. Ser. 7: 362. 1931. Ladenbergia puberula Steyerm., Mem. New York Bot. Gard. 23: 270, fig. 56. 1972. Ladenbergia venamoensis Steyerm., Mem. New York Bot. Gard. 23: 271. 1972. Shrub or tree to 20 m tall; leaves 9–28 × 3.5–15 cm; secondary veins 5–14 pairs; petioles 1.5–6 cm; stipules interpetiolar, 1.5–5 cm long; inflorescence 8–24 × 6–15 cm; calyx limb 1.5–3 mm long, lobed shallowly to deeply; corolla tube 14–25 mm long, lobes 6– 10 mm long; capsules 30–70 × 4–8 mm; seeds 12–17 × 3–4 mm long. Savannas, borders of savannas, gallery and secondary forests, forested rocky slopes, over sandstone and igneous substrate, fern thickets, 50–2000 m; common in Delta Amacuro, Bolívar, and Amazonas. Guyana, Amazonian Colombia, Brazil. ◆Fig. 499.

Ladenbergia lambertiana is the most commonly collected species in the genus in the flora area. Ladenbergia puberula was separated based on its leaves that are pubescent abaxially. This pubescence is found in a few plants from the region of Sierra de la Neblina (Amazonas), Perai-tepui (Bolívar), and the Utshe River in the country of Guyana (McDowell and Gopaul 2800, MO), but otherwise no other features separate these plants. This pubescence is apparently sometimes variable (and/or possibly caducous) on an individual plant (e.g., Liesner 6730, MO), consequently Andersson (1997) concluded that this name is best considered a synonym of L. lambertiana. Ladenbergia lucens was separated from L. lambertiana only by its pubescent rather than glabrous style, but as noted by Andersson (1997) this character varies continuously and is not taxonomically informative. Ladenbergia venamoensis was separated from L. lambertiana only by its glabrescent to sparsely pubescent hypanthium, another character that is now known to be continuously variable. Ladenbergia oblongifolia (Mutis) L. Andersson, Fl. Ecuador 50: 19. 1994. —Cinchona oblongifolia Mutis, Papel Periódico de Santa Fé 111: 465. 1793. Cinchona magnifolia Ruiz & Pav., Fl. Peruv. 2: 53, t. 196. 1799. —Ladenbergia magnifolia (Ruiz & Pav.) Klotzsch in Hayne, Getr. Darst. Gew. 14: ad t. 15. 1846.

Fig. 498. Ladenbergia amazonensis

Ladenbergia 637

Fig. 499. Ladenbergia lambertiana

Tree to 10 m tall; leaves 20–33 × 15–30 cm; secondary veins 8–12 pairs; petioles 2–6 cm long; stipules interpetiolar, 2–3 cm long; inflorescences 15–30 × 10–30 cm; calyx limb ca. 2 mm long, lobed for 1/2–3/4; corolla tube 6– 11 mm long, lobes 7–9 mm long; capsules 25– 70 × 7–10 mm; seeds not seen. Ridge tops of montane forests over quartzite and granite, ca. 1300 m; Amazonas (Sierra Parima near Brazilian frontier). Colombia, Peru, Bolivia,

Brazil (Roraima). The plants of Ladenbergia oblongifolia found in the flora area are significantly disjunct from the principal range of this species, in the Andes from Colombia through Bolivia. Both Steyermark (1972; 1974) and Andersson (1997) considered these plants conspecific, but when more material is available the Venezuelan and Brazilian plants probably deserve re-evaluation.

638

R UBIACEAE

Ladenbergia sp. A Tree to 16 m tall; leaves 9–27 × 4.5–17 cm; secondary veins 6–11 pairs; petioles 1.5–5.5 cm long; stipules 1–2 cm long; infructescences 17–25 × 10–15 cm; capsules 25–120 × 4–5 mm;

seeds ca. 20 × 2 mm. Evergreen lowland to lower montane forests, 100–600 m; Amazonas (Caño Majagua, Raudal Ceguera, Río Autana). These plants await discovery of flowering material for a definite identification.

49. LIMNOSIPANEA Hook. f. in Hook., Icon. Pl. pl. 1050. 1868. by Piero G. Delprete and Julian A. Steyermark Herbs, terrestrial and often aquatic, unarmed, with the stems slender, terete, erect or the basal internodes sometimes prostrate. Leaves opposite or 3–6-verticillate, sometimes heterophyllous with the submerged leaves linear, sessile or subsessile, venation not lineolate; stipules obsolete, reduced to a seta subtended by a group of glands, or interpetiolar with the sheath united to the petioles, narrowly triangular, persistent, the arrangement in bud unknown. Inflorescence terminal, pedunculate, bracteate, few-flowered to multiflowered, with loosely 2–manybranched cymes. Flowers small, subsessile, bibracteolate, homostylous, protandrous. Hypanthium pubescent, ellipsoid to subglobose. Calyx limb pubescent, deeply lobed, lobes 5, generally equal and well developed, without calycophylls; corolla salverform or narrowly funnelform, white and yellow or sometimes roseate, internally pubescent in the mouth of the tube, lobes 5, convolute in bud. Stamens 5, inserted in the corolla throat; anthers narrowly oblong, exserted, dorsifixed. Ovary 2-locular; ovules numerous in each locule, on axile placentas. Fruit capsular, subglobose to ellipsoid, crustaceous to chartaceous, loculicidal from the apex. Seeds small, angled, foveolate or minutely scrobiculate. Panama, Colombia, Venezuela, Guyana, Brazil, Bolivia; 4 species, 2 in Venezuela, both in the flora area. Limnosipanea is similar to and sometimes confused with Perama; their separation is discussed under Perama. Plants of Limnosipanea sometimes grow partially submersed in wet savannas and along pond and stream edges, with the submersed leaves often quite different in form from the aerial leaves. Limnosipanea, particularly L. spruceana, is often overlooked as a genus of Rubiaceae because the stipules are so reduced they appear to be absent, and the aquatic habit and whorled leaves of some species are unusual in this family. Galium, found in cool montane and premontane habitats, also has whorled leaves without evident stipules. Key to the Species of Limnosipanea 1. 1.

Leaves opposite or ternate, usually ciliate and sparsely pilose on the veins on the lower surface ........................................................ L. palustris Leaves 3–6-verticillate, glabrous on the lower surface ............. L. spruceana

Limnosipanea palustris (Seem.) Hook. f. in Hooker’s Icon. Pl. 11: 38, pl. 1050. 1868. —Sipanea palustris Seem., Bot. Voy. Herald 136. 1854, non (A. Rich.) J.H. Kirkbr. 1997, nom. illeg. hom. Limnosipanea schomburgkii Hook. f. in Hook., Icon. Pl. 1050. 1868. Herb 5–40 cm tall, sparsely branched; leaves 2(3) per node, 5–18 × 1–8 mm; calyx

lobes 3–3.5 mm long; corolla tube 3.5–6.5 mm long, the lobes 2.2–3.5 mm long; capsules 2.5–3.5 × 2–2.5 mm. Granitic exposures along streams and savannas, 100–1200 m; Bolívar (between mouth of Río Horeda and Cerro Gavilán, near Roraima-tepui), Amazonas (56 km northeast of Puerto Ayacucho). Guárico; Panama, Colombia, Venezuela, Guyana, Brazil. ◆Fig. 500.

Machaonia 639

Steyermark distinguished Limnosipanea schomburgkii based on its entire calyx lobes, its leaves 2 per node, and its range in northeastern South America, versus L. palustris with biaristate calyx lobes, 3 or 4 leaves per node, and a distribution in Panama. The characters are variable throughout the range of this complex, and these two names are here considered synonyms. Limnosipanea spruceana Hook. f. in Hook., Icon. Pl. 1050. 1868. Limnosipanea kuntzei Standl., Publ. Field Columbian Mus., Bot. Ser. 7: 280. 1931. Limnosipanea guaricensis Pittier, Bol. Soc. Venez. Cienc. Nat. 8: 145. 1943. Limnosipanea ternifolia Pittier, Bol. Soc. Venez. Cienc. Nat. 8: 144. 1943. Herb 7–45 cm tall, lower part of the stem often submersed; aerial leaves 3–6(–8) per node, 5–20 × 0.3–4 mm, submersed leaves linear; calyx lobes 1.5–4 mm long; corolla tube 2–2.5 mm long, the lobes 1–3 mm long; capsules 2–4 × 1.5–2.5 mm. Wet savannas, borders of Mauritia palm swamps, 50–900 m; Bolívar (between km 12 and 120 on road from Caicara del Orinoco toward Puerto Ayacucho, Hato Divina Pastora), Amazonas (basin of Río Manapiare on lower and middle Río Parucito). Cojedes, Guárico; Colombia, Guyana, Brazil, Bolivia. ◆Fig. 501. Steyermark distinguished Limnosipanea ternifolia by its lower and upper cauline leaves that are generally uniform in shape, its middle cauline leaves ternate, its calyx lobes 4–5.5 mm long, and its corollas shorter than or equal to the calyx lobes, versus markedly different in shape, 4–6-verticillate, 1.5–2 mm long, and markedly longer, in L. spruceana. However, in field studies the variation along the stem in leaf form and arrangement was observed to be due largely to the heterophylly found on partially submersed stems, while the plants with the leaf form and arrangement generally similar along the stem were not partially submersed. The other characters used by Steyer-

Fig. 500. Limnosipanea palustris Fig. 501. Limnosipanea spruceana

mark were observed to have continuous variation. Consequently, these names are here treated as synonymous.

50. MACHAONIA Humb. in Humb. & Bonpl., Pl. Aequinoct. 1: 101, pl. 29. 1806 [1808]. by Charlotte M. Taylor and Julian A. Steyermark Shrubs or trees, terrestrial, often armed with axillary spines or short spinescent branches. Leaves opposite or verticillate, petiolate, venation not lineolate; stipules interpetiolar, triangular, persistent, generally valvate to imbri-

640

R UBIACEAE

cate in bud. Inflorescence terminal, multiflowered, pedunculate, paniculate to cymose, bracteate. Flowers sessile or pedicellate, small, white, homostylous, protandrous. Hypanthium turbinate to ellipsoid, often laterally compressed. Calyx limb deeply lobed, lobes 4 or 5, equal to markedly unequal but without calycophylls; corolla shortly funnelform, white or cream to yellow, internally densely villous, the lobes 4 or 5, imbricate in bud. Stamens 4 or 5, inserted in the corolla throat; anthers narrowly oblong, dorsifixed, included or exserted. Ovary 2-locular; ovules solitary in each locule, pendulous from the locule apex. Fruit schizocarpous (or samaroid and indehiscent in M. lindeniana Baill. of Mexico and Central America), dry, ellipsoid; mericarps 2, indehiscent, separating from a central, generally persistent carpaphore. Seeds cylindrical, small. Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Ecuador, Brazil, Bolivia, Paraguay; ca. 30 species, 2 species in Venezuela, 1 of these in the flora area. Machaonia is sometimes confused with Chomelia, but that genus can be recognized by its fleshy drupaceous fruits with a single multilocular pyrene. Additionally, most of the species of Chomelia that are similar in general aspect to Machaonia also have lineolate tertiary leaf venation, which Machaonia lacks. Machaonia brasiliensis (Hoffmanns. ex Humb.) Cham. & Schltdl., Linnaea 4: 12. 1829. —Cinchona brasiliensis Hoffmanns. ex Humb., Berlin Mag. 1: 104, 119. 1807. Shrub or small tree to 6 m tall; leaves 5– 10 × 2–4 cm; stipules 1–2 mm long; inflorescences 4–12 × 4–12 cm; calyx lobes ca. 1 mm long; corolla with the tube 1.5–2 mm long, the lobes 1.5–2 mm long; fruits ca. 4.5 × 1.5–2 mm. Gallery forests, lowland forests, 100– 300 m; Bolívar (eastern portion west to Río Paragua). Brazil, Paraguay. ◆Fig. 502. Species distinctions are subtle (or not entirely clear) in this genus, and Machaonia brasiliensis may not be distinct from M. spinosa Cham. & Schltdl. of Brazil and Paraguay.

Fig. 502. Machaonia brasiliensis

51. MAGUIREOCHARIS Steyerm., Mem. New York Bot. Gard. 23: 230, fig. 50. 1972. by Julian A. Steyermark and Charlotte M. Taylor Shrubs, terrestrial, unarmed, the stems and leaves usually pale and glaucous. Leaves opposite, petiolate, venation not lineolate; stipules interpetiolar, caducous, oblanceolate, obtuse to rounded, generally erect and flatly appressed. Inflorescence terminal, in the uppermost leaf axils and/or pseudoaxillary, pedunculate, cymose,

Maguireocharis 641

many-flowered, bracteate. Flowers medium-sized, sessile to shortly pedicellate, floral biology unknown. Hypanthium turbinate. Calyx limb lobed nearly to base, lobes 5, without calycophylls; corolla funnelform, roseate, internally glabrous except hirtellous in upper part of tube and on lobes, lobes 5, valvate in bud. Stamens 5, inserted in upper part of corolla tube; anthers narrowly oblong, dorsifixed near base, partially exserted, hirtellous. Ovary 2-locular; ovules numerous in each locule, on axile placentas. Fruit capsular, ellipsoid to oblong, chartaceous to woody, septicidal from base, each valve often then splitting from base. Seeds small, oblong to subelliptic, flattened, marginally winged. Venezuela, Brazil; 1 species. Chalepophyllum, Duidandia, and Maguireocharis are sometimes sympatric and similar in general aspect, in particular in their similar cymose inflorescences, capsular fruits, and tough-textured leaves with rounded to obtuse apices. In flower, Chalepophyllum differs in its corolla lobes convolute in bud, versus valvate in the other genera. Duidania differs from Maguireocharis in its yellow corollas, versus roseate ones in Maguireocharis. In fruit Chalepophyllum differs in its angled seeds, versus winged fruits in the other genera; Duidania differs from Maguireocharis in its loculicidal capsules, versus septicidal ones in Maguireocharis. Steyermark originally described the lower surface of the leaf of Maguireocharis as “covered with a gray cellular network with orbicular scales.” Scales are otherwise unreported in the Rubiaceae; no material of this genus has been seen to amplify his description. Steyermark also described the ovary placentation as “central,” as did Andersson (1995), who considered the placentation of Maguireocharis similar to that of Ladenbergia and Remijia in his morphological study of the tribe Cinchoneae. The placentation of Ladenbergia was not described by Andersson

Fig. 503. Maguireocharis neblinae

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R UBIACEAE

(1997) in his synopsis of that genus; that of Cinchona was described by Andersson (1998) as “placentae attached to the septum,” which corresponds to the arrangement generally described in Rubiaceae as “axile.” Therefore, the arrangement of all three of these genera is here described as axile. Steyermark (1974) illustrated the ovaries of species of these three genera as having the placentas inserted on the septum. However, Steyermark’s original illustration of Maguireocharis depicts the placentas as basal. Andersson (1997) concluded that the taxonomic affinities of Maguireocharis are with Cinchonopsis, found in the flora area, and with Pimentelia Wedd. and Stilpnophyllum Hook. f. of Andean Peru and Bolivia. Maguireocharis neblinae Steyerm., Mem. New York Bot. Gard. 23: 230, fig. 50. 1972. Shrub to 4 m tall; leaves 2.5–10 × 1.2–4.5 cm; stipules 15–20 mm long; inflorescences 5.5–6 × 5–6 cm; calyx limb 1.5–2 mm long;

corolla tube 7.5–8 mm long, lobes ca. 4.5 mm long; capsules 12–14 × 4–6 mm. Upper tepui slopes, 1300–1500 m; Amazonas (southern canyon slopes of Sierra de la Neblina). Adjacent northern Brazil (Sierra Pirapucú). ◆Fig. 503.

52. MAGUIREOTHAMNUS Steyerm., Mem. New York Bot. Gard. 10(5): 220. 1963. by Piero G. Delprete and Julian A. Steyermark Shrubs, erect or prostrate, terrestrial, unarmed, sparsely branched, the internodes usually short and the leaves consequently crowded, sometimes only present in groups at stem apices. Leaves opposite, subsessile, thickly coriaceous, venation not visible; stipules interpetiolar, persistent, in bud generally valvate, truncate with a central triangular or subulate lobe. Inflorescences with flowers solitary in upper leaf axils, sessile to shortly pedunculate, bracteate, the bracts foliaceous. Flowers large, showy, very fragrant, bibracteolate, homostylous, protandrous. Hypanthium ellipsoid. Calyx limb deeply lobed, lobes 5, well developed, equal or unequal but without calycophylls; corolla salverform with the tube elongated, white becoming cream-colored when old, internally tomentose or villosulous in the upper part of the tube and glabrous or puberulous in the lower part, lobes 5 or 6, convolute in bud. Stamens 5, inserted in upper part of corolla tube; anthers included, narrowly oblong, sagittate, dorsifixed; filaments short. Ovary 2-locular; ovules numerous in each locule, on axile placentas. Fruit capsular, ellipsoid, chartaceous to woody, loculicidal from the apex. Seeds small, angular, irregularly subrhomboid, subcompressed, subalate; testa foveolate. Venezuela, Guyana; 2 species, both in the flora area. The flowers have a fragrance reminiscent of gardenias and are probably pollinated by hawk moths. Key to the Species of Maguireothamnus 1. 1.

Ovary, hypanthium, and capsule glabrous ................................. M. speciosus Ovary, hypanthium, and capsule pubescent ....................................... M. tatei

Maguireothamnus speciosus (N.E. Br.) Steyerm., Mem. New York Bot. Gard. 10(5): 223, fig. 76A–C. 1963. —Chalepophyllum speciosum N.E. Br., Trans. Linn. Soc. London, ser. 2, 6: 33, pl. 5, figs. 10–17. 1901.

Shrub 0.5–3 m tall, sometimes prostrate; leaves (1.5–)2.2–6.5 × 0.5–2.5 cm; stipules triangular and gradually tapering at apex or sometimes broadly triangular at base and aristate at apex, 2–10 mm long; calyx lobes 10–32 mm long; corolla tube 6.5–13 cm long,

Maguireothamnus 643

Fig. 504. Maguireothamnus speciosus subsp. jauaensis

Fig. 505. Maguireothamnus speciosus subsp. speciosus

Fig. 506. Maguireothamnus tatei

the lobes acute or round at apex, 7–55 × 3–23 mm; capsules 10–25 × 8–15 mm. Tepui scrub, 1700–2600 m. Guyana. 2 varieties, both in the flora area.

Maguireothamnus jauaensis var. breweri Steyerm., Fl. Venez. 9: 311. 1974. Tepui scrub and meadows, 1700–2000 m; Bolívar (Cerro Jaua, near Río Marajano). Endemic. ◆Fig. 504.

Key to the Subspecies of M. speciosus 1. Corolla tube internally puberulent in the lower half ................ subsp. jauaensis 1. Corolla tube internally glabrous in the lower half ................. subsp. speciosus M. speciosus subsp. jauaensis Steyerm., Mem. New York Bot. Gard. 23: 883. 1972. —Maguireothamnus jauaensis (Steyerm.) Steyerm., Fl. Venez. 9: 309. 1974.

M. speciosus subsp. speciosus Chalepophyllum coriaceum Gleason, Brittonia 3: 192. 1932. Tepui scrub and meadows, 1800–2600 m; Bolívar (Roraima-tepui west to Cerro Jaua). Guyana. ◆Fig. 505. Maguireothamnus tatei (Standl.) Steyerm., Mem. New York Bot. Gard. 10(5): 227. 1963. —Chalepophyllum tatei Standl.,

644

R UBIACEAE

Publ. Field Columbian Mus., Bot. Ser. 7: 380. 1931. Chalepophyllum latifolium Standl., Publ. Field Columbian Mus., Bot. Ser. 7: 379. 1931. —Maguireothamnus tatei var. latifolius (Standl.) Steyerm., Mem. New York Bot. Gard. 10(5): 228. 1963. Shrub 0.5–2 m tall; leaves 1.2–4 × 0.3–2 cm; stipules broadly triangular at base and

aristate at apex, 0.7–7.5 mm long; calyx lobes 8–25 mm long; corolla tube 6.5–11 cm long, the lobes acute at apex, 15–36 × (4–)7–16 mm; capsules 10–25 × 8–11 mm. Tepui slope and summit scrub and meadows, 1200–2600 m; Amazonas (Cerro Duida, Cerro Huachamacari, Cerro Marahuaka, Cerro Sipapo). Endemic. ◆Fig. 506.

53. MALANEA Aubl., Hist. Pl. Guiane 106, t. 41. 1775. by Charlotte M. Taylor and Julian A. Steyermark Lianas or occasionally twiners or shrubs, terrestrial, unarmed. Leaves opposite, petiolate, venation lineolate; stipules interpetiolar, ligulate to triangular, caducous, generally erect and flatly appressed in bud. Inflorescence axillary, pedunculate, multiflowered, paniculate to spiciform, bracteate. Flowers sessile, small, homostylous or distylous, apparently protandrous. Hypanthium generally turbinate. Calyx limb truncate or with lobes 4, without calycophylls; corolla rotate to funnelform, white to green or yellow, internally densely villous in throat and on bases of lobes, the lobes 4, valvate (or reportedly sometimes slightly imbricate) in bud. Stamens 4, inserted in the corolla throat; anthers narrowly oblong, exserted, dorsifixed near the base. Ovary 2-locular; ovules solitary in each locule, pendulous from the apex of the locule. Fruit drupaceous, ellipsoid to cylindrical or fusiform, fleshy, red to purple or black; pyrenes 1 or 2, 1- or 2-locular, smooth to longitudinally ridged. Seeds cylindrical, small. Southern Central America, Antilles, Colombia, Venezuela, Trinidad-Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; ca. 35 species, 21 in Venezuela, 15 of these in the flora area. Malanea. Malanea is similar to Chomelia and is in need of study; see comments on their distinction under Chomelia. Key to the Species of Malanea 1. 1. 2(1).

2.

3(1).

3.

Lower surface of leaf blade moderately to densely and evenly, silverygray-velutinous ...................................................................................... 2 Lower surface of leaf blade glabrous or pubescent, but not silvery-graypubescent ............................................................................................... 3 Leaves ovate to elliptic, 4.5–10.5 × 2.5–7.5 cm; style pubescent in the lower half; hypanthium glabrous; calyx limb densely to moderately strigillose, sinuate to shallowly lobed ................................... M. hypoleuca Leaves elliptic to broadly so, 6–17 × 4–10 cm; style completely glabrous; hypanthium glabrous; calyx limb glabrous to sparsely strigillose, truncate to sinuate .................................................................... M. macrophylla Leaf blades 1.7–3.3 cm long with the margins usually revolute and the secondary veins 4 or 5; calyx lobes 1–1.5 mm long, longer than broad; bracts subtending the flowers glabrous or nearly so, decussate in 2 separate, erect, appressed pairs ...................................... M. microphylla Leaf blades 5–16 cm long with the margins flat or revolute and the secondary veins 5–13; calyx lobes, if developed, < 0.7 mm long, as broad as or broader than long; bracts subtending the flowers crowded, proxi-

Malanea 645

4(3). 4. 5(4). 5. 6(5).

6.

7(6). 7. 8(7). 8. 9(8). 9. 10(8). 10.

11(10). 11. 12(11).

12.

13(11). 13. 14(13).

14.

15(14).

mate, spreading, not separate nor decussate ....................................... 4 Leaf blades glabrous on both sides or the costa and secondary veins sparsely strigillose below .......................................................... M. obovata Leaf blades strigose to hirsute on one or both sides at least on the costa and lateral veins .................................................................................... 5 Stipules rounded or obtuse at apex ....................................... M. macrophylla Stipules acute, acuminate, or subcaudate at apex ................................... 6 Pubescence of the costa and secondary veins on the lower leaf surface of hirsute, spreading, or subspreading hairs mainly 1–1.5 mm long ...................................................................................................... M. ursina Pubescence, when present, of the costa and secondary veins on the lower leaf surface of appressed to strongly ascending hairs 1 mm or less long, or, if spreading trichomes then these < 1 mm long .............................. 7 Leaves with costa and secondary veins impressed or elevated on the upper surface ............................................................................ M. sipapoensis Leaves with the costa and secondary veins obscurely to prominently sulcate on the upper surface ...................................................................... 8 Leaf lower surface densely covered and hidden by a crowded tomentum ................................................................................................................ 9 Leaf lower surface visible, not hidden by a crowded tomentum ............ 10 Leaves rounded or obtuse at base and apex; calyx limb 0.8–1 mm long; corolla lobes without setulose trichomes at apex ............... M. sarmentosa Leaves narrowed to an acute or subacute base and apex; calyx limb ca. 1.5 mm long; corolla lobes with setulose trichomes at apex .... M. setulosa Lower surface of leaf blade glabrous or the costa and secondary veins pubescent ................................................................................ M. jauaensis Lower surface of leaf blade shortly appressed- or spreading-pubescent, the costa and secondary veins appressed-pubescent or glabrescent .............................................................................................................. 11 Leaves with the tertiary venation smooth on the lower surface ........... 12 Leaves with the tertiary venation elevated on the lower surface ......... 13 Anthers 1–1.5 mm long; corolla ca. 6 mm long, the lobes 3–3.2 mm long; petiole 6–14 mm long; stipules acuminate to caudate; secondary leaf veins (6–)8–11 pairs ............................................................ M. gabrielensis Anthers 0.8 mm long; corolla 5–5.5 mm long, the lobes 2.5–2.8 mm long; petiole 5–7 mm long; stipules acute; secondary leaf veins 5–8 pairs .................................................................................................... M. ueiensis Anthers subsessile, the filaments 0.1–0.2 mm long ....... M. auyantepuiensis Anthers with filaments 0.4–2 mm long .................................................. 14 Leaf blades rounded, obtuse, or rarely subacute at apex, on the lower surface with the tertiary venation inconspicuously reticulate and only weakly rugulose-elevated ................................................. M. chimantensis Leaf blades acute to acuminate at apex, on the lower surface with the tertiary venation prominently and uniformly reticulate and rugulose-elevated ................................................................................................... 15 Individual floral clusters all sessile on the primary inflorescence axis, the complete inflorescence (measured from base of peduncle to uppermost flowers) 3–6 cm long; style ca. 4.5 mm long; filaments 0.4–0.5 mm long, 2–3 times shorter than anther ....................................M. guaiquinimensis

646

15.

R UBIACEAE

Lowest two tiers of floral clusters borne on secondary inflorescence axes 5–20 mm long, the complete inflorescence 8–12 cm long; style ca. 3 mm or less long; filaments 1.8–2 mm long, longer than the anther ................................................................................................. M. ptariensis

Malanea auyantepuiensis Steyerm., Mem. New York Bot. Gard. 12(3): 256. 1965. Low shrub or climber; leaves 7–11.5 × 3– 5.5 cm; stipules 1–1.2 cm long; inflorescences 3.5–7.5 cm long; calyx limb ca. 1.5 mm long; corolla with tube 1.5–2 mm long, lobes ca. 2 mm long; fruit 10–11 × 5–6 mm. Tepui scrub, 1700–1800 m; Bolívar (Auyán-tepui). Endemic. Malanea chimantensis Steyerm., Mem. New York Bot. Gard. 12(3): 259. 1965. Low climber; leaves 5–11 × 3–7 cm; stipules 6–13 mm long; inflorescences and flowers not seen; infructescences 6–7.5 cm long; fruits 9–10 × 5–8 mm. Tepui forests and scrub, ca. 1400 m; Bolívar (Macizo del Chimantá [Abacapá-tepui]). Endemic. Malanea gabrielensis Müll. Arg., Flora 58: 453. 1875. —Guianaguiarra (Guahibo). Low climber; leaves 6–13 × 3–7.5 cm; stipules 7–16 mm long; inflorescences 3.5–7 cm long; calyx limb ca. 1 mm long; corollas with tubes ca. 3 mm long, the lobes ca. 3 mm long; fruit 6–8 × 3–5 mm long. Riparian forests, around Mauritia palm swamps, forests on alluvial, igneous, or sandstone substrates, 50–700 m; Bolívar (basin of Río Cuyuní and Río Orinoco), southwestern Amazonas. Apure, Zulia; northwestern Amazonian Brazil (Amazonas: basin of upper Río Negro, Río Guainía, and Río Orinoco). Eastern Colombia, Guyana. ◆Fig. 509. Malanea gabrielensis is relatively widespread and commonly collected compared to other species of Malanea in the flora region. Several species separated by Steyermark (notably M. ptariensis an M. ueiensis) may not actually be distinct from M. gabrielensis. Malanea guaiquinimensis Steyerm., Mem. New York Bot. Gard. 12(3): 253. 1965. Small tree; leaves 9–12 × 3–6 cm; stipules 7–10 mm long; inflorescences 3–6 cm long; calyx limb ca. 1.5 mm long; corollas with tubes ca. 3 mm long, the lobes ca. 3 mm long; fruits 11–12 × 5–6 mm. Riparian forests on tepui summits, ca. 1200 m; Bolívar (Cerro Guaiquinima). Endemic.

Malanea hypoleuca Steyerm., Mem. New York Bot. Gard. 12(3): 251. 1965. Liana; leaves 4.5–10.5 × 2.5–7.5 cm; stipules 5–12 mm long; inflorescences 2–4 cm long; calyx limb ca. 1 mm long; corollas with tube ca. 2 mm long, lobes ca. 2.5 mm long; fruits 6–10 × 3–6 mm. Evergreen lowland to lower montane forests, montane wet forests over sandstone, 100–900 m; Bolívar (Cerro Venamo, El Dorado). Guyana, French Guiana. Malanea hypoleuca is similar to the form of M. macropylla that has leaves silveryvelutinous abaxially. Malanea jauaensis Steyerm., Mem. New York Bot. Gard. 23: 885, fig. 16. 1972. Low climber; leaves 5–7 × 1.5–3.5 cm; stipules ca. 5 mm long; inflorescences 1.5–3 cm long; calyx limb ca. 2 mm long; corolla with tube 2–2.5 mm long, lobes ca. 2.5 mm long; fruits not seen. Brushy slopes along streams on tepui summits and upper slopes, 1900–2000 m; Bolívar (Cerro Jaua, Cerro Sarisariñama). Endemic. Malanea macrophylla Griseb., Fl. Brit. W. Ind. 337. 1861. —Malanea macrophylla Bartl. in M.R. Schomb., Reis. Br.-Guiana 947. 1848 [1849], nom. nud. Liana or shrub; leaves 6–15 × 3–9 cm; stipules 1–2.5 cm long; inflorescences 1–8 cm long; calyx limb 0.5–1 mm long; corolla with tubes 2–4 mm long, lobes 2–2.5 mm long; fruits 8–10 × 3–5 mm. Evergreen lowland to lower montane forests, 50–600 m; Delta Amacuro (east of Caño Sacupana), Bolívar (Altiplanicie de Nuria, Amaruay-tepui). Apure, Miranda, Monagas; Antilles, Guyana, Suriname, French Guiana, Peru, Brazil. Malanea macrophylla is a widespread, morphologically variable species. Accordingly, Steyermark (1965) recognized three varieties and three forms based on density of the velutinous, usually silvery pubescence on the lower surface of the leaves, the shape of the leaves, and the size of the flowers. Large flowers distinguished var. megalantha (Wernham) Steyerm. of Tobago from the remaining plants. Dense pubescence on the abaxial leaf surfaces distinguished f. macro-

Malanea 647

phylla; acute leaf bases distinguished f. cuneata Steyerm.; and the remaining plants were placed in f. bahiensis (Müll. Arg.) Steyerm. Plants of the flora area fall into Steyermark’s f. macrophylla, although the variation among these features is probably continuous in the forms and likely the varieties so they are not considered taxonomically informative. Malanea microphylla Standl. & Steyerm., Fieldiana, Bot. 28(3): 582. 1953. Sprawling low shrub; leaves 1.7–3.3 × 1– 2.5 cm; stipules 3–5 mm long; inflorescences 1.5–3 cm long; calyx limb 1.5–2 mm long; corolla with tubes ca. 2.5 mm long, lobes ca. 2.5 mm long; fruits ca. 10 × 5 mm. Dwarf tepui summit forests, 1900–2300 m; Bolívar (Macizo del Chimantá). Endemic. ◆Fig. 508. Malanea obovata Hochr., Bull. New York Bot. Gard. 6: 289. 1910. Malanea angustifolia Bartl. in M.R. Schomb., Reis. Br.-Guiana 3: 947. 1848 [1849], nom. nud. Malanea roraimensis Wernham, J. Bot. 50: 243. 1912. Low shrub or woody vine; leaves 7–12.5 × 2–5 cm; stipules 5–10 mm long; inflorescences 4.5–10 cm long; calyx limb ca. 1 mm long; corolla with tubes 1.5–2 mm long, lobes 2–2.5 mm long; fruits 7–10 × 3–6 mm. Riparian forests, borders of savannas, 400–1000 m; Bolívar (Río Parupa, near Roraima-tepui, Sabana de Topopo on lower Río Caroní). Adjacent Guyana. ◆Fig. 507. Steyermark distinguished Malanea schomburgkii Steyerm. of Guyana from M. obovata based on its more numerous secondary leaf veins (8–11 pairs, versus 7–9 pairs in M. obovata), its larger corollas (4.5–5.5 mm long, versus 3.5–4.2 mm long in M. obovata), and its “usually conspicuously revolute” leaf margins (versus “plane to subrevolute” in M. obovata). Steyermark identified one specimen (Steyermark et al. 131854, MO, Bolívar) as M. schomburgkii based on its corolla size, but its leaf characters conform to those of M. obovata. Specimens determined by Steyermark as M. obovata (e.g., Pinkus 218, MO) actually have corollas that range to 4–5 mm long, so this feature does not seem to separate these species. Consequently the Venezuelan plants referred by others to M. schomburgkii are here included in M. obovata.

Malanea ptariensis Steyerm., Mem. New York Bot. Gard. 12(3): 255. 1965. Low climber; leaves 8–14 × 4–7 cm; stipules 9–16 mm long; inflorescences 8–12 cm long; calyx limbs ca. 1.5 mm long; corolla with tube 2–2.5 mm long, lobes ca. 3.5 mm long; fruits ca. 10 × 6 mm. Tepui slope and summit forests and scrub, 1000–2400 m; Bolívar (Ptari-tepui), Amazonas (Cerro Marahuaka). Endemic. Malanea sarmentosa Aubl., Hist. Pl. Guiane 106, pl. 41. 1775. Malanea sarmentosa f. tomentosa Steyerm., Mem. New York Bot. Gard. 12(3): 258. 1965. Low climber; leaves 3.5–16 × 2–10 cm; stipules 8–12 mm long; inflorescences 5–14 mm long; calyx limb 0.8–1 mm long; corolla with tube ca. 2 mm long, lobes 2–3 mm long; fruits 6–10 × 4–8 mm. Evergreen lower montane to montane forests, 500–1400 m; eastern Bolívar, Amazonas (Cerro Aratitiyope, Sierra de Maigualida, Sierra Parima by Brazil border). Trinidad-Tobago, Guyana, Suriname, French Guiana, Brazil. ◆Fig. 510. Malanea sarmentosa is often recognizable by its leaves that are densely tomentulose on the lower surface and have revolute margins and the venation sulcate adaxially. It is commonly collected and rather widespread. Steyermark distinguished several varieties based on density of pubescence on the leaves and calyces, but these characters do not seem informative and these varieties are not recognized here. Malanea setulosa Steyerm., Mem. New York Bot. Gard. 12(3): 260. 1965. Low climber; leaves 9–16 × 3.8–7 cm; stipules 9–12 mm long; inflorescences 8–13 cm long; calyx limb ca. 1.5 mm long; corolla with tube ca. 2 mm long, lobes ca. 2 mm long; fruits 10–11 × 4.5–5 mm. Montane gallery forests, 1100–1200 m; Bolívar (along tributary of Río Kukenán, near Roraima-tepui). Endemic. Malanea sipapoensis Steyerm., Mem. New York Bot. Gard. 12(3): 252. 1965. Woody climber; leaves 12–16 × 6.5–9.5 cm; stipules 8–13 mm long; inflorescences 5–9.5 cm long; calyx limb ca. 0.8 mm long; corolla not seen; fruits 10–11 × 4 mm. Wet lower montane forests, ca. 600 m; Amazonas (Cerro Sipapo). Endemic, or possibly also in Guyana (Clarke et al. 6391, MO, US).

648

R UBIACEAE

Fig. 507. Malanea obovata

Fig. 508. Malanea microphylla

Fig. 509. Malanea gabrielensis

Malanea ueiensis Steyerm., Mem. New York Bot. Gard. 12(3): 255. 1965. Low climber or shrub; leaves 7–13 × 2.5–8 cm; stipules 5–16 mm long; inflorescences 7– 9 cm long; calyx limbs ca. 1 mm long; corolla with tube ca. 3 mm long, lobes ca. 2.5 mm long; fruits not seen. Wet montane forests,

1100–2300 m; Bolívar (eastern Río Cuyuní basin, Sororopán-tepui). Guyana. Malanea ursina Standl., Publ. Field Columbian Mus., Bot. Ser. 8: 62. 1930. Low climber; leaves 5–8.5 × 2–4.5 cm; stipules ca. 7 mm long; inflorescences ca. 4

Manettia 649

cm long; calyx limb ca. 1 mm long; corollas and fruits not seen. Evergreen lowland forests, 100–200 m; Amazonas (Río Pasiba). Endemic.

Fig. 510. Malanea sarmentosa

54. MANETTIA Mutis ex L., Mantissa 553. 1771. by Charlotte M. Taylor and Julian A. Steyermark Herbaceous or suffruticose vines or climbers, twining or rarely climbing by adventitious roots, unarmed, terrestrial. Leaves opposite, petiolate, venation not lineolate; stipules interpetiolar, triangular to bidentate or setose, often adnate to petioles, erect, persistent, generally valvate to imbricate in bud. Inflorescence axillary and rarely also terminal, bracteate, pedunculate, multiflowered and fasciculate to cymose or rarely with flowers solitary. Flowers small to medium, pedicellate, homostylous or distylous, protandrous. Hypanthium turbinate to subglobose or cylindrical. Calyx limb 4–8-lobed, without calycophylls; corolla salverform or funnelform, white to pale green, red, blue, or purple, often barbate in throat and internally usually with a pubescent ring in the tube, lobes 4, valvate in bud. Stamens 4, inserted in corolla throat; anthers dorsifixed, narrowly oblong, included or exserted. Ovary 2-locular; ovules numerous in each locule, on axile placentas. Fruit capsular, obovoid to turbinate, septidical, chartaceous to papery. Seeds small, subcircular to discoid, flattened, surrounded by a broad often erose wing. Mexico, Central America, West Indies, South America; ca. 130 species, 20 in Venezuela, 4 of these in the flora area. Manettia and Sabicea are commonly confused, largely because of their similar habit. They can be distinguished easily by their stipules, triangular and erect in Manettia versus ligulate to ovate and spreading in Sabicea. Species of Manettia are often variable in pubescence and leaf size and shape, as in most neotropical Rubiaceae, and notably variable in calyx morphology. In most Manettia species the calyx lobes vary in length on an individual flower, often to

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100%, and sometimes in shape. More unusual, in several species the lobes may vary in number among individual flowers on a single plant as well as among plants. The most variable species in this regard is M. reclinata, which may have as few as four or as many as eight lobes on an individual flower, with these lobes nearly equal in length or with some so reduced that they can be overlooked. The flowers are often borne along pendulous stems. Key to the Species of Manettia 1. 1. 2(2). 2. 3(2). 3.

Calyx lobes 4, 0.7–1.5 mm long; corolla pale green to white, the tube 2– 2.5 mm long ...................................................................................... M. alba Calyx lobes 4–8, (2–)3.5–10 mm long; corolla red, the tube 10–18 mm long ................................................................................................................ 2 Calyx lobes 4, elliptic to ovate, 1.5–2.5 mm wide ........................ M. calycosa Calyx lobes 5–8 or occasionally 4 in a few flowers, linear, lanceolate, or oblanceolate, 0.5–1.2 mm wide ............................................................. 3 Corolla throat densely yellow-bearded, the tube 3–4 mm diameter, the lobes 5–12 mm long .................................................................. M. coccinea Corolla throat glabrous, the tube ca. 2 mm diameter, the lobes 4–6 mm long ........................................................................................... M. reclinata

Manettia alba (Aubl.) Wernham, J. Bot. 57(Suppl.): 29. 1918. —Nacibea alba Aubl., Hist. Pl. Guiane 95, pl. 37, fig. 2. 1775. Herbaceous twiner; leaves 3.5–7 × 1.5–3.5 cm; secondary veins 3–10 pairs; petioles 5–12 mm long; stipules ca. 1 mm long; inflorescences cymose, 2.5–3.5 × 2–3 cm; calyx lobes 4, 0.5–1.5 mm long; corolla pale green to white, tube 2–2.5 mm long, lobes ca. 2.5 mm long; capsules 4–6 × 3–5 mm; seeds 1–1.5 mm diameter. Wet montane or lower montane forests, 300–1200 m; common in eastern Bolívar (basin of Río Cuyuní and Río Caroní west to Macizo del Chimantá and Amaruaytepui). Guyana, Suriname, French Guiana, northeastern Brazil. ◆Fig. 511. Manettia calycosa Griseb., Fl. Brit. W. Ind. 330. 1861. Colombia, Venezuela, Hispaniola; 3 varieties, 1 in Venezuela. M. calycosa var. karsteniana K. Schum. in Mart., Fl. Bras. 6(6): 175. 1889. Herbaceous twiner; leaves 3.5–10.5 × 1.2– 3.5 cm; secondary leaf veins 3 pairs; petioles 2–8 mm long; stipules 1–2 mm long; inflorescences umbelliform or fasciculate, few-flowered; pedicels 4–6 mm long; calyx lobes 3–4 mm long; corolla red, tube ca. 15 mm long, lobes ca. 4 mm long; capsules ca. 6 × 6 mm.

Openings on forested slopes in evergreen seasonally dry montane forests, 1500–1800 m; Amazonas (west of Cerro Yutajé). Northern Venezuela; eastern Colombia. Manettia coccinea (Aubl.) Willd., Sp. Pl. 1: 625. 1797. —Nacibea coccinea Aubl., Hist. Pl. Guiane 96, pl. 37, fig. 1. 1775. Herbaceous twiner; leaves 3–10 × 1–4 cm; secondary leaf veins 4 or 5 pairs; petioles 3– 10 mm long; stipules 1–2 mm long; flowers 1 or 2 per axil; peduncles 2–4 cm long; calyx lobes 4–8, 2–7 mm long; corolla red, tube 10– 18 mm long, lobes 5–12 mm long; capsules ca. 10 × 8 mm; seeds 1–2 mm long. Riparian and lowland forests, 50–200 m; Bolívar (Río Caura at Salto Pará), Amazonas (lower Río Ventauri at Salto Yureba). Carabobo, Yaracuy, Zulia; Colombia, Guyana, Suriname, French Guiana, Brazil. Steyermark recognized two varieties of this species, var. coccinea and var. spraguei (Wernham) Steyerm. He distinguished these by number of calyx lobes, 8 in var. coccinea versus 4–6 in var. spraguei. However, as noted above this character is variable within several species of Manettia, and does not seem taxonomically informative. Manettia coccinea is commonly confused with M. reclinata, particularly when corollas are not available.

Merumea 651

Fig. 511. Manettia alba

Fig. 512. Manettia reclinata

Manettia reclinata Mutis ex L., Mantissa 558. 1767. Manettia uniflora Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 3: 387. 1818 [1820]. Herbaceous twiner; leaves 3–8 × 2–3.5 cm; secondary leaf veins 4–6 pairs; petioles 3–10 mm long; stipules 0.8–1 mm long; flowers 1– 3 per axil; peduncles and/or pedicels 1–4 cm long; calyx lobes 5–8, 2–7 mm long; corolla red to red-purple, tube 12–18 mm long, lobes 4–6 mm long; capsules ca. 10 × 8 mm; seeds 1–2 mm diameter. Riparian forests, evergreen seasonally dry forests bordering bodies of water, 200–400 m; Bolívar (Canaima, basin of Río Caroní), Amazonas (upper Río Orinoco at Ugueto). Anzoátegui, Sucre, Zulia; Mexico, Central America, West Indies, Guyana, Suriname, French Guiana. ◆Fig. 512.

David Lorence (personal communication) considers Manettia uniflora a synonym of M. reclinata, which is a morphologically variable species found in secondary habitats. Manettia reclinata shows variation in particular in number of flowers and degree of development of the peduncles and/or pedicels. Steyermark distinguished M. uniflora by its generally glabrescent leaves and stems, shorter corolla tubes, and inflorescences with fewer flowers. These features are all continuously variable, however, and these names are here considered synonymous. Apparently, Steyermark confuses M. reclinata to some extent with M. coccinea, which may have contributed to his recognition of a third taxon in this group.

55. MERUMEA Steyerm., Mem. New York Bot. Gard. 23: 232, fig. 51. 1972. by Julian A. Steyermark and Charlotte M. Taylor Perennial herbs, terrestrial, unarmed. Leaves opposite, petiolate, venation not lineolate; stipules interpetiolar, triangular, acute to erose, persistent, apparently imbricate to valvate in bud. Inflorescence axillary or terminal, several-flowered, subsessile to pedunculate, capitate to congested-cymose, bracteate, with the exter-

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R UBIACEAE

nal bracts well developed. Flowers medium-sized, subsessile to shortly pedicellate, floral biology unknown. Hypanthium turbinate. Calyx limb deeply lobed, lobes 4 or 5, unequal with 2 or 3 lobes as much as 3 times as long as the shortest lobe, but without calycophylls; corolla salverform to tubular, white to blue, internally glabrous except puberulous in middle and upper part of tube, tube curved, lobes 5, induplicate-valvate in bud. Stamens 5, inserted near middle of the corolla tube; anthers narrowly oblong, included, position of the filament insertion (i.e., dorsifixed or basifixed) unknown. Ovary at least weakly 2-locular; ovules numerous in each locule, on basal placentas. Fruit capsular, turbinate and somewhat flattened perpendicularly to septum, cartilaginous, apparently indehiscent. Seeds small, angled, pilosulous, shortly winged at one end. Venezuela, Guyana; 2 species, 1 in Venezuela. The second species of Merumea is known from the Merume mountains of Guyana. Both of these have been very rarely collected. Merumea coccocypseloides Steyerm., Mem. New York Bot. Gard. 23: 233, fig. 51. 1972. Young stems densely hirtellous; leaves 5– 7 × 2–4 cm; stipules 5–10 mm long; inflores-

Fig. 513. Merumea coccocypseloides

cences 1.5–2 × 2–2.5 cm; calyx lobes 3.5–9 mm long; corolla tube ca. 1.5 mm long, lobes ca. 3.5 mm long; fruit ca. 6 × 6 mm. Montane forested slopes along streams, ca. 1600 m; Amazonas (Cerro Sipapo). Endemic. ◆Fig. 513.

Mitracarpus 653

56. MITRACARPUS Zucc. ex Schult. & Schult. f., Mant. 210, 399. 1827. by Charlotte M. Taylor and Julian A. Steyermark Weedy annual or perennial herbs or low shrubs, terrestrial, unarmed. Leaves opposite, subsessile or shortly petiolate, venation not lineolate; stipules interpetiolar and fused to the petiole bases, persistent, truncate to broadly rounded, setose, generally erect and flatly appressed in bud. Inflorescences terminal and/or axillary, capitate or glomerulate, sessile, multiflowered, bracteate. Flowers small, homostylous, protandrous, sessile. Hypanthium turbinate to subglobose. Calyx limb deeply 4(5)-lobed, the lobes equal or usually unequal in pairs with 2 shorter and 2 longer, without calycophylls; corolla salverform to funnelform, white, internally glabrous or variously pubescent, lobes 4, valvate in bud. Stamens 4, inserted in the corolla throat; anthers narrowly oblong, dorsifixed, included or exserted. Ovary 2locular; ovules solitary in each locule, axile. Fruit capsular, ellipsoid to subglobose, papery to cartilaginous, circumscissile at the equator. Seeds small, ellipsoid-oblong to depressed-ovoid, ventrally with an impressed cruciform scar. Mexico, Central America, West Indies, South America, and introduced in Africa, Madagascar, and Asia; ca. 30 species, 6 in Venezuela, all in the flora area. Mitracarpus is clearly separated from other members of the tribe Spermacoceae by its circumscissile fruits and seeds with a cruciform or X-shaped sulcus on the adaxial face. Plants of Mitracarpus without mature fruits are frequently confused with Borreria and Spermacoce, but most species of Mitracarpus are recognizable by their calyx lobes, which are relatively well developed, at least a little unequal in pairs, and scabrid and often whitened along their margins. Because of the difficulty of separating flowering specimens of these genera, the species of Mitracarpus known from the flora region are included not only in the key to this genus below but also in the key to Borreria. Key to the Species of Mitracarpus 1. 1.

2(1).

2. 3(1). 3. 4(3). 4. 5(4).

5.

Slender annual herbs to 10 cm tall; leaves 4–26 × 0.5–5 mm ................. 2 Taller, more robust annuals, perennial herbs, or subshrubs to 150 cm tall, generally flowering when 15 cm tall or taller; leaves 7–80 × 3–20 mm ................................................................................................................ 3 Leaves glabrous or sometimes scabrous adaxially; corolla with the tube ca. 2 mm long, the lobes thickened or shortly horned abaxially .......................................................................................... M. microspermus Leaves hirsutulous to pilose and scabrose on both surfaces; corolla with the tube 1–1.5 mm long, the lobes smooth abaxially ............. M. parvulus Stipules with 1–3 narrowly triangular lobes 2–5 mm long; calyx lobes 3– 5 mm long ........................................................................................ M. sp. A Stipules with 3–7 linear setae 1–4 mm long; calyx lobes 1–2.5 mm long ... 4 Corolla 3.5–6.5 mm long ................................................................ M. frigidus Corolla 1–3 mm long .................................................................................. 5 Corolla ca. 1.2 mm long, shorter than the longer pair of calyx lobes; shorter pair of calyx lobes 0.8–1 mm long; stems pubescent with uniformly sub-appressed upcurved short trichomes ..................... M. diffusus Corolla 2–2.5 mm long, equaling or a little exceeding the longer pair of calyx lobes; shorter pair of calyx lobes ca. 1.5 mm long; stems pubescent with short trichomes on the sides and long spreading trichomes on the angles ...................................................................................... M. hirtus

654

R UBIACEAE

Mitracarpus diffusus (Willd. ex Roem. & Schult.) Cham. & Schltdl., Linnaea 3: 1828. —Spermacoce diffusa Willd. ex Roem. & Schult., Syst. Veg. 3: 531. 1818. Suffrutescent plant to 35 cm tall; leaves 7–30 × 3–7 mm, scaberulous to glabrescent on both surfaces; secondary veins not visible or 1 or 2 pairs; stipule with sheath 1–2 mm long, setae 3–5, 1.5–4.5 mm long; glomerules 3–7 mm diameter; calyx lobes 4, 1–1.5 mm long, unequal; corolla tube 0.8–1 mm long, lobes 0.3–0.5 mm long; capsules ca. 0.8 × 0.8 mm. Sandy banks along streams, wet open ground, 50–200 m; Amazonas (southwestern part of basin of Río Atabapo and Río Orinoco). Apure, Barinas, Guárico; adjacent Colombia. ◆Fig. 515. Mitracarpus frigidus (Willd. ex Roem. & Schult.) K. Schum. in Mart., Fl. Bras. 6(6): 81. 1888. —Spermacoce frigida Willd. ex Roem. & Schult., Syst. Veg. 3: 531. 1818. Colombia, Venezuela, Guyana, Peru, Brazil, Paraguay; 10 varieties, 3 in Venezuela, 1 of these in the flora area. Mitracarpus frigidus is very similar to M. scabrellus Benth. of Guyana and adjacent Brazil. Mitracarpus scabrellus has narrower, smaller leaves, 7–28 × 1–5 mm, and corollas ca. 3 mm long, and grows in open savannas. The other varieties of Mitracarpus frigidus reported from Venezuela are var. frigidus, distinguished from the other varities by its pubescent stipule sheaths, and var. andinus Steyerm., distinguished from the other varieties by its glabrous leaves, stems, and stipule. Both of these varieties are reported from 1400–2600 m in the Coastal Cordillera and northern Andes. The pubescence patterns used to separate these have been found to be continuous in some other Rubiaceae species, and these two varieties may not actually be distinct. Var. orinocensis of the flora region differs significantly from these two varieties of M. frigidus in its markedly lower elevational distribution, and its relationship to M. frigidus probably deserves re-evalution also. M. frigidus var. orinocensis Steyerm., Mem. New York Bot. Gard. 23: 780. 1972. Herb or subshrub to 0.5 m tall; leaves 10– 70 × 3–20 mm, scaberulous adaxially, pilosulous or hirtellous to glabrescent abaxially;

secondary veins 2–4 pairs; stipules with sheath 1.5–2 mm long, setae 5–9, 1.5–4 mm long; glomerules 8–15 mm diameter; calyx lobes 4, 1–2 mm long, unequal; corolla tube 2.5–3 mm long, lobes 1–1.2 mm long; capsules 1–1.5 × 1–1.5 mm. Sandy banks of streams bordering white-sand savannas, 100– 200 m; Amazonas (basin of Río Atabapo). Endemic. ◆Fig. 516. Mitracarpus hirtus (L.) DC., Prodr. 4: 572. 1830. —Spermacoce hirta L., Sp. Pl. ed. 2, 1: 148. 1762. Mitracarpus villosus (Sw.) DC., Prodr. 4: 572. 1830. —Spermacoce villosa Sw., Prodr. 29. 1788. Annual herb to 40 cm tall; leaves 10–80 × 5–20 mm, tomentulose to hirsute or glabrescent on both surfaces; secondary veins 3–6 pairs; stipules with sheath 1–4 mm long, setae 3–9, 1–5 mm long; glomerules 5–20 mm diameter; calyx lobes 4, 1–2.5 mm long, unequal; corolla tube 1.5–2 mm long, lobes 0.5– 1 mm long; capsules 2–3.5 mm diameter. Igneous outcrops, upland rocky savannas, waste places, 50–500 m; Bolívar (Altiplanicie de Nuria, near Upata and Puerto Ordaz), Amazonas (near Puerto Ayacucho and Isla Ratón, basin of Río Orinoco). Anzoátegui, Aragua, Carabobo, Cojedes, Falcón, Guárico, Mérida, Portuguesa, Trujillo, Zulia; widely distributed in Mexico, the West Indies, Central America, Colombia, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 514. Mitracarpus hirtus is variable morphologically, particularly in the size of the plant and its leaves, which apparently vary to a significant degree in relation to available water. There has been much controversy as to the correct name for this species, due to disagreement about the identity of Linnaeus’s name. However most recent authors have agreed that the names Spermacoce hirta and S. villosa do in fact apply to the same species (e.g., Nicolson 1977). Mitracarpus microspermus K. Schum. in Mart., Fl. Bras. 6(6): 83. 1888. Herbaceous annual to 10 cm tall; leaves 4–16 × 0.5–3 mm, scabrous adaxially, glabrous abaxially; secondary veins not evident or 1 pair; stipules with sheath 1–1.5 mm long, setae 5–7, 1–2 mm long; glomerules 1–5 mm diameter; calyx lobes 4, 1.5–3 mm long, unequal; corolla tube ca. 2 mm long, lobes 0.5–1 mm long; fruit ca. 1 × 1.2 mm. Sandy

Mitracarpus 655

savannas, tepui slopes, 400– 1100 m; Bolívar (Auyán-tepui, Salto Hacha). Guyana, Suriname, French Guiana, Brazil. Mitracarpus parvulus K. Schum. in Mart., Fl. Bras. 6(6): 83. 1888. Herbaceous annual to 10 cm tall; leaves 4–26 × 1–5 mm, hirsutulous to pilose and scabrous on both sides; secondary veins not visible; stipule with sheath 1–1.5 mm long, setae 3–7, 1–5 mm long; glomerules 3–5 mm diameter; calyx lobes 4, 1–2 mm long, unequal; corolla tube 1–1.5 mm long, lobes 0.5– 0.8 mm long; capsules ca. 1 × 1 mm; seeds ca. 0.5 mm diameter. Igneous outcrops along rivers and rocky sandstone savannas, 50–500 m; Bolívar (Altiplanicie de Nuria), Amazonas (Raudal de Atures, Río Orinoco). Guárico, Yaracuy; Guyana, Brazil. ◆Fig. 517. Fig. 514. Mitracarpus hirtus

Mitracarpus sp. A Shrub to 1.5 m tall; leaves 30–65 × 5–10 mm, smooth and glabrous on both surfaces; secondary veins 3 or 4 pairs; stipules with sheath 3–4 mm long, lobes 1–3, narrowly triangular, 2–5 mm long; calyx lobes 4, 3–5 mm long, unequal; corolla tube ca. 3 mm long, lobes ca. 2 mm long; capsules ca. 1.5 × 1.5 mm. Open thickets bordering streams, 50– 100 m; Amazonas (Río Sipapo). Endemic. Mitracarpus sp. A cannot be conclusively identified at present because of the poor knowledge of Mitracarpus, but may belong to a species described from Brazil. Fig. 516. Mitracarpus frigidus var. orinocensis

Fig. 515. Mitracarpus diffusus

Fig. 517. Mitracarpus parvulus

656

R UBIACEAE

57. MORINDA L., Sp. Pl. 176. 1753. Appunia Hook. f. in Benth. & Hook. f., Gen. Pl. 2: 120. 1873. Bellynkxia Müll. Arg., Flora 48: 465. 1875. by Charlotte M. Taylor and Julian A. Steyermark Trees or shrubs (in the Paleotropics also climbers), terrestrial, sometimes armed with spines. Leaves opposite or ternate, subsessile to petiolate, venation not lineolate; stipules interpetiolar or usually shortly united around the stem, persistent, triangular to shortly bilobed, in bud generally imbricate to valvate. Inflorescences axillary and/or terminal, multiflowered, capitate, with the flowers free or fused by their ovaries, the heads 1 to various, sessile to pedunculate or paniculate, bracteate or the bracts reduced. Flowers sessile, distylous, medium-sized. Hypanthium when free turbinate to ellipsoid. Calyx limb truncate to sinuate, without calycophylls; corolla salverform to funnelform, white, internally, usually pubescent in the tube, lobes 4 or 5(–7), valvate in bud, sometimes fleshy and triangular in cross section. Stamens 4 or 5(–7), inserted in the corolla tube or throat; anthers narrowly ellipsoid, included or partially exserted, dorsifixed near middle, sometimes with connective prolonged into a short apical appendage. Ovary 2- or 4-locular; ovules axile, 2 in each of 2 locules or 1 in each of 4 locules. Fruits drupaceous, fleshy, blue-black, free and ellipsoid to subglobose or fused into a subglobose multiple fruit; pyrenes 2 or 4, cartilaginous to bony, 1-locular, each with one seed. Seeds small, angled. Tropical Mexico, Central America, the Antilles, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Bolivia, Brazil, Old World tropics; ca. 90 species, 7 in Venezuela, 4 of these in the flora area. The genera Morinda and Appunia were distinguished by the arrangement of their flowers, with those of each inflorescence head all fused producing a single multiple fruit in Morinda versus the flowers all free with each producing a free drupe in Appunia. Additionally these were separated by their stigmas, reportedly 2 and linear in Morinda versus solitary and clavate in Appunia. Steyermark (1972, 384–386) reviewed the relationships between these genera, and noted that paired stigmas are found in several species with free fruits (e.g., A. triphylla Ducke) and that several species have fruits that are “coherent only in their basal half,” notably M. hoffmannioides Standl., which he considered to be intermediate between these genera. Accordingly Steyermark formally synonymized these genera, as done here. However, this conclusion has not been accepted by some authors (e.g., Taylor 2001a), who consider the single versus multiple fruits to be a clear and apparently meaningful taxonomic distinction. In the fruits of M. hoffmannioides the individual fruits are evident, and it is unclear whether they are closely set or actually fused at the base in that they appear to be dispersed separately in either case. Variation in stigma form is found in some other Rubiaceae genera (e.g., Borreria), and as shown by Steyermark this character is not correlated with fruit type. The “Appunia” group is found almost throughout the neotropical range of Morinda but has its greatest concentration of species in northeastern South America; all the species in the flora region have free, separate fruits. Two species of the “Morinda” group might be found in the flora area: the pantropical species M. citrifolia L. grows naturally on tropical sea coasts but is infrequently also cultivated, and the Central American species M. panamensis Seem. is occasionally cultivated for its edible fruits.

Morinda 657

Steyermark (1972) set much weight on details of the pubescence of the leaves, peduncles, calyx limbs, and corollas to circumscribe species of Morinda sensu lato. However this character has not been well evaluated as to its variability in local populations, and does seem to be widely variable in some species of Morinda (e.g., M. royoc L., and David Lorence personal communication). Key to the Species of Morinda 1. 1. 2(1). 2. 3(2). 3.

Peduncles ca. 20 cm long; leaves 17–30 cm long, with secondary veins 9 or 10 pairs ....................................................................... M. longipedunculata Peduncles 0.2–8.5 cm long; leaves 7–20 cm long, with secondary veins 5– 9 pairs ..................................................................................................... 2 Flowers free; peduncles 4–8.5 cm long, terminating in a single head or sometimes 3 closely grouped heads ................................. M. peduncularis Flowers free to partially coherent or fused; peduncles 0.1–6 cm long, terminating in 1–5 heads ........................................................................... 3 Anthers at the apex with an acute, lanceolate, prominent appendage; peduncles 18–55 mm long ...................................................... M. tenuiflora Anthers not appendiculate at the apex; peduncles 6–15 mm long ........................................................................................... M. venezuelensis

Morinda longipedunculata Steyerm., Ann. Missouri Bot. Gard. 75: 340, fig. 9. 1988. Shrub ca. 2 m tall; leaves 17–30 × 6–8 cm; secondary veins 9 or 10 pairs; petioles 1.5–2 cm long; stipules deltoid to deeply bifid, ca. 4 mm long; inflorescences pseudoaxillary, peduncles ca. 20 cm long, terminating in a pair of bracteate leaves, these 7–9 × 3–4 cm; heads 3 or 4, fasciculate; bracts ca. 2 mm long, shallowly 3- or 4-lobed; flowers free; calyx limb reduced; corolla in bud with color not noted, tube ca. 7.5 mm long, lobes ca. 0.5 mm long; fruits free, subglobose, 7–8 × 8–8.5 mm, dark red. Wet lower montane forests, 300–400 m; Amazonas (Sierra de la Neblina). Eastern Colombia. Morinda debilis (Sandwith) Steyerm. of Guyana has similarly long peduncles and large leaves, but it can be distinguished from M. longipedunculata by its calyx limbs 1.5–2 mm long and its corolla lobes ca. 6 mm long, as long as or longer than the corolla tube. Morinda peduncularis Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 3: 380. 1818 [1819]. Ixora angulata Benth. in Benth. & Hook. f., Gen. Pl. 2: 114. 1873. —Bellynkxia angulata (Benth.) Müll. Arg., Flora 58: 465, 475. 1875. —Appunia angulata (Benth.) Baill., Hist. Pl. 7: 415. 1880. Shrub to 6 m tall, sometimes weakstemmed; leaves 13–16 × 4–10.5 cm; second-

ary veins 5–9 pairs; petioles 5–17 mm long; stipules 1.5–3 mm long; peduncles 4–8.5 cm long; inflorescence with 1 head; flowers free; calyx limbs 1.5–2 mm long; corolla in bud to 1 cm long; fruits ca. 1 cm diameter. Riparian and evergreen lowland forests, 100–200 m; Amazonas (western part of basin of Río Casiquiare and Río Orinoco, Río Sipapo). Apure. ◆Fig. 518. Steyermark (1974) treated Mapouria spruceana Müll. Arg. as a synonym of M. pedunculata, but as noted elsewhere in that same work the specimen shown in the photo of the type collections of M. spruceana (Spruce 3343, C; Rockefeller neg. #22855, MO!) is Rudgea laurifolia. Morinda tenuiflora (Benth.) Steyerm., Mem. New York Bot. Gard. 23: 386. 1972. —Coffea tenuiflora Benth., J. Bot. (Hooker) 3: 232. 1841. —Appunia tenuiflora (Benth.) Jacks. & Hook. f., Kew Index 1: 166. 1893. [incorrectly attributed to Benth. & Hooker f., Gen. Pl. 2: 120. 1873]. —Cafecillo, Ciruela de playa, Guayabita. Appunia tenuiflora var. leiophylla Steyerm., Mem. New York Bot. Gard. 17: 358. 1967. —Morinda tenuiflora var. leiophylla (Steyerm.) Steyerm., Mem. New York Bot. Gard. 23: 386. 1972. Shrub or small tree to 4 m tall; leaves 7– 20 × 2.5–10 cm; secondary veins 5–9 pairs;

658

R UBIACEAE

Fig. 518. Morinda peduncularis

petioles 2–10 mm long; stipules 1.5–3 mm long; inflorescences with 1–5 heads; peduncles 18–55 mm long; flowers free or very shortly fused or coherent at base; calyx limb 0.5–1.5 mm long; corolla with tube 5.5–9 mm long, lobes 4–7 mm long; fruits 7–8 mm long. Riparian forests, evergreen seasonally dry forests, rocky forested dry slopes, borders of savannas and forests, open tree savannas with Caraipa and Platycarpum, bases of igneous rocks, floodplain forests, 50–200 m; Bolívar (common), Amazonas (common in Río Orinoco basin). Apure, Lara, Zulia; Guyana, Brazil. ◆Fig. 519. Steyermark (1967; 1974) recognized two varieties of Morinda tenuiflora, distinguished by pubescence characters: puberulent or pilose-puberulous on leaf lower surfaces, stipules, and peduncles in var. tenui-

flora, versus glabrous or sparsely puberulous on those structures in var. leiophylla. Because the variation is continuous between the two, separate varieties are not recognized here. Morinda tenuiflora is quite variable, particularly in leaf size, peduncle length, and number of flower heads per peduncle. This species typically grows in the edges of thickets and gallery forests in savannas, and no doubt variation in available water is responsible for much of this morphological variation. Morinda surinamensis (Bremek.) Steyerm. is very similar to M. tenuiflora and perhaps not actually distinct; Steyermark distinguished M. surinamensis by its fully glabrous vegetative and reproductive structures and its range in Suriname rather than Venezuela, Guyana, or Brazil.

Neblinathamnus 659

Morinda venezuelensis (Steyerm.) Steyerm., Mem. New York Bot. Gard. 23: 386. 1972. —Appunia venezuelensis Steyerm., Mem. New York Bot. Gard. 17(1): 356. 1967. Tree to 5 m tall; leaves 11–17.5 × 3–8 cm; secondary veins 5 or 6 pairs; petioles 3–5 mm long; stipules 2.5–3 mm long; inflorescences with 1–3 heads; peduncles 6–15 mm long; calyx limbs 0.5–0.7 mm long; corolla with tube 8–8.5 mm long; lobes 5–6 mm long; fruits 9– 10 × 7–8 mm. Edges of Masutiyia palm swamps, semideciduous forests, 400–600 m; Bolívar (northeastern Altiplanicie de Nuria). Endemic. Two species of Morinda known from the Guianas but not reported from the flora area have the flowers in 1–several heads that are

subsessile to fascicled on peduncles to 2 cm long. Morinda calycina (Benth.) Steyerm. can be recognized by its calyx limbs 3–5 mm long, while M. brachycalyx (Bremek.) Steyerm. can be recognized by its membranaceous acuminate leaves with the tips rather long and slender and its stipules 4–6 mm long.

Fig. 519. Morinda tenuiflora

58. NEBLINATHAMNUS Steyerm., Mem. New York Bot. Gard. 10(5): 229. 1963. by Piero G. Delprete and Julian A. Steyermark Shrubs, terrestrial, unarmed, erect. Leaves 3-verticillate, relatively small, subsessile to petiolate, tessellate beneath, venation not lineolate; stipules reduced,

660

R UBIACEAE

interpetiolar, truncate or shortly bilobed with broadly triangular lobes, persistent, in bud erect and flatly appressed. Inflorescences terminal and in the uppermost leaf axils, pedunculate, bracteate with the bracts setaceous, with flowers solitary or 2 and fasciculate. Flowers medium-sized, pedicellate, homostylous, protandrous, bibracteolate with the bract setaceous. Hypanthium ellipsoid to turbinate. Calyx limb deeply lobed, lobes 5, linear or spatulate, equal or unequal but without calycophylls; corolla salverform, salmon to brick red on tube and pale green to yellow green on lobes, internally pubescent in the upper part of the tube, lobes 5, contorted in bud. Stamens 5, inserted near middle of the corolla tube; anthers narrowly oblong to linear, dorsifixed, included. Ovary 2-locular; ovules numerous in each locule, on axile placentas. Fruit capsular, obovoid, loculicidal from the base, woody. Seeds small, angulate, reticulate-rugose. Venezuela, Brazil; 2 species, both in the flora area. Key to the Species of Neblinathamnus 1. 1.

Leaves 1–2.5(–3.3) × 0.5–1.3 cm; corolla tube 14–16 mm long ... N. argyreus Leaves 3.5–6 × 1–3 cm; corolla tube ca. 12 mm long .............. N. brasiliensis

Neblinathamnus argyreus Steyerm., Mem. New York Bot. Gard. 10(5): 229, fig. 77. 1963. Neblinathamnus glabratus Steyerm., Mem. New York Bot. Gard. 10(5): 231. 1963. Shrub 0.3–2 m tall; leaves 1–2.5(–3.3) × 0.5–1.3 cm; calyx lobes 4–8 mm long; corolla with tube 14–16 mm long, the lobes 5.5–6 × 4.5–5 mm; capsules 4–5 × 3.5–4 mm. Open scrub forests, escarpments and summits of tepuis, 1700–1900 m; Amazonas (Sierra de la Neblina). Endemic. ◆Fig. 520. Steyermark distinguished Neblinathamnus glabratus by details of its leaf shape and pubescence and by the absence of vermiform trichomes on its lower leaf surface. However, as he predicted in his description of N. glabratus, recent collections have shown that these characters intergrade. Consequently these names are here considered as synonymous. Neblinathamnus brasiliensis Steyerm., Mem. New York Bot. Gard. 23: 309. 1972. Neblinathamnus aracamunianus Steyerm., Ann. Missouri Bot. Gard. 76: 968. 1989. Shrub 0.5–1 m tall; leaves 3.5–6 × 1–3 cm; calyx lobes 4–7 mm long; corolla with tube ca. 12 mm long, lobes 5–6 × 3–3.7 mm; capsules ca. 5 × 4 mm. Meadows along escarpment on tepui summits, 1500–1600 m; Amazonas (Cerro Aracamuni). Brazil (Serra Pirapucú).

Fig. 520. Neblinathamnus argyreus

Notopleura 661

59. NOTOPLEURA (Benth. & Hook. f.) Bremek., Recueil Trav. Bot. Néerl. 31: 289. 1934. —Psychotria sect. Notopleura Benth. & Hook. f., Gen. Pl. 2: 124. 1873. by Charlotte M. Taylor Herbs or subshrubs, unarmed, rather succulent, terrestrial or epiphytic, the terrestrial species usually unbranched. Leaves opposite, petiolate, venation not lineolate; stipules persistent to caducous, generally valvate to flatly appressed in bud, united around stem at least shortly, the interpetiolar sheaths truncate to triangular, smooth or usually with 1–several appendages attached at center or base, these appendages persistent or caducous, conical to linear or sometimes expanded into a flattened blade. Inflorescences terminal and/or pseudoaxillary, multiflowered, capitate to congested-cymose or paniculate, sessile to pedunculate, bracteate. Flowers sessile to pedicellate, generally small, protandrous, distylous or homostylous. Hypanthium subglobose to turbinate. Calyx limb reduced or developed, truncate to lobed, lobes 4 or 5, without calycophylls; corolla funnelform to tubular, white to pale yellow, orange, or purple, glabrous or variously pubescent internally, lobes 4–6, valvate in bud, often dorsally thickened to horned. Stamens 4 or 5, inserted near or above middle of corolla tube; anthers narrowly oblong, exserted or included, dorsifixed near middle. Ovary 2–4(–6)-locular; ovules 1 in each locule, basal. Fruit subglobose to ellipsoid, drupaceous, carnose or spongy, white, orange, red, or black; pyrenes 2–4(–6), chartaceous to bony, 1-locular, hemispherical to triangular. Seeds ellipsoid, small. Mexico, Central America, the Antilles, Colombia, Venezuela, Trinidad-Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Bolivia, Brazil, with most species in wet forests in the Andes and southern Central America; ca. 100 species, 23 in Venezuela, 8 of these in the flora area. This genus was included by most previous authors in Psychotria, but as discussed by Taylor (2001b) both morphological and molecular sequence evidence support its separation. Notopleura comprises two subgenera, both found in the flora area: subgenus Notopleura includes terrestrial, usually unbranched plants, while subgenus Viscagoga (Baill.) C.M. Taylor includes the epiphytic species, which are usually branched. Plants of subgenus Viscagoga have been rather infrequently collected in the flora area, but as with most epiphytes they are probably more common than shown by collection frequency. Several species recognized by Steyermark are here treated provisionally; their distinctions are rather minor and when more collections are available these may be found to intergrade and be better combined. In many species the maturing fruits pass through a red stage and then become black and are usually removed rapidly from the plant by frugivores. This black fruit stage is often not found by collectors and overlooked in descriptions. Key to the Species of Notopleura 1. 1. 2(1). 2.

Plants terrestrial, usually unbranched; inflorescences pseudoaxillary ...... 2 Plants epiphytic or hemiepiphytic, branched; inflorescences terminal and/or pseudoaxillary ............................................................................ 5 Leaves elliptic to narrowly elliptic, with the secondary veins not visible on the abaxial surface including when dry .......................................... 3 Leaves elliptic to broadly so, with the secondary veins visible on the abaxial surface ....................................................................................... 4

662

3(2). 3. 4(2). 4. 5(1). 5. 6(5). 6. 7(6). 7.

R UBIACEAE

Inflorescences capitate, surrounded by involucral bracts ...... N. sandwithiana Inflorescences branched to several orders, bracteate but the bracts not involucral ............................................................................ N. tapajozensis Inflorescences capitate or subcapitate, subsessile .............. N. thesceloantha Inflorescences branched to several orders, pedunculate ............ N. uliginosa Calyx limb truncate to undulate ........................................... N. multiramosa Calyx limb lobed ........................................................................................ 6 Calyx lobes triangular to narrowly triangular, acute to acuminate, 0.5– 1.5 mm long; corolla 6–7 mm long ............................................... N. crassa Calyx lobes broadly triangular, obtuse to deltoid, 0.2–0.8 mm long; corolla 3–5 mm long .................................................................................. 7 Calyx limb ciliolate and corolla externally smooth ........................ N. aligera Calyx limb entire and corolla externally papillose-puberulous ................. ........................................................................................... N. perpapillifera

Notopleura aligera (Steyerm.) C.M. Taylor, Ann. Missouri Bot. Gard. 88: 512. 2001. —Psychotria aligera Steyerm., Mem. New York Bot. Gard. 23: 572, fig. 75. 1972. Fleshy epiphyte; leaves 4–5 × 1.5–1.8 cm; stipules 1–1.5 mm long, truncate; inflorescences 1.5–2.5 × 2–4 cm; calyx limb ca. 0.8 mm long; corolla white, tube ca. 2 mm long, lobes 4, ca. 1 mm long; fruit unknown. Wet montane and Clusia forests over sandstone substrate, 700–1900 m; Bolívar (basin of Río Caroní and Río Cuyuní, southeastern portion west to Aparda-tepui). Guyana. Notopleura crassa (Benth.) C.M. Taylor, Ann. Missouri Bot. Gard. 88: 512. 2001. —Psychotria crassa Benth., J. Bot. (Hooker) 3: 227. 1841. Psychotria crassa f. alba Steyerm., Ann. Missouri Bot. Gard. 71: 331. 1984. Psychotria crassa f. angustior Steyerm., Ann. Missouri Bot. Gard. 71: 331. 1984. Fleshy epiphyte or hemiepiphyte; leaves 2–11.5 × 1–4 cm; stipule 1.5–3.5 mm long, truncate; inflorescences 2–9 × 2–8 cm; calyx limb 1–2 mm long; corolla white to pink, tube 4–8 mm long, lobes 4, 1.5–2.5 mm long; fruit 6–7 mm diameter, becoming red then black. Recumbent mossy forests, Bonnetia roraimae forests, dwarf and wet forests, riparian forests, in sandstone crevasses, Schefflera groves, and groves of Mallophyton, common on the slopes and summits of sandstone table mountains, (400–)1600–2600 m;. Bolívar (Cerro Jaua, Ilú-tepui, Kukenán-tepui, Macizo del Chimantá, Roraima-tepui, Urimán), Amazonas (Cerro Duida, Cerro Huachamacari, Cerro Marahuaka, Cerro Sipapo,

Cerro Yaví, Sierra de la Neblina). Guyana, adjacent Brazil. Steyermark described several infraspecific taxa for this species at the level of form, but their differences are incomplete and minor so these were synonymized by Taylor (2001b). Notopleura multiramosa (Steyerm.) C.M. Taylor, Ann. Missouri Bot. Gard. 88: 573. 2001. —Psychotria multiramosa Steyerm., Mem. New York Bot. Gard. 23: 568. 1972. Fleshy epiphyte; leaves 6–10 × 2.5–5 cm; stipules 1–2 mm long, truncate; inflorescences 4.5–7 × 8–13 cm; calyx limb 0.5–0.8 mm long; corolla white to pink, tube 2.5–3 mm long, lobes 4, 1.2–1.8 mm long; fruit 5–6 × 2.5–3 mm, becoming red then black. Tepui slope forests, ca. 1300 m; Bolívar (Macizo del Chimantá). Guyana. Notopleura perpapillifera (Steyerm.) C.M. Taylor, Ann. Missouri Bot. Gard. 88: 513. 2001. —Psychotria perpapillifera Steyerm., Mem. New York Bot. Gard. 23: 571. 1972. —Psychotria pendula subsp. trinitensis Urb., Symb. Ant. 1: 447. 1900, not Psychotria trinitensis Urb. 1913. Fleshy epiphyte; leaves 4–7.5 × 2–4 cm; stipules 1.5–2 mm long, truncate; inflorescences 1–5 × 2.5–12 cm; calyx limb 0.5–1.5 mm long; corolla white, tube 3–4 mm long, lobes 5 or 6, 2–2.5 mm long; fruit 4–4.5 × 2.8– 3 mm, becoming red then black. Semideciduous lower montane forests, 600–700 m; Bolívar (Altiplanicie de Nuria). Monagas, Sucre; Trinidad, Tobago, Guyana.

Notopleura 663

Notopleura sandwithiana (Steyerm.) C.M. Taylor, comb. nov. —Cephaelis sandwithiana Steyerm., Mem. New York Bot. Gard. 17(1): 428, fig. 4. 1967. —Psychotria sandwithiana (Steyerm.) Steyerm., Mem. New York Bot. Gard. 23: 555. 1972. Fleshy terrestrial herb to 2.5 m tall; leaves 26–37 × 10–12 cm; stipules 2–3 mm long, truncate to broadly rounded; peduncle ca. 4.5 cm long; inflorescence capitate, ca. 2.8 × 1.7 cm; involucral bracts white, ca. 2 × 2.5 cm; calyx limb ca. 2 mm long; corolla white, tube 9–11 mm long, lobes 4, ca. 2 mm long; fruit unknown. Semideciduous slope forests, ca. 400 m; Bolívar (Cerro Pitón). Endemic. Notopleura tapajozensis (Standl.) Bremek., Recueil Trav. Bot. Néerl. 34: 290. 1934. —Psychotria tapajozensis Standl., Publ. Field Columbian Mus., Bot. Ser. 8: 215. 1930. Psychotria amplinoda Steyerm., Mem. New York Bot. Gard. 17(1): 431. 1967. Psychotria amplinoda var. ayangannae Steyerm., Mem. New York Bot. Gard. 17(1): 433. 1967. Fleshy terrestrial to 4 m tall; leaves 8–34 × 2.5–8 cm; stipules 2.5–3.5 mm long, truncate; inflorescences with peduncles 4–10 cm long, branched portion 2.5–4.5 × 3–6 cm; calyx limb 0.5–1 mm long; corolla white to lilac, tube ca. 7 m long, lobes 5, ca. 3 mm long; fruit 8–9 × 7–8 mm, becoming red-purple then black. Frequent on the sandstone table mountains in wet montane forests, forested

Fig. 521. Notopleura tapajozensis

slopes along streamlets, dwarf Bonnetia roraimae forests, 1100–2300 m; Bolívar (Amaruay-tepui, Aprada-tepui, Auyán-tepui, Carrao-tepui, Cerro Guaiquinima, Cerro Venamo, Ilú-tepui, Kukenán-tepui, Macizo del Chimantá, Ptari-tepui, Serranía Marutaní, Sierra de Lema), Amazonas (Cerro Sipapo, Cerro Yapacana, Sierra de la Neblina). Amazonian Colombia, Guyana, Suriname, French Guiana, northeastern Brazil. ◆Fig. 521. Notopleura thesceloantha (Steyerm.) C.M. Taylor, Ann. Missouri Bot. Gard. 88: 509. 2001. —Psychotria thesceloantha Steyerm., Ann. Missouri Bot. Gard. 75: 347, fig. 13. 1988. Fleshy terrestrial to 1 m tall; leaves 21–24 × 10–11 cm; stipules ca. 5 mm long, broadly triangular; inflorescence ca. 2 × 4 cm; calyx limb 2–4 mm long; corolla and mature fruit unknown. Evergreen lowland forests, swampy areas, 100–200 m; Amazonas (upper Río Baría below base of Sierra de la Neblina). Colombia, Ecuador. Notopleura uliginosa (Sw.) Bremek., Recueil Trav. Bot. Néerl. 31: 289. 1934. —Psychotria uliginosa Sw., Prodr. Veg. Ind. Occ. 43. 1788. —Uragoga uliginosa (Sw.) Kuntze, Revis. Gen. Pl. 2: 963. 1891. —Ají de gato. Fleshy terrestrial to 1.5 m tall; leaves 17– 35 × 5–14.5 cm; stipules 3–6 mm long, broadly triangular; inflorescences with peduncle 2.5–1.5 cm long, branched portion

664

R UBIACEAE

2.5–10 × 3–10 cm; calyx limb 0.5–1.5 mm long; corolla white to sometimes pink-purple, tube 2–3.5 mm long, lobes 5, 0.5–1.5 mm long; fruit 7–8 × 4.5–6 mm, becoming red then black. Evergreen lowland to montane forests, 50–1600 m; Delta Amacuro (lower

Río Orinoco, Serranía de Imataca), Bolívar (Altiplanicie de Nuria, Aprada-tepui, Macizo del Chimantá, basin of Río Cuyuní). Zulia and portions of the Coastal Cordillera of Venezuela west to Yaracuy; Mexico, Central America, the West Indies, Colombia.

60. OLDENLANDIA L., SP. PL. 119. 1753. by Charlotte M. Taylor and Julian A. Steyermark Annual or perennial herbs or low subshrubs, terrestrial, unarmed. Leaves opposite, subsessile to petiolate, venation not lineolate; stipules interpetiolar and often also fused to petiole bases, persistent or caducous, triangular, entire to bilobed or setose, generally valvate to imbricate in bud. Inflorescence terminal, pseudoaxillary, or axillary, fasciculate or cymose to paniculate, bracteate or the bracts reduced, few- to multiflowered or with flowers solitary. Flowers usually small, homostylous or distylous, protandrous, pedicellate. Hypanthium obconic to subglobose. Calyx limb with lobes 4, generally equal, without calycophylls; corolla rotate or funnelform, white to pink, internally glabrous or barbate in throat, lobes 4, valvate in bud. Stamens 4, inserted in the corolla throat; anthers dorsifixed, included or exserted, ellipsoid. Ovary 2-locular; ovules few to numerous in each locule, on axile placentas. Fruit capsular, generally subglobose, papery to chartaceous, often becoming prolonged into an apical beak above the insertion of the calyx limb, loculicidal in this beak and sometimes later septicidal. Seeds subglobose to angled, small, smooth to minutely alveolate or granular. Widespread in the New World and Old World tropics and subtropics; ca. 300 species, 4 in Venezuela, all in the flora area. Oldenlandia is related to the pantropical genus Hedyotis L. The distinctions between these genera are not completely clear, and Oldenlandia is sometimes combined under Hedyotis. Oldenlandia is sometimes confused with Perama. Perama can be separated by its corolla lobes that are convolute in bud and its capsules that are not apically beaked. Key to the Species of Oldenlandia 1. 1. 2(1). 2. 3(1). 3.

Leaves 2–12 mm wide ................................................................................ 2 Leaves 0.1–1 mm wide ............................................................................... 3 Flowers 2–5 in each leaf axil, each one pedicellate in a pedunculate cyme; capsules 1.5–2.5 mm diameter .............................................. O. corymbosa Flowers 1–3 in each leaf axil, individually pedunculate directly from the stem; capsules 2.5–4 mm diameter ......................................... O. lancifolia Branching of stems opposite; capsules 1.2–1.5 mm diameter; corollas 2– 3 mm long, 2 or more times as long as the calyx lobes ........... O. filicaulis Branching of stems mainly alternate; capsules 1.8–2 mm diameter; corollas 1–2 mm long, a little longer than the calyx lobes but < 2 times their length ............................................................................................. O. tenuis

Oldenlandia corymbosa L., Sp. Pl. 119. 1753. Slender annual; stems erect to prostrate; leaves 1–3.5 × 0.2–0.5 cm, subsessile; stipules with sheath 0.5–2 mm long, setae 3–5,

0.5–1.5 mm long; peduncles 3–10 mm long; pedicels 3–5 mm long; calyx lobes 0.5–1 mm long; corolla pink to white, tube 0.8–1 mm long, lobes ca. 1 mm long; capsules 1–1.5 × 1.5–2.5 mm. Open banks along streams, sa-

Oldenlandia 665

vannas, waste ground and disturbed areas, 50–400 m; Delta Amacuro (Sacupana), eastern Bolívar, Amazonas (Puerto Ayacucho, Santa Barbara del Orinoco). Aragua, Portuguesa, Táchira; apparently native to Africa, adventive and widespread in lowland and middle-elevation tropical regions worldwide. ◆Fig. 522. Oldenlandia filicaulis K. Schum. in Mart., Fl. Bras. 6(6): 271, pl. 127, fig. 2. 1889. Slender annual; stems erect; leaves 3–15 × 0.1–0.3 mm, subsessile; stipules ca. 0.2 mm long, triangular; flowers solitary in each leaf axil; peduncles 2–6 mm long; calyx lobes ca. 1 mm long; corolla white, 2–3 mm long; capsules 1.2–1.5 mm diameter. Igneous outcrops, ca. 100 m; Amazonas (near Puerto Ayacucho). Brazil (Bahia, Pará, Paraiba, Piauhy). The distinctions between Oldenlandia filicaulis and O. tenuis were detailed and illustrated by Steyermark (1988, 736–738).

One specimen from Venezuela (Huber et al. 1399, VEN) was provisionally referred to Oldenlandia filicaulis but said to be intermediate between this and O. tenuis by Steyermark (1988). This specimen reportedly has the relatively small capsules and opposite branching of O. filicaulis, together with the shorter corollas and sessile included stamens of O. tenuis. Oldenlandia lancifolia (Schumach.) DC., Prodr. 4: 425. 1830. —Hedyotis lancifolia Schumach., Beskr. Guin. Pl. 72. 1827. —Araña, Paja aguja, Sardina.

Fig. 523. Oldenlandia tenuis Fig. 522. Oldenlandia corymbosa

Fig. 524. Oldenlandia lancifolia

666

R UBIACEAE

Herbaceous annual or perennial; stem procumbent to suberect, sometimes rooting at nodes; leaves 1–7 × 0.15–0.8 cm, subsessile; stipules with sheath ca. 1 mm long, setae 2–4, 1.5–3 mm long; peduncles 5– 30 mm long; calyx lobes 1–1.5 mm long; corolla white to infrequently pink, tube ca. 1 mm long, lobes 1–1.5 mm long; capsules 2.5– 3 × 2.5–4 mm. Sandy stream banks, open areas near watercourses, white-sand areas with shrubs and small trees (bana formation), open rocky places, 50–600 m; Delta Amacuro (Sacupana, Serranía de Imataca), Bolívar (Río Cuyuní, Río Paragua, Serranía de Imataca), Amazonas (near Puerto Ayacucho, upper Río Orinoco, San Carlos). Apparently native to Africa, now adventive and widespread in lowland tropics worldwide. ◆Fig. 524. When dry the leaves of Oldenlandia lancifolia sometimes have their margins markedly revolute, thus appearing to be narrower than the measurements given here. Oldenlandia lancifolia was misidentified by several authors as O. herbacea (L.) Roxb.

However true O. herbacea is found only in East Africa and has not yet been reported as adventive in any other regions. Oldenlandia tenuis K. Schum. in Mart., Fl. Bras. 6(6): 273. 1889. Slender annual; stems erect; leaves 2–10 × 0.5–1 mm, subsessile; stipules 0.5–0.8 mm long, triangular to bilobed; flowers solitary in leaf axils; peduncles 3–15 mm long; calyx lobes ca. 1 mm long; corolla white, 1.8–2 mm long, lobes about equal to tube; capsules 1.8– 2 mm diameter. Moist rocky igneous savannas or open forested slopes over igneous substrate in deciduous forests, usually in wet depressions, 50–200 m; Bolívar (Cerro El Medano south of Caicara, Los Pijiguaos). Guyana, Brazil (Ceará, Piauhy, Roraima). ◆Fig. 523. The Bolívar specimens agree best with Oldenlandia tenuis, but this species is infrequently collected throughout its range and is poorly known. Oldenlandia tenuis is similar to O. filicaulis; see additional comments under that species.

61. PAGAMEA Aubl., Hist. Pl. Guiane 112, t. 44. 1775. by Alberto Vincentini & Julian A. Steyermark Shrubs or trees, unarmed, terrestrial. Leaves opposite, petiolate, membranaceous to coriaceous, glabrous to densely pubescent, venation not lineolate; stipules fused into a cylindrical sheath with 4–8 setae at the apex, usually only the upper terminal sheaths present, the lower deciduous. Inflorescences axillary or rarely pseudoterminal (i.e., in an apparently terminal head formed by a cluster of axillary inflorescences), pedunculate or rarely sessile, multiflowered and capitate, spicate, thyrsoid, paniculate, a compound cyme, or with flowers 1–3; bracts reduced, simple or lobed (leaf-like). Flowers small, distylous or homostylous, sessile to pedicellate. Hypanthium cup-shaped. Calyx truncate to lobed, lobes 4–6, without calycophylls; corolla rotate, campanulate or tubular, white, cream-colored, or greenish, 2–4.5 (–10) mm long and 1–1.5(–3.5) mm diameter at the base, the tube always glabrous inside, the lobes 4(5), densely pubescent adaxially (glabrous) with the hairs usually longer at their bases, valvate in bud. Stamens 4(5), inserted near top of corolla tube; anthers oblong to lanceolate, dorsifixed near the base, exserted. Ovary superior, 2locular; ovules solitary in each locule, ascending, basal or axile near the base. Fruit a fleshy drupe, globose to ellipsoid, 3.5–13 mm long, shiny black or purple-black at maturity, partially inserted in a cup-shaped calyx or completely exserted in a shallow, plate-like calyx, this fruiting calyx red (orange) at maturity. Pyrenes 2, hard, obovate to obdeltoid, with one seed each. Humid tropical South America: Colombia, Venezuela, French Guiana, Guyana, Suriname, Peru, Brazil, Bolivia; ca. 40 species, 17 in Venezuela, all in the flora area. Pagamea can be recognized by its distinctive stipules, which are tubular and setose. The ovaries and fruits are secondarily superior, i.e., this condition repre-

Pagamea 667

sents an evolutionary reversal from the inferior ovaries of the Rubiaceae. These structures often make it difficult to recognize Pagamea species as Rubiaceae. Key to the Species of Pagamea 1. 1. 2(1).

Leaves 12–30 × 3.5—8.5 cm ...................................................................... 2 Leaves 1–11 × 0.5–3.7 cm .......................................................................... 5 Lower surface of leaves glabrous or nearly so, with the trichomes minute (< 0.2 mm long); petiole 1/3–1/2 the length of the leaf blade; inflorescence 11–22 cm long, with the secondary axes (1.5–)3–5 cm long ...... P. coriacea 2. Lower surface of leaves hirsute or pilose, with the trichomes 0.5–4.5 mm long; petiole < 1/4 the length of the leaf blade; inflorescence 5–14 cm long, with the secondary axes to 1 cm long .......................................... 3 3(2). Upper surface of leaves glabrous or strigose along the midrib, the trichomes < 0.5 mm long; petiole (1.6–)2–3.8 cm long; leaves strongly coriaceous ...................................................................................... P. plicata 3. Upper surface of leaves hirsute or pilose, the trichomes 1–4.5 mm long; petiole 1–2 cm long; leaves membranaceous ........................................ 4 4(3). Leaves with 7–10 pairs of secondary veins and trichomes 2–4.5 mm long; inflorescence 7–12 cm long, the peduncle 5–7 cm long; ovary glabrous at apex .......................................................................................... P. hirsuta 4. Leaves with ca. 6 pairs of secondary veins and trichomes 1–1.5 mm long; inflorescence 6–7 cm long, the peduncle 3–4 cm long; ovary strigulose at apex ......................................................................................... P. velutina 5(1). Stipules persistent on all nodes with leaves .................................. P. capitata 5. Stipules soon caducous or their basal portions persistent ....................... 6 6(5). Inflorescence with 1–35 flowers, monocephalous or nearly so, i.e., the primary axis above the peduncle 0–5 mm long ......................................... 7 6. Inflorescence with 10–600 flowers, branched, the primary axis above the peduncle 5–130 mm long ..................................................................... 14 7(6). Inflorescence sessile and pseudoterminal, i.e., in an apparently terminal head formed by a cluster of sessile axillary inflorescences...... P. sessiliflora 7. Inflorescence pedunculate and axillary, the peduncle 0.4–4.5 cm long ... 8 8(7). Stipules 1.1–3.6 cm long; leaves obovate, 1.6–2.5 cm broad ....... P. jauaensis 8. Stipules 0.2–1 cm long; leaves elliptic, 0.6–1.5 cm broad ........................ 9 9(8). Corolla tube 5.5–7 mm long, 2–3 times as long as the lobes ... P. magniflora 9. Corolla tube 1–3 mm long, shorter than or at most as long as lobes .... 10 10(9). Barbate tufts present along lower midrib of leaf blade ......................... 11 10. No barbate tufts present along lower midrib of leaf blade .................... 13 11(10). Inflorescence with the peduncle > 2 cm long and the secondary axes 1.5– 4.5 mm long ................................................................................ P. montana 11. Inflorescence with the peduncle < 1.6 cm long and the secondary axes 0– 1.4 mm long .......................................................................................... 12 12(11). Stipule setae 1–2.5 mm long; petiole 5–9 mm long, corresponding to 1/3–1/5 the leaf blade length; twigs slender, 0.9–2 mm thick; calyx lobes absent or to 0.6 mm long; inflorescence usually 1-flowered or to 4-flowered .................................................................................................... P. duidana 12. Stipule setae < 1.2 mm long; petiole 2–6.5 mm long, corresponding to < 1/5 the leaf blade length; twigs stout, 1.4–2.5 mm thick; calyx lobes 0.4–

668

R UBIACEAE

13(10). 13. 14(6).

14. 15(14). 15. 16(15).

16.

17(15).

17.

18(17). 18. 19(18). 19.

20(18).

20. 21(20). 21.

1.8 mm long; inflorescence with 3–12 flowers or rarely 1–flowered .............................................................................................. P. standleyana Leaf margin strongly revolute; leaf blade broadly elliptic, 1.7–3 times longer than broad ....................................................................P. pauciflora Leaf margin flat or slightly revolute; leaf blade elliptic, 2.5–5 times longer than broad ...................................................................... P. montana Flowers and fruits distributed contiguously on the two sides of the distal part of primary axis, its internodes not elongated and indistinguishable ................................................................................................ P. diceras Flower and fruit clusters separated by internodes, the first internode above peduncle always > 5 mm long ................................................... 15 Upper surface of leaf blades conspicuously sulcate-veined, lower surface hirsute with the trichomes 0.7–1.6 mm long ...................................... 16 Upper surface of leaf blades not sulcate-veined, lower surface glabrous or if pubescent then with the trichomes 0.1–0.7 mm long ..................... 17 Leaf blade strongly coriaceous, 2–2.7 times longer than broad, with trichomes 0.7–1.1 mm long; petiole 1.5–4 cm long; stipule setae 0–2 mm long; twigs below nodes with leaves 5–9 mm thick ..................... P. plicata Leaf blade membranaceous to subcoriaceous, 2.6–3.5 times longer than broad, with trichomes 1.2–1.6 mm long; petiole 1–1.5 cm long; stipule setae 2–4.5 mm long; twigs below nodes with leaves < 4 mm thick ............................................................................................. P. plicatiformis Petiole 2–4.4 cm long, corresponding to 1/3–1/2 of the blade length; inflorescences 11–22 cm long, the primary axis above the peduncle 5–13 cm long, the peduncle 2.5–6 mm broad at base .............................. P. coriacea Petiole 0.2–2.2 cm long, corresponding to < 1/3 of the blade length; inflorescences 1.5–10 cm long, the primary axis above peduncle 0.5–4 cm long, the peduncle 1–2.2(–3) mm broad at base ................................. 18 Petiole 1–2.2 cm long; leaf blade 2–3.6 cm broad ................................... 19 Petiole 0.3–1 cm long; leaf blade 0.8–2.2 cm broad ................................ 20 Calyx densely pubescent inside, truncate or with lobes to 0.4 mm long; ovary glabrous ...................................................................... P. anisophylla Calyx glabrous inside or with sparse trichomes, the calyx lobes 0.6– 1.2 mm long; ovary strigulose on top, the trichomes usually visible only between the septa ................................................................... P. guianensis Inflorescence with the primary axis above the peduncle 1.6–3.2 cm long, the secondary axes 0.6–1.8 cm long and with 1–3 internodes ................................................................................................. P. thyrsiflora Inflorescence with the primary axis above the peduncle 0.5–1.5 cm long, the secondary axes 0–0.4 cm long and with to 1 internode ............... 21 Stipules 1.1–3.6 cm long; leaves obovate, 1.6–2.5 cm broad ....... P. jauaensis Stipules 0.4–1 cm long; leaves elliptic, 0.8–1.5 cm broad ............ P. montana

Pagamea anisophylla Standl. & Steyerm., Fieldiana, Bot. 28: 584, fig. 128. 1953. Shrub or tree 1.8–5(–9) m tall; twigs 2.4– 4.5 mm diameter; leaves 4–14 × 1.6–5.4 cm, margins flat, blade below glabrous or nearly so, with 0–4 barbate tufts of trichomes along each side of midrib; secondary veins 3–7

pairs; petiole 0.75–2.2 cm long; stipules 13– 34 mm long, soon caducous, setae to 1.5 mm long; inflorescence with 15–80(–160) flowers, primary axis 3.6–10.7 cm long, peduncle 1.9– 7.7 cm long, secondary axes 0.3–3 cm long; ovary glabrous; calyx densely pubescent inside, lobes 0–0.6 mm long; fruits spherical,

Pagamea 669

4.4–7 mm diameter. White-sand savannas and scrublands, slopes over sandstone substrate at the base of tepuis, 100–300(–1600) m; Amazonas (Canaripó, Carmelitas, Cerro Camani, Cerro Moriche, Cerro Yutajé, La Esmeralda, basin of Río Atabapo, Río Autana, Río Manapiare, Río Orinoco, Río Samariapo, Río Ventuari, Río Yudi). Endemic. ◆Fig. 526. Pagamea capitata Benth., J. Linn. Soc. Bot. 1: 85, 109. 1857. Pagamea brevipedunculata Steyerm., Mem. New York Bot. Gard. 12(3): 283. 1963. Shrub or tree 1–4 m tall; twigs 1.8–5 mm diameter; leaves 2.6–10 × 0.5–2.9 cm, the margin usually strongly revolute, the lower surface of blade glabrous or nearly so or the midrib sparsely pubescent, trichomes 0.1–1 mm long; secondary veins 4–10 pairs; petiole 0.2–1.3 cm long, densely pubescent; stipules 2.5–6.7 mm long, persistent, setae 0.5–5.5 mm long; inflorescence monocephalous, with 3–26(–50) flowers, primary axis 0.3–5 cm long; ovary glabrous to strigulose at apex; calyx glabrous inside, the lobes 0.4–3 mm long; fruits 3–7 mm diameter. Scrub formations, savannas, and dwarf forests over white sands or sandstone substrates, 50–1900 m. Guyana, Suriname, Brazil; 4 subspecies, 2 in Venezuela, both in the flora area. Pagamea brevipedunculata is treated here as synonym of P. capitata. The differences used by Steyermark (1965) to distinguish these species are largely overlapping (i.e., ovary glabrous in P. brevipedunculata), or are associated with the higher altitude where the type of P. brevipendunculata was collected (i.e., its smaller leaves and inflorescences). Key to the Subspecies of P. capitata 1. Inflorescences with long leaf-like bracts subtending the clusters of flowers, the peduncle (1–)1.3–5 cm long; lower surface of leaves usually with 2–6 barbate tufts of hairs along each side of the midrib, rarely absent ....... subsp. capitata 1. Inflorescences without leaf-like bracts subtending the clusters of flowers, the peduncle 0.5–1.3(–1.9) cm long; lower surface of leaves usually without barbate tufts of hairs, or if present only 1 or 2 per side .................... subsp. conferta

P. capitata subsp. capitata. —Tabirau-yék (Arekuna). Leaves 3.2–9.1 × 0.8–2.9 cm, the lower surface with (0–)2–6 barbate tufts of hairs along midrib; secondary veins 4–8 pairs; inflorescence primary axis (1–)1.5–5 cm long. 50–1400 m; eastern Bolívar (Auyán-tepui, Gran Sabana, Ilú-tepui, Luepa, Santa Elena de Uairén, Uarama-tepui). Guyana, Suriname. ◆Fig. 527. P.

capitata subsp. conferta (Standl.) Steyerm., Mem. New York Bot. Gard. 12(3): 281. 1965. —Pagamea conferta Standl., Publ. Field Colombian Mus., Bot. Ser. 7: 419. 1931. Pagamea capitata subsp. capitata f. breviloba Steyerm., Mem. New York Bot. Gard. 12(3): 281. 1965 Leaves 2.6–10 × 0.5–2.9 cm, the lower surface with 0–2 barbate tufts of hairs along one side of the midrib; inflorescence primary axis 0.3–1.9 cm long. 100–1900 m; western Bolívar (Cerro Guacanoco, Cerro Guaiquinima, Cerro Ichún, Jaua-Sarisariñama massif), Amazonas (Cerro Aracamuni, Cerro Autana, Cerro Duida). Endemic. Steyermark (1965) recognized two forms of subsp. conferta, f. conferta and f. breviloba Steyerm., the latter supposedly differing from the former only in the shape and size of the calyx lobes. These characters vary continuously and independently, and these forms are not recognized here. Pagamea coriacea Spruce ex Benth., J. Linn. Soc. Bot. 1: 110. 1857. —Cupi. Pagamea coriacea var. pubescens Steyerm., Mem. New York Bot. Gard. 12(3): 273. 1965. Shrub or tree 2–13 m tall; twigs 4.8–9 mm diameter; leaves (6.3–)12–18.3 × (1.8–)3.3– 6.6 cm, margin flat, the lower surface blade glabrous or nearly so, rarely with barbate tufts of hairs along the midrib; secondary veins 4–7 pairs; petiole 1.6–5.3 cm long; stipules 8–36 mm long, soon caducous, setae 0.5–4 mm long; inflorescences with 100–600 flowers, primary axis 9–21 cm long, peduncle 4.6–11 cm long, secondary axes (9–) 3.5–8.7 cm long; ovary glabrous; calyx glabrous or nearly so inside, lobes 0–0.6(–1) mm long; fruits spherical, 4–6.5 diameter. Low gallery forests on temporarily flooded soils, scrublands (bana), white-sand savannas, 50–

670

R UBIACEAE

200(–700) m; Amazonas (Río Autana, Río Casiquiare, Río Negro, Río Orinoco, Río Pasimoni, Río Sipapo, Río Ventuari). Amazonian Colombia and Brazil, Guyana. ◆Fig. 534. Steyermark (1965) recognized two varieties in the flora area: var. pubescens Steyerm. and var. coriacea. Variety pubescens was described based on only two, extremely distantly located collections, one from Central Amazon in Brazil, and the other from Venezuela. Additional collections indicate that the differences Steyermark (1965) cited distinguishing var. pubescens from var. coriacea, the presence of barbate hairs in the axils of veins, and the density and size of trichomes of the calyx tube and corolla lobes, vary continuously and independently. The plants from the flora area conform to a single morphological entity, and they match the type collection of P. coriacea (Spruce 2026, BM). Thus here, var. pubescens is not recognized, and the name P. coriacea is used in its strict sense.

1–3(–5) flowers, primary axis 0.7–1.7 cm long, peduncle 0.7–1.6 cm long; ovary glabrous; calyx glabrous inside, the lobes 0.3– 0.7 mm long; fruits spherical, 4.2–6.8 mm diameter. Wet montane forests and dwarf mossy recumbent forests on tepui summit, 1400–1800 m; Amazonas (Cerro Duida). Endemic. ◆Fig. 536.

Pagamea diceras Steyerm., Mem. New York Bot. Gard. 12(3): 281, fig. 30A–D. 1965. Shrub or small tree 2–4 m tall; twigs 1.6– 4.8 mm diameter; leaves 2.5–5.2 × 1–2 cm, margin flat to slightly revolute, no barbate tufts along midrib below, blade glabrous; secondary veins 3–9 pairs; petiole 0.3–0.6 cm long; stipules 5.2–18 mm long, soon caducous, setae 0.7–2.2 mm long; inflorescences with 11–20 flowers, spicately arranged on the sides of the primary axis, this 2.4–5 cm long, peduncle 1.4–3.8 cm long; ovary glabrous; calyx glabrous inside, the lobes (0.1–)0.4–1 mm long; fruits unknown. Along sandstone escarpment on tepui summit, ca. 2000 m; Amazonas (Cerro Parú). Endemic. ◆Fig. 533.

Pagamea guianensis Aubl., Hist. Pl. Guiane 113, pl. 44. 1775. —Ají de paloma. Shrub or tree 2–6(–13) m tall; twigs 2.6– 6.5 mm diameter; leaves 6.5–14.3 × 1.8–5.2 cm, margin flat, with 0–7 barbate tufts of hairs along one side of the midrib below, blade glabrous to sparsely pubescent, trichomes 0.4–0.7 mm long; 4–10 pairs of secondary veins; petiole 1–2.8 cm long; stipules 2–17 mm long, soon caducous, setae 1.6–9 mm long; inflorescences with 14–80(–124) flowers, primary axis 2.5–11.3 cm long, peduncle 1.5–8.3 cm long, secondary axes 0–7 cm long; ovary strigulose at apex, conspicuously so between the septa, rarely glabrous; calyx glabrous or sparsely pubescent inside, the lobes 0.5–1.7 mm long; fruits ellipsoid, 4.4–8.5 × 4–7.2 mm. Riparian forests and scrublands (bana) on temporarily flooded white sands, 25–300 m; Amazonas (basins of Río Guainía, Río Manapiare, Río Negro, Río Orinoco, Río Pasimoni, and Río Ventuari). Apure; Guyana, Suriname, French Guiana, Brazil. ◆Fig. 529. The name Pagamea guianensis has been used in a very broad sense to designate many species throughout the range of the genus, and sometimes has been misapplied to lineages that are not immediately related. The Guayana plants match the type specimen, so this name is considered in its strict sense here.

Pagamea duidana Standl. & Steyerm., Fieldiana, Bot. 28: 586. 1953; emend. Steyerm., Mem. New York Bot. Gard. 12(3): 282. 1965. Shrub or small tree 1.5–1.8 m tall; twigs 0.8–2.8 mm diameter; leaves 1.4–4.4 × (0.45–) 1–1.8 cm, margin flat to slightly revolute, with 1–3 barbate tufts of hairs along on side of midrib below, blade glabrous or nearly so; secondary veins 2–5 pairs; petiole 0.3–1.1 cm long; stipules 2–7 mm long, soon caducous, setae 1.1–3.1 mm long; inflorescences with

Pagamea hirsuta Spruce ex Benth., J. Linn. Soc. Bot. 1: 111. 1857. —Vieja. Shrub or tree 1.5–8 m tall; twigs 4.5–7 mm diameter; leaves (8.8–)12.5–24 × (2.6–) 3.6–6.8 cm, margin flat, blade, midrib, and veins hirsute below, trichomes 2–4.2 mm long; secondary veins 6–11 pairs; petiole (0.8–)1.3–2.5 cm long; stipules (10–)21–40 mm long, soon caducous, setae 0.5–2 mm long; inflorescences with 20–60 flowers, primary axis 5.2–13 cm long, peduncle 3.8–10 cm long, secondary axes 0.15–0.6 cm long;

Pagamea 671

ovary glabrous; calyx densely pubescent inside, the lobes 0–0.2 mm long; fruits ellipsoid, 5.5–10 × 3.3–5.5 mm. Dwarf forests and dense scrub formations (bana) over white sands, 50–200 m; southwestern Amazonas (basins of Río Atabapo, Río Guainía and Río Negro). Colombia. Pagamea jauaensis Steyerm., Mem. New York Bot. Gard. 23: 883, fig. 15. 1972. Shrub to small tree 2.5–4 m tall; twigs 2.1–6 mm diameter; leaves 3.7–8.6 × 1.2–2.6 cm, margin flat, without barbate tufts of hairs along the midrib below, blade glabrous to sparsely strigose below, trichomes to 0.4 mm long; secondary veins 4–7 pairs; petiole 0.4–1.4 cm long; stipules 9.5–35 mm long, soon caducous, setae 0.3–1.7 mm long; inflorescences with 3–18 flowers, primary axis 2.1–5.3 cm long, peduncle 1.7–4.5 cm long, secondary axes 0–0.4 cm long; ovary glabrous; calyx sparsely pubescent inside, the lobes 0.2–0.6 mm long; fruits ellipsoid, 5.5–7 × 4.5–6.2 mm. Shrub thickets on tepui summits, 1600–2000 m; Bolívar (Cerro Jaua), Amazonas (Cerro Yutajé, Sierra de Maigualida). Endemic. ◆Fig. 525. Pagamea magniflora Steyerm., Mem. New York Bot. Gard. 12(3): 285. 1965. Shrub to 2.5 m tall; twigs 2.2–3.2 mm diameter; leaves 2.9–4 × 0.8–1.5 cm, margin strongly revolute, without barbate tufts of hairs along midrib below, blade glabrous; secondary veins 4 or 5 pairs; petiole 0.6–0.9 cm long; stipules 4.6–11 mm long, soon caducous, setae 0–1 mm long; inflorescences with 1–3 flowers, primary axis 0.1–0.2 cm long; ovary glabrous; calyx glabrous inside, the lobes 0.5–1.5 mm long; fruits unknown. Dense scrub formation on tepui summits, ca. 2500 m; Bolívar (Ilú-tepui). Endemic. ◆Fig. 530. Pagamea montana Gleason & Standl., Publ. Field Columbian Mus., Bot. Ser. 7: 421. 1931. Shrub or tree 2–8 m tall; twigs 1.1–3.4 mm diameter; leaves (1.8–)3.2–6.8 × 0.6–1.8 cm, margin flat to slightly revolute, without (rarely with up to 3) barbate tufts of hairs along one side of the midrib below, blade glabrous or nearly so; secondary veins 0–7 visible pairs; petiole 0.3–1 cm long; stipules 2.3– 11 mm long, soon caducous, setae 0.2–2.1

mm long; inflorescences with (1–)3–70 flowers, primary axis 0.8–4.2 cm long, peduncle 0.5–3.1 cm long, secondary axes 0–0.6 cm long; ovary glabrous; calyx glabrous or nearly so inside, the lobes 0.2–1.5 mm long; fruits ellipsoid, 5–6.5 × 3.4–6 mm. Dwarf forests, gallery forests, and scrublands, on tepui slopes and summits, 600–1900 m; western Bolívar (Cerro Marutaní), Amazonas (Cerro Aracamuni, Cerro Duida, Cerro Guanay, Cerro Marahuaka, Cerro Sipapo, Serranía Parú, Cerro Yutajé, Sierra de la Neblina). Brazil (Amazonas: Serra Aracá) and Colombia (Vaupés: Mount Chiribiquete). ◆Fig. 528. Pagamea pauciflora Standl. & Steyerm., Fieldiana, Bot. 28: 588. 1959; emend. Steyerm., Mem. New York Bot. Gard. 12(3): 283. 1965. Shrub 1–5 m tall; twigs 1.3–4 mm diameter; leaves 1.6–5.1 × 0.4–2.1 cm, margin strongly revolute, without barbate tufts of hairs along midrib below, blade glabrous or nearly so; secondary veins 3–5 pairs, sometimes not visible; petiole 0.2–1.1 cm long; stipules (1–)3–9 mm long, soon caducous, setae 0.7–4.5 mm long; inflorescence with 1–6 flowers, primary axis 0.6–3.1 cm long, peduncle 0.4–2.7 cm long; ovary glabrous; calyx glabrous inside, the lobes 0.2–2.2 mm long; fruits ellipsoid, 6.5–10 × 3.8–6.4 mm. Wet dwarf montane forests, dwarf forested thickets below sandstone bluffs, among rocky crevasses, and dwarf forests along streams on upper slopes and summits of tepuis, 1400– 2500 m; eastern Bolívar (Aparamán-tepui, Carrao-tepui, Macizo del Chimantá, Murisipán-tepui, Ptari-tepui, Tereké-Yurén-tepui). Endemic. ◆Fig. 531. Pagamea plicata Spruce ex Benth., J. Linn. Soc. Bot. 1: 109. 1857. —Ají de paloma, Cachira-huaro (Baniva), Garrapata. Pagamea plicata var. multinervia Steyerm., Mem. New York Bot. Gard. 12(3): 274. 1965. Shrub to tree 3–20 m tall; twigs 4.8–8.5 mm diameter; leaves (7.2–)10–23.5 × (2.7–) 3.7–8.3 cm, margin flat, without barbate tufts of hairs along midrib below, blade sparsely pubescent below, trichomes 0.7–1.1 mm long; secondary veins 5–11 pairs; petiole (1–)1.6–4.7 cm long; stipules 3.5–51 mm long, soon caducous, setae 0.8–2 mm long; in-

672

R UBIACEAE

florescences with 40–250 flowers, primary axis 4.8–13.5 cm long, peduncle 2.5–8.4 cm long, secondary axes 0–1.5 cm long; ovary usually strigulose at apex, rarely glabrous; calyx glabrous or nearly so inside, the lobes 0.5–1.4(–2) mm long; fruits ellipsoid, 4.8–8.2 × 4.8–8.5 mm. Scrub formation and savannas on white sands, forest borders, periodically flooded forests, talus slopes of tepuis, 50–700 m; Bolívar (near Urimán), Amazonas (Río Autana, Río Casiquiare, Río Guainía, Río Negro, Río Orinoco, Río Pasimoni, Río Sipapo). Colombia, Brazil. ◆Fig. 539. Steyermark (1965; 1974) recognized two varieties in the flora area, var. plicata and var. multinervia Steyerm., the latter differing by having leaves with > 10 lateral veins, and the veins below with trichomes denser and longer (0.8–1 mm) than var. plicata. These characters vary independently, and leaves of the NY isotype of var. plicata (Spruce 2342) also have 10 veins. These varieties are not recognized here. The dried leaves of Pagamea plicata are twisted and used in wrapping tobacco. Pagamea plicatiformis Steyerm., Mem. New York Bot. Gard. 12(3): 274. 1965. Tree to 5 m tall; twigs 2.7–4.2 mm diameter; leaves (4.6–)8–16 × (1.6–)2.5–4.7 cm, margin flat, barbate tufts absent, blade sparsely pubescent below, trichomes 1.2–1.6 mm long; secondary veins 5–10 pairs; petiole 0.6–2.4 cm long; stipules (4–)14–24 mm long, soon caducous, setae 2–5.6 mm long; inflorescences with 20–120 flowers, primary axis 1.85–6.4 cm long, peduncle 0.9–4.6 cm long, secondary axes 0–0.4 cm long; ovary strigulose at apex; calyx glabrous inside, the lobes 0.7–2.1 mm long; fruits ellipsoid, 6–9 × 4–7 mm. Riparian forests, white-sand scrublands, dwarf forests, 50–200 m; Amazonas (Maroa, Río Guainía, Río Mucuriapi, Santa Barbara del Orinoco). Brazil (Amazonas: basin of Rio Negro). Pagamea sessiliflora Spruce ex Benth., J. Linn. Soc. Bot. 1: 110. 1857. Shrub or tree 1–5 m tall; twigs 1.2–1.9 mm diameter; leaves 2–6.7 × 0.7–2.2 cm, margin flat, the lower surface of blade glabrous, rarely pubescent, with (0)1–4 barbate tufts of hairs along one side of midrib, trichomes to 0.4 mm long; secondary veins 4–7

pairs; petiole 0.2–0.6 cm long; stipules 2.2–4 mm long, soon caducous, setae 0.5–1.4 mm long; inflorescences with 1–10 flowers, sessile or nearly so, clustered together forming a many-flowered pseudoterminal head; ovary glabrous or strigulose at apex; calyx glabrous inside, the lobes 1.5–3.2 mm long; fruits ellipsoid, 7.5–10 × 5–8 mm. Evergreen lowland and riparian forests, exposed igneous substrates and sandy soil, 50–200 m; southwestern Amazonas (basins of Río Baría, Río Negro, Río Pasimoni, and Río Yatúa). Endemic. ◆Fig. 538. Pagamea standleyana Steyerm., Fieldiana, Bot. 28: 589, fig. 130. 1953; emend. Steyerm., Mem. New York Bot. Gard. 12(3): 282. 1965. Pagamea reducta Steyerm., Mem. New York Bot. Gard. 12(3): 283. 1965. Pagamea uniflora Steyerm., Mem. New York Bot. Gard. 12(3): 284. 1965. Shrub or tree 2–12 m tall; twigs 1–3 mm diameter; leaves 1.2–5.3 × 0.4–1.6 cm, margin flat to slightly revolute, the lower surface of blade glabrous or nearly so, with 1–4 barbate tufts of hairs along each side of midrib; secondary veins 0–6 visible pairs; petiole 0.2–0.8 cm long; stipules 1.5–9 mm long, soon caducous, setae 0.4–2.1 mm long; inflorescences with 1–12(–19) flowers, primary axis 0.3–3 cm long, peduncle 0.2–3 cm long, secondary axes absent or rarely to 0.3 cm long; ovary glabrous; calyx glabrous inside, the lobes 0.4–2.5 mm long; fruits ellipsoid, 4–7.5 × 3–5.3 mm. Tepui slope and summit scrub forests, 1200–2200 m; Bolívar (Auyán-tepuí, Camarcaibarai-tepuí, Cerro Guaiquinima, Cerro Uaipán, Cerro Venamo, Macizo del Chimantá, Ptari-tepui, Sarvén-tepuí, Ueitepuí). Endemic. ◆Fig. 535. Pagamea thyrsiflora Spruce ex Benth., J. Linn. Soc. Bot. 1: 110. 1857. —Sakauyék (Arekuna). Shrub or tree 1.5–5(–10) m tall; twigs 1.5– 2.8 mm diameter; leaves 3.5–10.5 × 0.8–2.4 cm, margin flat to slightly revolute, the lower surface of blade glabrous, rarely with sparse trichomes to 0.4 mm, with 1–5 barbate tufts of hairs along each side of midrib; secondary veins 4–10 pairs; petiole 0.3–1.2 cm long; stipules 3.9–10 mm long, soon caducous, setae (0.8–)1.3–3.7 mm long; inflorescences

Pagamea 673

with 11–90(–120) flowers, primary axis 2.2– 5.6 cm long, peduncle 0.4–2.8 cm long, secondary axes (0.2–)0.6–2 cm long; ovary glabrous; calyx glabrous inside, the lobes 0.1– 0.5 mm long; fruits ellipsoid 3.8–6 × 2.6–5 mm. Scrub formations, shrublands, dwarf forests at bases of and slopes of tepuis, 100– 1400 m; Bolívar (Cerro Arepuchi in upper Río Caroní basin opposite mouth of Río Icabarú, El Paují, Río Caroní, Río Paragua, Salto Uraima, Sierra de Lema); Amazonas (Cerro Duida, Cerro Huachamacari, Cerro Moriche, Cerro Parú, Cerro Vinilla, Río Casiquiare, Río Guainía, Río Guayapo, Río Negro, Río Orinoco, Río Sipapo, San Fernando de Atabapo, Sierra de la Neblina, Yavita). Guyana, Amazonian Colombia, Brazil, and Bolivia. ◆Fig. 532.

Pagamea velutina Steyerm., Mem. New York Bot. Gard. 12(3): 275. 1965. Tree to 12 m tall; twigs 3–6 mm diameter; leaves (8.4–)13–19 × (3–)4–6.4 cm, margin flat, without barbate tufts of hairs below, blade densely pubescent below, trichomes 0.8–1 mm long; secondary veins 4–7 pairs; petiole 1.4–2.3 cm long; stipules 25–69 mm long, soon caducous, setae 0.3–2 mm long; inflorescences with 8–50 flowers, primary axis 4.6–9 cm long, peduncle 2.4–4.6 cm long, secondary axis 0.3–1.4 cm long; ovary strigulose at apex; calyx densely pubescent inside, the lobes 0.2–1.4 mm long; fruits globose to ellipsoid, 6.6–8.5 × 5.7–8.3 mm. White-sand savannas, tepui slope forests, 100–1500 m; Amazonas (Cerro Sipapo). Endemic. ◆Fig. 537.

Fig. 525. Pagamea jauaensis

674

R UBIACEAE

Fig. 526. Pagamea anisophylla

Fig. 527. Pagamea capitata subsp. capitata

Pagamea 675

Fig. 528. Pagamea montana

Fig. 530. Pagamea magniflora

Fig. 529. Pagamea guianensis

676

R UBIACEAE

Fig. 531. Pagamea pauciflora

Fig. 533. Pagamea diceras

Fig. 532. Pagamea thyrsiflora

Pagamea 677

Fig. 534. Pagamea coriacea

Fig. 535. Pagamea standleyana

Fig. 536. Pagamea duidana

678

R UBIACEAE

Fig. 537. Pagamea velutina

Fig. 538. Pagamea sessiliflora

Fig. 539. Pagamea plicata

62. PAGAMEOPSIS Steyerm., Mem. New York Bot. Gard. 12(3): 267. 1965. by Julian A. Steyermark and Charlotte M. Taylor Shrubs, sometimes pachycaulous, terrestrial, unarmed. Leaves opposite, commonly congested at stem apices, subsessile, venation not visible; stipules interpetiolar and fused to petioles, truncate, persistent, shortly ciliate or shortly setose, generally valvate to imbricate in bud. Inflorescences terminal and pseudoaxillary, multiflowered, pedunculate, subcapitate to paniculate, bracteate, the bracts often fused. Flowers small, sessile, floral biology unknown. Hypanthium turbinate, those of adjacent flowers sometimes partially to completely fused. Calyx limb deeply

Pagameopsis 679

lobed, lobes 4–6, without calycophylls; corolla shortly tubular, blue to purple, internally hirtellous in middle to upper part of tube, tube fenestrate, lobes 5 or 6, valvate in bud. Stamens 5, inserted near middle of corolla tube; anthers narrowly oblong, dorsifixed near middle, partially exserted. Ovary 2-locular; ovules 1 in each locule, basal. Fruit drupaceous, obovoid to subglobose, indehiscent, fleshy, sometimes fused to the other fruits; pyrene 1, 2-celled, bony. Seeds obovoid, smooth, small. Venezuela, Brazil; 2 species, both in the flora area. Pagameopsis is similar to and apparently sometimes sympatric with Aphanocarpus and Coryphothamnus; its separation from these are discussed under each of these other genera. Piesschaert et al. (2001) detailed many aspects of the morphology of this poorly known genus. The pubescent disk and fenestrate (i.e., longitudinally slitted) corolla of Pagameopsis were first reported by them. These conditions are unusual in the Rubiaceae. As noted by these authors, Pagameopsis is clearly not closely related to Pagamea and also does not belong to the tribe Psychotrieae. Key to the Species of Pagameopsis 1.

1.

Leaves with marginal cilia divaricate to subascending, 0.5–1 mm long; inflorescence with flowering portion 1.5–4 × 2–4 cm, comprising 1– 5 capitula or dense glomerules .............................................. P. garryoides Leaves with marginal cilia appressed to ascending, < 0.5 mm long; inflorescence with flowering portion 5–11 × 3–6 cm, comprising numerous small glomerules ........................................................................ P. maguirei

Pagameopsis garryoides (Standl.) Steyerm., Mem. New York Bot. Gard. 12(3): 267. 1965. —Pagamea garryoides Standl., Publ. Field Columbian Mus., Bot. Ser. 7: 420. 1931. Shrub to 1 m tall; leaves 2–4 × 1–1.8 cm; stipule sheath 2–7 mm long, cilia or setae 0.5–1 mm long; peduncles 1.5–6.5 cm long; calyx limb 1.5–3.5 mm long; corolla tube 1.5– 2 mm long, lobes 2–3 mm long; fruit not seen. Tepui meadows, scrub savannas with bush islands, borders of savannas, 1100–2500 m; Bolívar (Cerro Guanacoco, Cerro Sarisariñama, Gran Sabana, Ilú-tepui, TerekéYurén-tepui), Amazonas (Cerro Duida, Cerro Huachamacari, Cerro Marahuaka, Cerro Parú, Cerro Yutajé, Sierra de la Neblina). Endemic. ◆Fig. 540. Pagameopsis maguirei Steyerm., Mem. New York Bot. Gard. 12(3): 267, fig. 37. 1965. Pagameopsis maguirei subsp. neblinensis Steyerm., Mem. New York Bot. Gard. 12(3): 270, fig. 37. 1965. Pagameopsis maguirei subsp. neblinensis var. angustifolius Steyerm., Mem. New York Bot. Gard. 12(3): 270. 1965. Pagameopsis maguirei subsp. pusillus Steyerm., Ann. Missouri Bot. Gard. 74:

106, fig. 8. 1987. Pagameopsis maguirei subsp. pusillus var. glabrus Steyerm., Ann. Missouri Bot. Gard. 74: 108. 1987. Shrub to 3 m tall; leaves 2.5–10 × 0.5–2.5 cm; stipule sheath 5–6 mm long, cilia or setae 0.5–1 mm long; peduncles 2–10 mm long; calyx limb 2.5–4.5 mm long; corolla tube 2–2.5 mm long, lobes 2–3 mm long; fruit 1.8–2 × 2 mm. Dwarf open forests, Bonnetia forests, thickets on rocky escarpments, rocky borders of streams, open rock outcrops, shrubby borders of savanna formations, summits of tepuis, 1500–2300 m; Bolívar (Auyán-tepui, Macizo del Chimantá), Amazonas (Sierra de la Neblina summit of Cerro Avispa). Brazil. ◆Fig. 541. The extensive infraspecific classification presented by Steyermark (1984) recognized three subspecies and six varieties, distinguished by characters that are continuously variable and probably strongly environmentally influenced rather than taxonomically informative: degree of development of the internodes, stipule sheath length, leaf size, length and distribution of trichomes, and degree of expansion of the inflorescences. Consequently, Steyermark’s infraspecific taxa are not recognized here.

680

R UBIACEAE

Fig. 540. Pagameopsis garryoides

Fig. 541. Pagameopsis maguirei

63. PALICOUREA Aubl., Hist. Pl. Guiane 172, pl. 66. 1775. by Charlotte M. Taylor and Julian A. Steyermark Shrubs or trees, unarmed, terrestrial, sometimes subshrubs from a xylopodium, sometimes with hollow stems housing ants. Leaves opposite or 3- or 4-verticillate, petiolate or sessile, venation not lineolate; stipules persistent, united around the stem or sometimes laminar (i.e., subinterpetiolar due to the reduced intrapetiolar portion), in bud generally valvate to imbricate, in the interpetiolar portion bilobed with the lobes usually well developed. Inflorescence terminal, multiflowered, congested-cymose to paniculate, pedunculate, bracteate, usually brightly colored. Flowers sessile or at least some pedicellate, distylous, medium-

Palicourea 681

sized and often showy. Hypanthium turbinate to subglobose. Calyx limb reduced to well developed, lobes (4)5, without calycophylls; corolla tubular to funnelform, white, yellow, orange, red, blue, or purple, internally glabrous except with a dense ring of trichomes in the lower part of the tube, the tube swollen at the base, often asymmetrically (i.e., gibbous), often curved, lobes (4)5, valvate in bud. Stamens (4)5, inserted near or above middle of corolla tube; anthers narrowly oblong, dorsifixed near middle, partially exserted to exserted in short-styled flowers, included in longstyled flowers. Ovary 2(–5)-locular; ovules solitary in each locule, basal. Fruit subglobose to ovoid, drupaceous, carnose or spongy, blue or purple-black; pyrenes 2(–5), 1-locular, bony, hemispherical to triangular. Seeds ellipsoid, rather small. Mexico, Central America, West Indies, Colombia, Venezuela, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina; ca. 250 species, about 50 in Venezuela, 27 of these in the flora area. Palicourea is sometimes confused with Hamelia and Isertia. Its separation is discussed under each of the other genera, respectively. Palicourea is closely related to Psychotria subgenus Heteropsychotria, and these will probably eventually be combined. They are separated only by their corollas, which in Palicourea are colorful with well developed tubes that are swollen at the base, with this swelling closed off by a dense ring of trichomes; versus the corollas white with straight bases and no ring of trichomes in the bottom half in Psychotria. The unusual corolla form of Palicourea appears to be an adaptation for hummingbird pollination: Palicourea flowers produce a comparatively large amount of nectar, which accumulates in the swollen corolla base where it is protected from most insects by the relatively long tube and the trichomes. This nectar is accessible to hummingbirds, which are strong enough to push their beaks through the trichome ring. In contrast, the flowers of subgenus Heteropsychotria are generally insect-pollinated. Without flowers these genera are virtually impossible to separate. Taylor (1997) provided an overview of Palicourea. The genus comprises two subgenera, both of which are found in the flora area. In the flora area, subgenus Montanae C.M. Taylor is represented by P. perquadrangularis, while the remaining species belong to subgenus Palicourea. The color patterns of the inflorescence axes and corollas are usually informative in Palicourea. However, Taylor (1997) considered details of the pubescence to be much less informative than Steyermark did. The infructescence axes of most species are purple, even when the inflorescence is a different color; these structures attract different birds because of the inflorescences, and change their color as the fruits develop. The flowers are 5merous, but occasional variant flowers on a plant may be 4-merous and/or 6merous. Key to the Species of Palicourea (includes Psychotria leiantha) 1. 1. 2(1). 2. 3(2).

Leaves sessile or very shortly petiolate, cordulate at the base ............... 2 Leaves subsessile to petiolate, truncate to rounded, obtuse, or acute at the base .................................................................................................. 4 Leaves bullate; stems and lower surface of leaves hirtellous .......... P. huberi Leaves plane; stems and lower surface of leaves glabrous to puberulous ................................................................................................................ 3 Leaves lanceolate to ovate, 2.5–7 cm wide, with the secondary veins 9– 12 pairs, with the margins flat; peduncles 2–2.5 cm long ........ P. foldatsii

682

R UBIACEAE

3. 4(1). 4. 5(4). 5. 6(5).

6.

7(6). 7. 8(4). 8. 9(8). 9. 10(9).

10.

11(10).

11.

12(11).

12. 13(12). 13. 14(11).

Leaves lanceolate, 1.2–2.5 cm wide, with the secondary veins 5–9 pairs, with the margins revolute; peduncles 0.8–4 cm long .............. P. lancigera Leaves 3 or 4 per node ............................................................................... 5 Leaves 2 per node ...................................................................................... 8 Inflorescences pyramidal to narrowly so, 2 or more times as long as wide; stipule lobes 10–16 mm long ..................................................... P. triphylla Inflorescences corymbiform, wider than long or about as wide as long; stipule lobes 0.2–3 mm long .................................................................. 6 Floral bracts shorter than or equal to calyx limb (i.e., not exceeding hypanthium plus calyx limb), subtending but not surrounding the flower; corollas with tuberculate projections at the sinuses between the lobes; pyrenes 2–5 per fruit ............................................................ P. corymbifera Floral bracts equal to or longer than the calyx limb, at least partly surrounding the base of the flower; corolla with the sinuses between the lobes smooth; pyrenes 2 per fruit .......................................................... 7 Peduncles 1.8–2 cm long; corolla tube ca. 6 m long ..................... P. cardonae Peduncles 4–11.5 cm long; corolla tube 8–12 mm long ............ P. quadrifolia Stems, lower surface of leaves, peduncles, and inflorescence axes densely velutinous or pilosulous ........................................................ P. longistipula Stems, lower surface of leaves, peduncles, and inflorescence axes glabrous or sparsely to moderately puberulous, strigillose, or hirtellous .......... 9 Older stems with thick corky bark, plants often from a xylopodium; leaves subsessile to shortly petiolate ............................................ P. rigida Older stems with thin smooth bark, plants without xylopodium; leaves subsessile to long-petiolate ................................................................. 10 Stipules united around the stem into a sheath that is longer in the intrapetiolar portion than in the center of the emarginate interpetiolar portion ................................................................................. P. wurdackiana Stipules longer in the interpetiolar portion than the intrapetiolar portion, united around the stem into a continuous sheath or laminar (i.e., with the intrapetiolar portion reduced and the interpetiolar portion developed) ............................................................................................ 11 Stipules laminar and lobed for as much as 2/3 of their length, with the sinus between the lobes acute to rounded; plants of higher elevations, 1800–2800 m ........................................................................................ 12 Stipules united around the stem into a continuous sheath, this sheath sometimes somewhat reduced in the intrapetiolar portion, truncate to broadly rounded in the interpetiolar portion, with lobes as long as or longer than the sheath ........................................................................ 14 Leaves rounded to usually truncate at the base, with 10–15 pairs of secondary veins; petioles 6–45 mm long; calyx limb 0.8–2.2 mm long .................................................................................................... P. obtusata Leaves acute to rounded at the base, with 4–8 pairs of secondary veins; petioles 2–5 mm long; calyx limb 0.4–1 mm long............................... 13 Inflorescence axes red; corolla yellow, with the tube 5–5.5 mm long .................................................................................................. P. ottohuberi Inflorescence axes purple; corolla purple, with the tube ca. 14.5 mm long ...................................................................................................... P. pensilis Longest stipule lobes 5–16 mm long, narrowly triangular to lanceolate,

Palicourea 683

14.

15(14). 15. 16(15). 16. 17(14).

17. 18(17). 18. 19(17). 19. 20(19). 20. 21(20). 21. 22(19). 22.

23(22). 23. 24(22). 24. 25(24). 25. 26(24).

sharply acute, and the inflorescence narrowly pyramidal, 8–28 cm long and half or less as wide at the widest point as long ........................... 15 Longest stipule lobes 1–9 mm long, linear to ligulate, sharply acute to obtuse or rounded, and the inflorescence 1.5–18 cm long, roundedcorymbiform to pyramidal, at the widest point half or more as wide as long ....................................................................................................... 17 Stipule sheath 3–6 mm long; fruit subglobose, 3–4 mm diameter; plants of montane forests, 1200–2000 m ............................ P. perquadrangularis Stipule sheath 1–2 mm long; fruit laterally somewhat flattened, 3.5–7 × 3.5–7 mm; plants of lowland forests, 50–1000 m ............................... 16 Fruit 6–7 × 6–7 mm .................................................................... P. calophylla Fruit 3.5–5 × 3.5–4 mm ................................................................. P. triphylla Floral bracts 3–13 mm long, longer than the calyx limb (i.e., the total length of the hypanthium plus calyx), extending at least to the middle of corolla tube ....................................................................................... 18 Floral bracts reduced or to 1.5 mm long, shorter than the calyx limb ...... 19 Floral bracts 3–8 mm long, yellow to orange; peduncle 1–11 cm long; widespread at 650–800 m ......................................................... P. bracteosa Floral bracts 8–13 mm long, green; peduncle 0.3–1 cm long; 1900–2100 m on Sierra de la Neblina ................................................ Psychotria leiantha Stipule lobes ligulate to broadly triangular, obtuse to rounded, relatively wide at the middle, 1–4 mm wide at widest part ............................... 20 Stipule lobes narrowly triangular to linear, acute, narrow, widest at the base, 0.5–1 mm wide ........................................................................... 22 Stipule lobes 0.8–3 mm long; inflorescence broadly pyramidal to rounded corymbiform; fruit 7–8 × 7–8 mm ........................................ P. grandiflora Stipule lobes 3–13 mm long; inflorescences broadly to narrowly pyramidal; fruit 4–7 × 3.5–5 mm .................................................................... 21 Leaves with 17–21 pairs of secondary veins; stems terete or subterete; ovary locules 4–6 ................................................................... P. grandifolia Leaves with 10–17 pairs of secondary veins; stems subquadrate; ovary locules 2 ................................................................................... P. guianensis Corollas externally densely lanose-pubescent with triangular to linear trichomes to 0.5 mm long .................................................................... 23 Corollas externally glabrous to sparsely or moderately puberulous or hirtellous with linear trichomes to 0.3 mm long, or densely granulosepuberulous with trichomes < 0.1 mm long ......................................... 24 Corolla externally puberulous on tube but glabrous on lobes, the tube 11– 14 mm long; leaves with secondary veins 8–17 pairs .............. P. nitidella Corolla with the external pubescence similar on the tube and lobes, the tube 17–18 mm long; leaves with secondary veins 5–9 pairs ........ P. sp. A Corollas yellow at base usually grading to red, pink, or purple at apex, tubes 14–28 mm long; fruit 4–5 × 5–6 mm ......................................... 25 Corollas red, orange, or yellow, similarly colored throughout or becoming orange or red at apex, tubes 8–13 mm long; fruit 3–6 × 3.5–6 mm ...... 26 Corollas glabrous externally, with tubes ca. 16 mm long ......... P. glabriflora Corollas granulose-puberulous externally, with tubes 14–28 mm long .................................................................................................. P. longiflora Inflorescences corymbiform, with the secondary axes as well developed as

684

R UBIACEAE

the primary axis and strongly ascending ................................ P. fastigiata Inflorescences cylindrical, narrowly to broadly pyramidal, or subcorymbiform, with the secondary axes usually less well developed than the primary axis, ascending to spreading ...................................................... 27 27(26). Corollas deep orange to red; fruit 3–4.5 × 3.5–4.5 mm; inflorescences pyramidal ....................................................................................... P. crocea 27. Corollas yellow to orange; fruit 4–6 × 4.5–6 mm; inflorescences cylindrical, narrowly to broadly pyramidal, to subcorymbiform .................... 28 28(27). Leaves 4.5–18 × 1.5–7 cm; inflorescences cylindrical, narrowly to broadly pyramidal, or subcorymbiform; plants found from 35 to 1700 m but most common below 400 m ...................................................... P. croceoides 28. Leaves 13–19 × 4.5–10 cm; inflorescences pyramidal; plants found from 400 to 1500 m ............................................................................ P. tepuicola 26.

Palicourea bracteosa Standl., Publ. Field Columbian Mus., Bot. Ser. 8: 222. 1930. Psychotria edaphothrix Steyerm., Ann. Missouri Bot. Gard. 75: 342, fig. 10. 1988. Shrub to 8 m tall, puberulous or hirtellous to glabrescent; leaves 2 per node, 16–32 × 6– 15 cm; petioles 5–18 mm long; stipules united around stem, sheath 2–4 mm long, lobes 2–4 mm long, acicular; inflorescence yellow to orange, peduncle 1–11 cm long, branched portion corymbiform, 3–5.5 × 3.5– 11 cm; floral bracts 3–8 mm long; calyx limb 0.1–0.5 mm long; corolla white to yellow, tube 10–14 mm long, lobes 1.5–3 mm long; fruits 3–4 × 4–5 mm; pyrenes 2. Upland forests, 600–800 m; Bolívar (near El Paují). Peru, Amazonian Brazil. Although Steyermark classified this species in Psychotria based on its white flowers, it has an older name in Palicourea and is similar morphologically to Palicourea quadrifolia (Rudge) DC., also found in the flora area. Palicourea calophylla DC., Prodr. 4: 529. 1830. Shrub or tree to 5 m tall, glabrescent; leaves 2 per node, 12–40 × 3.5–11 cm; petioles 5–30 mm long; stipules united around stem, sheath 1–2 mm long, lobes 5–10 mm long; inflorescences orange to pink, peduncle 4.5–8 cm long, branched portion narrowly pyramidal, 8–12 × 3–6 cm; floral bracts 0.5–1 mm long; calyx limb 0.3–0.8 mm long; corolla pink to yellow or orange, tube ca. 9 mm long, lobes 1–2.5 mm long; fruit 6–7 × 6–7 mm; pyrenes 2. Riparian forests, 100–200 m; Bolívar (Haymary), Amazonas (upper Río Orinoco). Amazonian Colombia, Guyana, Suriname, French Guiana, Amazonian Brazil.

The name “Palicourea nicotianaefolia” was incorrectly applied to this species by some previous authors; however, P. nicotianifolia Cham. & Schltdl. is actually a synonym of P. macrobotrys (Ruiz & Pav.) Roem. & Schult., which is found outside the flora area from the south-central Amazon basin to southeastern Brazil and Paraguay. Palicourea cardonae Steyerm., Mem. New York Bot. Gard. 23: 759. 1979. Shrub or small tree, height not noted, glabrous to strigillose or tomentulose; leaves 3 or 4 per node, 18–22 × 3.5–5 cm; petioles 8– 10 mm long; stipules united around the stem, sheath 2–3 mm long, lobes 0.2–0.3 mm long, acicular; inflorescence color not noted, peduncle 1.8–2 cm long, branched portion corymbiform, ca. 1.3 × 2 cm; floral bracts 1– 1.5 mm long; corolla color not noted, tube ca. 6 mm long, lobes ca. 3 mm long; fruit unknown. Riparian forests, 100–200 m; Amazonas (Río Castanho, affluent of Rio Padauiri on Brazil-Venezuela border). Palicourea cardonae is known only from the type. Palicourea corymbifera (Müll. Arg.) Standl., Publ. Field Columbian Mus., Bot. Ser. 7: 127. 1930. —Psychotria corymbifera Müll. Arg. in Mart., Fl. Bras. 6(6): 247. 1881. —Carrizo, Carrizo de picure, Sorteja amarilla. Psychotria verrucosa Müll. Arg. in Mart., Fl. Bras. 6(6): 246. 1881. Palicourea expetens Standl., Lloydia 2: 216. 1939. Shrub or tree to 8 m tall, puberulous to glabrescent; leaves 4(–6) per node, 12–36 × 3.5–12 cm; petioles 4–30 mm long; stipules

Palicourea 685

united around stem, sheath 2–3 mm long, lobes 0.5–1 mm long, narrowly triangular to linear; inflorescences yellow to orange or red, peduncles 3–22 cm long, branched portion corymbiform, 3–12 × 4–16 cm; floral bracts 0.2–0.8 mm long; calyx limb ca. 0.3 mm long; corolla yellow to orange or red, tube 8–9 mm long, lobes 1.5–2 mm long; fruit 3–4 × 4.5 mm; pyrenes 2–5. Evergreen lowland to montane and riparian forests, 50–700(–1200) m; Bolívar (Gran Sabana, Sierra Pakaraima), Amazonas (Río Cunucunuma, Río Negro, basin of Río Orinoco, Río Ventuari, Sierra Parima). Amazonian Colombia, Guyana, Suriname, French Guiana, Peru, Brazil. ◆Fig. 543. Palicourea crocea (Sw.) Roem. & Schult., Syst. Veg. 5: 193. 1819. —Psychotria crocea Sw., Prodr. Veg. Ind. Occ. 44. 1788. Shrub or tree to 5 m tall, hirtellous or puberulous to glabrescent; leaves 2 per node, 6–18.5 × 2–7 cm; petioles 1–10 mm long; stipules united around stem, sheath 0.2–1 mm long, lobes narrowly triangular, 1–3 mm long; inflorescences red, peduncles 1.5–15 cm long, branched portion pyramidal, 2.5–12 × 1.5–8 cm; floral bracts 0.5–2 mm long; calyx limb 0.5–1 mm long; corolla red to deep orange, tube 8–12 mm long, lobes 1–2.5 mm long; fruit 3–4.5 × 3.5–4.5 mm; pyrenes 2. Evergreen lowland to upland and riparian forests, 50–1000 m; Delta Amacuro (Caño Orocoima, northeast of El Palmar), Bolívar (Río Caura, Río Nichare, Río Paramici), Amazonas (middle Río Casiquiare, near San Carlos de Río Negro, base of Sierra de la Neblina). Táchira, Yaracuay; Mexico, Central America, West Indies, Colombia, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina. ◆Fig. 545. Palicourea crocea, P. croceoides, P. fastigiata, and P. tepuicola are similar species, and together form a morphologically variable complex. These have not been studied critically in the field in South America; they are here maintained provisionally and separated quite arbitrarily pending such study, and specimens are sometimes difficult to identify with confidence. Palicourea croceoides Desv. ex Ham., Prodr. Pl. Ind. Occ. 29: 1825. Palicourea riparia Benth., J. Bot. (Hooker) 3: 224. 1841. —Palicourea

crocea var. riparia (Benth.) Griseb., Fl. Brit. W. Ind. 345. 1864. Palicourea crocea var. riparia f. heterodoxa Steyerm., Mem. New York Bot. Gard. 23: 739. 1972. Shrub or small tree to 5 m tall, puberulous to hirtellous or glabrescent; leaves 2 per node, 4.5–18 × 1.5–7 cm; petioles 3–20 mm long; stipules united around stem, sheath 0.5–1.5 mm long, lobes narrowly triangular, 1–8 mm long; inflorescence orange to red, peduncles 2–8 cm long, branched portion cylindrical, narrowly to broadly pyramidal, or subcorymbiform, 1.5–10 × 2–10 cm; floral bracts 0.5–1.5 mm long; calyx limb 0.5–1.5 mm long; corolla yellow to orange, tube 8–13 mm long, lobes 1.5–3.5 mm long; fruit 4–6 × 4.5–6 mm; pyrenes 2. Evergreen lowland to upland forests and clearings, sea level to 900 m; Delta Amacuro (Caño Orocoima, Jotajana), Bolívar (Ciudad Bolívar, Gran Sabana, Río Paragua, Santa Elena de Uairén, Urimán), Amazonas (Puerto Ayacucho, Río Casiquiare, Río Orinoco, Río Sipapo). Anzoátegui, Apure, Barinas, Guárico, Miranda, Sucre, Táchira, Zulia; Antilles, Trinidad, Guyana, Suriname, French Guiana, Colombia, Ecuador, Peru, Brazil, Bolivia. Steyermark (1972; 1974) included this species in his circumscription of Palicourea crocea. He recognized two forms of what is this species based on details of the pubescence, but pubescence pattern varies widely throughout the range of this species and these forms are not recognized here. Palicourea fastigiata Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 3: 368. 1819. —Psychotria fastigiata (Kunth) Spreng., Syst. Veg. 1: 742. 1825. Shrub to 5 m tall, hirtellous or puberulous to glabrescent; leaves 2 per node, 4–14 × 1.5– 7.5 cm; petioles 3–20 mm long; stipules united around stem, sheath 0.5–1 mm long, lobes narrowly triangular, 1–2 mm long; inflorescence red to orange-red, peduncle 3.5– 10 cm long, branched portion corymbiform, 2–7 × 1.5–8 cm; floral bracts 0.5–1 mm long; calyx limb 0.8–1.2 mm long; corolla yellow to orange, tube 8–13 mm long, lobes ca. 2 mm long; fruit 4–4.5 × 4 mm; pyrenes 2. Riparian thickets usually in the influence of black-water rivers, 50–100 m; Bolívar (Reserva Imataca, Río Caura), Amazonas (near Atures). Colombia, Ecuador, Peru, Brazil.

686

R UBIACEAE

Palicourea foldatsii Steyerm., Mem. New York Bot. Gard. 23: 77. 1972. —Palo de sortija. Shrub to 3 m tall, glabrescent; leaves 2 per node, 4.5–10.5 × 2.5–7 cm; petioles 0–5 mm long; stipules united around stem, sheath ca. 1 mm long, lobes 1–2 mm long, triangular, acute; inflorescence orange to red, peduncle 2–2.5 cm long, branched portion corymbiform, 1.2–1.5 × 1.3–3.5 cm; floral bracts 0.8– 1.5 mm long; calyx limb 1–1.3 mm long; corolla orange, tube ca. 6 mm long, lobes ca. 1 mm long; fruit ca. 3 × 3 mm; pyrenes 2. Low caatinga forests on white sand, savannas on white sand, riparian forests, 100–200 m; southwestern Amazonas (Río Autana, Río Casiquiare, Río Guainía, Río Negro). Eastern Colombia. Palicourea glabriflora Steyerm., Mem. New York Bot. Gard. 23: 735. 1972. Shrub to 1.5 m tall, pilosulous to glabrescent; leaves 2 per node, 11.5–12.5 × 4.5–5 cm; petioles 3–5 mm long; stipules united around stem, sheath 0.7–1 mm long, lobes deltoid, 0.8–1.3 mm long; inflorescences orange, peduncles 6–7 cm long, branched portion corymbiform to broadly triangular, ca. 4 × 5.5 cm; calyx limb ca. 0.6 mm long; corolla yellow-orange at base to pink at apex, tube ca. 16 mm long, lobes ca. 4.5 mm long; fruit unknown. Forests on granitic outcrops, 100– 300 m; southwestern Amazonas (Piedra Tururumeri in Río Casiquiare basin). Endemic. Palicourea glabriflora is known only from the type. It may eventually be shown to be a form of P. longiflora with externally glabrous corollas. Palicourea grandiflora (Kunth) Standl., Publ. Field Columbian Mus., Bot. Ser. 7: 466. 1931. —Nonatelia grandiflora Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 3: 422. 1819. —Palo de bachaco, Palo de sortija. Shrub or small tree to 6 m tall, puberulous to glabrescent; leaves 2 per node, 11–30 × 3– 14 cm; petioles 7–22 mm long; stipules united around stem, sheath 0.8–2 mm long, lobes 0.8–3 mm long; inflorescence orange to red, peduncle 3.5–15 cm long, branched portion broadly pyramidal, 2.5–6.5 × 2–11 cm; calyx limb 0.3–0.8 mm long; corolla yellow to orange, tube 14–20 mm long, lobes 2.5–4.5

mm long; fruit 7–8 × 7–8 mm; pyrenes 2. Riparian forests, evergreen forests on white sand, tepui slope forests, white-sand savannas, 100–400 m; Amazonas (basin of Río Guainía and Río Orinoco). Amazonian Colombia, Ecuador, Peru, Brazil, and Bolivia. ◆Fig. 550. Palicourea amapaensis Steyerm. of northeastern Brazil, French Guiana, and Guyana is very similar to this species; P. amapaensis can be distinguished by its purple flowers and inflorescences, corolla tubes 22–30 mm long, and fruits 9–11 × 8–10 mm. Palicourea grandifolia (Willd. ex Roem. & Schult.) Standl., Field Mus. Publ. Bot. 11: 226. 1936. —Psychotria grandifolia Willd. ex Roem. & Schult., Syst. Veg. 5: 190. 1819. Nonatelia macrophylla Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 3: 423. 1819. Psychotria amazonica Müll Arg. in Mart., Fl. Bras. 6(5): 225. 1881. Psychotria sprucei Müll. Arg. in Mart., Fl. Bras. 6(5): 226, pl. 30, fig. 1. 1881. —Palicourea sprucei (Müll. Arg.) K. Schum. in Engl. & Prantl, Nat. Pflanzenf. 4(4): 115. 1891, “spruceana.” —Palicourea grandifolia var. sprucei (Müll. Arg.) Steyerm., Mem. New York Bot. Gard. 23: 725. 1972. Tree or shrub to 10 m tall, hirtellous, pilosulous, puberulous, or glabrescent; leaves 2 per node, 20–34 × 9–16 cm; petioles 7–35 mm long; stipules united around stem, sheath 3–4 mm long, lobes 3–10 mm long; inflorescence red to orange, peduncles 6–20 cm long, branched portion pyramidal, 5–15 × 5– 10 cm; floral bracts 0.5–1.5 mm long; calyx limb 0.5–1.2 mm long; corolla orange to yellow, tube 11–17.5 mm long, lobes 3.5–5.5 mm long; fruits 4–5 × 4–5 mm; pyrenes 4–5. Forest understory, borders, and clearings, forested igneous outcrops, tree islands in savannas, open places along streams, 100–1000 m; Bolívar (base of Cerro Guaiquinima), Amazonas (region of Puerto Ayacucho and San Carlos). Amazonian Colombia, Peru, Brazil. ◆Fig. 546. Steyermark separated Palicourea grandifolia from P. guianensis by several quantitative characters, but these species intergrade to some extent and a number of specimens do not have all the features that distinguished one of these species. Palicourea grandifolia

Palicourea 687

comprises, in general, the relatively largest plants of this complex; it is recognized here provisionally. One of the characters that Steyermark used to distinguish P. grandifolia was its secondary leaf veins 18–21 pairs, but the type collection of this species has several leaves with only 17 pairs of secondary veins. Steyermark recognized two varieties of Palicourea grandifolia, distinguished by the presence (var. sprucei) versus absence or notable sparsity (var. grandifolia) of pubescence on the leaves. He reported both varieties from the flora area, but the pubescence is continuously variable and these varieties are not recognized here. Palicourea guianensis Aubl., Hist. Pl. Guiane 173, pl. 66. 1775. —Psychotria guianensis (Aubl.) Raeusch., Nomencl. Bot. ed. 3, 56. 1797. Mexico, Central America, Antilles, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, 2 subspecies, 1 in the flora area. Steyermark (1972) recognized three subspecies, two varieties, and two forms of Palicourea guianensis. Two of these subspecies were distinguished primarily by details of the pubescence. The third, subsp. guianensis, was distinguished by its ovaries with 3 or 4 locules and its allopatric range in French Guiana, Suriname, and northern Brazil, versus 2-locular and distributed elsewhere in the other two subspecies. P. guianensis subsp. guianensis Palicourea guianensis subsp. occidentalis Steyerm., Mem. New York Bot. Gard. 23: 729. 1972. Palicourea guianensis subsp. occidentalis var. occidentalis f. glabra Steyerm., Mem. New York Bot. Gard. 72: 730. 1972. Shrub or small tree to 8 m tall, puberulous or hirtellous to glabrescent; leaves 2 per node, 13–43 × 6–18 cm; petioles 10–45 mm long; stipules united around stem, sheath 1– 3 mm long, lobes 3–13 mm long; inflorescences yellow to orange or red, peduncle 8–15 cm long, branched portion pyramidal, 9–19 × 7–15 cm; floral bracts 0.5–1.5 mm long; calyx limb 0.3–0.8 mm long; corollas red to orange, yellow, or white, tube 12–23 mm long, lobe 3– 5 mm long; fruit 4–7 × 3.5–5 mm; pyrenes 2.

Lowland forests, montane forested slopes, savannas, borders of wet forests, 100–1300 m; eastern Bolívar, Amazonas (Cerro Duida, basin of Río Guainía and Negro Río). Zulia; Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 544. Palicourea huberi Steyerm., Pittieria no. 9: 181, fig. 2. 1981. Shrub to 2 m tall, hirtellous; leaves 2 per node, 8.5–21 × 4.5–9.5 cm; petioles 1–5 mm long; stipules united around stem, sheath 1– 2 mm long, lobes 5–12 mm long, narrowly triangular; inflorescence red, peduncle 3–6 cm long, branched portion corymbiform, 2.5–3 × 3.5–4.5 cm; floral bracts 2–5 mm long; calyx limb 1–1.2 mm long; corolla pink to deep red, tube 6–7 mm long, lobes 1–1.5 mm long; fruit ca. 4 × 4 mm; pyrenes 2. Among shrub islands in wet savannas and white-sand savannas along streams, 100–200 m; Amazonas (Caño Yagua, basin of Río Orinoco). Northern Brazil. Palicourea lancigera (Standl.) Steyerm., Mem. New York Bot. Gard. 23: 762. 1972. —Psychotria lancigera Standl., Field Mus. Nat. Hist., Bot. Ser. 11: 240. 1936. Shrub to 2.5 m tall, glabrescent; leaves 2 per node, 4–9 × 1.2–2.5 cm; petioles 0–1 mm long; stipules united around stem, sheath 0.5–1 mm long, lobes 2–3 mm long, narrowly triangular; inflorescence pink, peduncle 0.8– 4 cm long, branched portion corymbiform, 1– 1.5 × 1–2 cm; floral bracts 0.5–1 mm long; calyx limb 1–1.5 mm long; corolla pink, tube ca. 6.5 mm long, lobes ca. 2 mm long; fruit ca. 3 × 3 mm; pyrenes 2. Flooded white-sand savannas, 100–200 m; southwestern Amazonas (basin of Río Guainía and Río Orinoco). Endemic. Palicourea longiflora DC., Prodr. 4: 528. Sep. 1830. —Psychotria longiflora Poir. in Lam., Encycl. 5: 704. 1804, nom. illeg., not Willd. 1797. Nonatelia longiflora Aubl., Hist. Pl. Guiane 185, pl. 171. 1775. —Psychotria longiflora Willd., Sp. Pl. 1: 971. 1797, not Poir. 1804. —Palicourea longiflora (Aubl.) A. Rich., Mem. Rubiaceae 95, pl. 9, fig. 2. 1829 [Dec. 1830]. Shrub or tree to 3(–7) m tall, glabrescent; leaves 2 per node, 6–20 × 3.5–8 cm; petioles

688

R UBIACEAE

3–11 mm long; stipules united around stem, sheath 0.5–1 mm long, lobes 0.5–1 mm long; inflorescence orange to red, peduncle 2–8.5 cm long, branched portion broadly pyramidal to corymbiform, 3–6 × 4–8 cm; floral bracts 0.5–2 mm long; calyx limb 0.2–0.5 mm long; corolla yellow to orange at base, pink to red or purple at apex, tube 14–28 mm long, lobes 2–3 mm long; fruit 4–5 × 5–6 mm; pyrenes 2. Evergreen lowland forests, tepui slope forests, 100–1000 m; Amazonas (Cerro Aratitiyope, Río Guainía, Río Negro, Río Orinoco, Río Ventuari). Guyana, Suriname, French Guiana, Ecuador, Brazil. Palicourea longiflora is similar to Palicourea marcgravii St.-Hil., which appears to replace it in drier habitats in the southern Guianas through southeastern Brazil. These species appear to intergrade in intermediate habitats, and specimens can be difficult to identify. Palicourea marcgravii differs from P. longiflora in its stipule lobes 1–2 mm long and its calyx limb that is 0.5– 1.3 mm long and usually lobed for 1/2 or more of its length, versus subtruncate to lobed for less than 1/2 in P. longiflora. Palicourea longiflora is often distinctive in fruit because the pedicels frequently become quite thickened. Palicourea longistipulata (Müll. Arg.) Standl., Publ. Field Columbian Mus., Bot. Ser., 7: 140. 1930. —Psychotria longistipulata Müll. Arg. in Mart., Fl. Bras. 6(5): 248. 1881. Shrub or tree to 5 m tall, densely velutinous to pilosulous; leaves 2 per node, 12–40 × 3.5–15 cm; petioles 4–20 mm long; stipules united around stem, sheath 2–4.5 mm long, lobes 3–26 mm long; inflorescence yellow to orange or red, peduncle 1.5–10 cm long, branched portion corymbiform, 2–7.5 × 3–13 cm; floral bracts 1–7 mm long; calyx limb 1–1.3 mm long; corolla orange, red, purple, or violet, tube 8–12 mm long, lobes 1– 1.5 mm long; fruit 4–5 × 3–4 mm; pyrenes 2. Colombia, Venezuela, Guyana, Suriname, French Guiana, Peru, Brazil; 2 subspecies, both in the flora area. Palicourea longistipulata generally shows clinal morphological variation from west to east across its range. Its two subspecies are usually quite clearly separable and allopatric, though a few plants from their contact zone may be difficult to classify. The stipule length is generally consistent in subsp.

chrysorrhachis but varies widely and apparently locally in subsp. longistipulata. Steyermark distinguished these subspecies by stipule lobe length, but they are here distinguished largely by inflorescence characters, which are more consistent. Key to the Subspecies of P. longistipulata 1. Stipule lobes 3–7 mm long; floral bracts 1– 2 mm long; inflorescence axes yellow, corollas orange to red .................................. ........................ subsp. chrysorrhachis 1. Stipule lobes 3–26 mm long; floral bracts 3–6 mm long; inflorescence axes orange to red, corollas red to purple or violet .......................... subsp. longistipulata P. longistipulata subsp. chrysorrhachis (Bremek.) Steyerm., Mem. New York Bot. Gard. 72: 748. 1972. —Palicourea chrysorrhachis Bremek., Brittonia 8: 244. 1957. Palicourea aculeifera Steyerm., Act. Bot. Venez. 2: 331, fig. 25. 1967. Lowland to montane forests, 200–1500 m; southeastern Bolívar (Auyán-tepui, Macizo del Chimantá). Guyana, Suriname, French Guiana, northeastern Brazil (Pará). P. longistipulata subsp. longistipulata. —Carriso, Carriso de picure, Oreja de picure. Psychotria sancti-caroli Standl., Publ. Field Columbian Mus., Bot. Ser. 7: 458. 1931. Evergreen lowland and caatinga forests, 100–200 m; Amazonas (basin of Río Guainía and Río Negro, San Carlos, Yavita). Colombia, Amazonian Peru, central to western Brazil. ◆Fig. 549. Palicourea nitidella (Müll. Arg.) Standl., Publ. Field Columbian Mus., Bot. Ser. 7: 142. 1930. —Psychotria nitidella Müll. Arg. in Mart., Fl. Bras. 6(5): 250. 1881. —Carriso sortijo, Kurucosadíyu (Yekwana). Shrub or tree to 5 m tall, glabrous; leaves 2 per node, 10.5–31 × 3.5–12 cm; petioles 5– 30 mm long; stipules united around stem, sheath 1–1.5 mm long, lobes narrowly triangular, 2–7 mm long; inflorescence red to orange, peduncle 5–11.5 cm long, branched portion corymbiform, 2–5.5 × 3–8 cm; calyx limb 0.5–1 mm long; corolla red to salmon with

Palicourea 689

lobes yellow to white, tube 11–14 mm long, lobes 1.5–2 mm long; fruit 4 × 4–4.5 mm; pyrenes 2. Evergreen lowland and riparian forests, forested igneous outcrops, borders of savannas and forests, palm forests, 50–900 m; Bolívar (widespread), Amazonas (Río Casiquiare, Río Cunucunuma, Río Guainía, Río Negro, widespread in basin of Río Orinoco, Río Parú, Río Siapa). Amazonian Colombia, Peru, and Brazil. Palicourea obtusata K. Krause, Notizbl. Bot. Gart. Berlin-Dahlem 6: 210. 1914. Shrub or tree to 4 m tall, pilosulous to glabrescent; leaves 2 per node, 4.5–12 × 2–6 cm; petioles 6–45 mm long; stipules laminar, 4–5 mm long, lobed for 1/3–1/2, lobes acute; inflorescence yellow, orange, red, or purple, peduncle 2–7.5 cm long, branched portion pyramidal, 2–7.5 × 2–6.5 cm; floral bracts 1.5–2 mm long; calyx limb 0.8–2.2 mm long; corolla yellow to white, tube 11–12 mm long, lobes ca. 2.5 mm long; fruit ca. 4 × 3 mm; pyrenes 2. Dwarf forests, Bonnetia and wet dwarf forests on tepui summits and slopes, 1900–2800 m; southeastern Bolívar (Aparamán tepui, Ilú-tepui, Kukenán-tepui, Macizo del Chimantá west to Auyán-tepui, Murisipán tepui, Ptari-tepui, basin of Río Aponguao and Río Caroní, Roraima-tepui). Guyana. ◆Fig. 542. Palicourea ottohuberi J.H. Kirkbr., BioLlania Ed. Esp. 6: 400. 1997. Shrub to 1 m tall, puberulous to glabrescent; leaves 2 per node, 3–3.5 × 1.2–1.8 cm; petioles 2–3 mm long; stipules laminar, 0.8– 1.2 mm long, lobed to 1/2, lobes obtuse; inflorescence red, peduncle 1.2–1.4 cm long, branched portion ca. 1 × 1.5 cm, shape not noted; floral bracts 0.4–0.5 mm long; calyx limb 0.4–0.7 mm long; corolla yellow, tube 5– 5.5 mm long, lobes ca. 3.5 mm long; fruit 3.5– 4 × 2.5 mm; pyrenes 2. Dissected granitic mountain summits, ca. 2100 m; Bolívar (Sierra de Maigualida). Endemic. Palicourea ottohuberi is known only from the type; no material of this species has been seen by the authors of this flora treatment. Kirkbride reported that this species is similar in general to P. puberulenta Steyerm. and P. canaguensis Steyerm., which are found in montane forests in the Venezuelan Andes. Palicourea pensilis J.H. Kirkbr., BioLlania Ed. Esp. 6: 402. 1997.

Shrub to 1 m tall, glabrous; leaves 2 per node, 2.5–5 × 1.3–2.3 cm; petioles 3–4 mm long; stipules laminar, ca. 2.5 mm long, bilobed for almost 2/3, lobes acute; inflorescences purple, peduncle 0.9–1.4 cm long, branched portion ca. 2 × 1.5 cm, shape not noted; floral bracts 0.7–1.5 mm long; calyx limb ca. 0.6 mm long; corolla purple, tube ca. 14.5 mm long, lobes ca. 2 mm long; fruit unknown. Shrub formations and low forests on tepui summits, 2100–2200 m; Bolívar (Macizo del Chimantá). Endemic. Palicoura pensilis is known only from the type; no material of this species has been seen by the authors of this flora treatment. Kirkbride said this species is similar to P. pittieri Standl. of the Coastal Cordillera of northern Venezuela. In addition to the characters cited by him, these can be distinguished by their stipules, which are united around the stem into a continuous sheath in P. pittieri. Palicourea perquadrangularis Wernham, J. Bot. 55: 341. 1917. Panama, Colombia, Venezuela, Brazil; 4 varieties, all in Venezuela, 1 of these in the flora area. P. perquadrangularis var. guayanensis Steyerm., Mem. New York Bot. Gard. 23: 774. 1972. Shrub or small tree to 4 m tall, puberulous to glabrescent; leaves 2 per node, 13–30 × 5.5–12 cm; petioles 1.5–4 cm long; stipules united around stem, sheath 3–6 mm long, lobes 3–12 mm long, acute; inflorescence purple, peduncle 2–5 cm long, branched portion 14–28 × 5–12 cm, pyramidal; floral bracts 0.5–1 mm long; calyx limb 1–1.5 mm long; corolla purple, tube 8–11.5 mm long, lobes 1.5–2 mm long; fruit 3–4 mm diameter; pyrenes 2. Tepui summit and slope forests, 800–2000 m; Bolívar (Cerro Jaua, Cerro Venamo, Macizo del Chimantá, basin of Río Cuyuní), Amazonas (Cerro Duida, Cerro Huachamacari, Cerro Marahuaka, Sierra de la Neblina). Guyana, Amazonian Brazil. Steyermark recognized several varieties of Palicourea perquadrangularis that he separated by different combinations of some quantitative characters, notably pubescence density, pedicel length, and calyx lobe shape. These characters are probably continuously variable and the varieties thus not taxonomi-

690

R UBIACEAE

cally very informative, but their evaluation is outside the scope of this flora. The plants included in var. guayannensis are geographically quite isolated from the rest of the species, which is found in the Andes of Venezuela and Colombia. Steyermark distinguished var. guayannensis morphologically only by its obtuse calyx lobes, versus acuminate, acute, or obtuse in the other varieties. Palicourea quadrifolia (Rudge) DC., Prodr. 4: 526. 1830. —Psychotria quadrifolia Rudge, Pl. Guian. 1: 27, pl. 42. 1805. Colombia, Venezuela, Guyana, French Guiana, Ecuador, Peru, Bolivia; 2 subspecies, 1 in Venezuela. Subsp. leticiana C.M. Taylor is found in the western Amazon basin, and has leaves generally 2 per node. P. quadrifolia subsp. quadrifolia Shrub or small tree to 2(–5) m tall, puberulous to glabrescent; leaves 3 or 4 per node, 10.5–35 × 3–9 cm; petioles 0.3–1.5 cm long; stipules united around stem, sheath 2– 6 mm long, lobes 0.5–5 mm long, acute to subulate; inflorescence yellow, peduncle 4– 11.5 cm long, branched portion 4.5–9 × 6–12 cm, corymbiform; floral bracts 1.2–4 mm long; calyx limb 0.2–0.3 mm long; corolla yellow, tube 8–12 mm long, lobes 1–2 mm long; fruit ca. 3 × 5 mm; pyrenes 2. Evergreen lowland forests, 100–200 m; Amazonas (Río Putaco near the mouth of the Río Ocamo). Guyana, Suriname, French Guiana, Peru, Amazonian Brazil. Palicourea rigida Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 3: 370. 1819. —Capa rosa. Psychotria rigida Bredem. ex Willd. ex Roem. & Schult., Syst. Veg. 5: 192. 1815. [Although these are often cited as synonyms, this name is based on a different type from Palicourea rigida.] Palicourea rigida subsp. rigida var. hirtibacca Steyerm., Mem. New York Bot. Gard. 23: 732. 1972. Erect shrub or subshrub to 2.5(–8) m tall, often from a xylopodium, hirtellous to puberulous or glabrescent, older stems with thick corky bark; leaves 2 per node, 5–30 × 2.5–20 cm; petioles 1.5–10 mm long; stipules united around stem, sheath 1.5–2 mm long, lobes

ligulate to triangular, 2–4.5 mm long, acute to rounded; inflorescence yellow, orange, or red, peduncle 2–20 cm long, branched portion narrowly to broadly pyramidal, 2–16 × 1.5– 10 cm; floral bracts 1–3 mm long; calyx limb ca. 0.3 mm long; corolla yellow to orange, tube 9–11 mm long, lobes 2.5–3.5 mm long; fruit 5–6 × 4–5 mm; pyrenes 2. Savannas and open or exposed low vegetation, bordering streams, 50–1100 m; Delta Amacuro (Los Castillos, Piacoa), Bolívar (common throughout, especially in the Gran Sabana), Amazonas (near Puerto Ayacucho and Isla Ratón). Colombia, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia, Paraguay. Steyermark, as well as many previous authors, recognized a number of infraspecific taxa of Palicourea rigida. This species is quite variable morphologically, notably in pubescence, size and shape of the leaves, and degree of development of their secondary and tertiary venation. This variation has not been critically studied, so no well grounded conclusions can yet be drawn. Steyermark (1974) recognized two varieties in the flora area, var. rigida with “usually glabrous” leaves, inflorescences, and fruits, and var. hirtibacca with these structures puberulous. Pubescence varies widely and apparently continuously in P. rigida, and consequently separating these varieties does not seem informative and they are not recognized here. Palicourea tepuicola Steyerm., Mem. New York Bot. Gard. 23: 742. 1972. Shrub to 3 m tall, puberulous to glabrescent; leaves 2 per node, 13–19 × 4.5–10 cm; petioles 9–13 mm long; stipules united around stem, sheath 1–1.5 mm long, lobes narrowly triangular, 2–4 mm long; inflorescence red, peduncle 6–11 cm long, branched portion pyramidal, 4–8.5 × 5–8 cm; floral bracts 0.5–1 mm long; calyx limb ca. 0.8 mm long; corolla yellow to orange, tube 8–13 mm long, lobes ca. 2 mm long; fruit ca. 5 × 4 mm; pyrenes 2. Tepui slope and summit forests, 400–1500 m; Bolívar (Aprada-tepui, Auyántepui, basin of Río Caroní and Río Cuyuní, Uaipán-tepui). Endemic. ◆Fig. 547. Palicourea triphylla DC., Prodr. 4: 526. 1830. —Psychotria triphylla (DC.) Müll. Arg. in Mart., Fl. Bras. 6(5): 233. 1881. Shrub to 3 m tall, hirtellous to glabres-

Palicourea 691

cent; leaves (2)3(4) per node, 12–22 × 3.5–8.5 cm; petioles 5–10 mm long; stipules united around stem, sheath 1.5–2 mm long, lobes lanceolate, 10–16 mm long, acute; inflorescence red to orange, peduncle 4.5–15.5 cm long, branched portion narrowly pyramidal, 9–25 × 5–9 cm; floral bracts 1–4 mm long; calyx limb ca. 0.5 mm long; corolla red to orange or yellow, tube 7–10 mm long, lobes ca. 1 mm long; fruit 3.5–5 × 3.5–4 mm; pyrenes 2. Tall riparian forests, gallery forests, forest borders, base of forested talus slopes of table mountains overlying Roraima sandstone, 50–1000 m; Bolívar (eastern portion northeast to Altiplanicie de Nuria and west to Río Caura), Amazonas (Río Negro, basin of Río Orinoco). Mexico, Central America, Cuba, Colombia, Trinidad, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia. ◆Fig. 548. Palicourea wurdackiana J.H. Kirkbr., BioLlania Ed. Esp. 6: 403. 1997. Shrub to 1.5 m tall, puberulous to glabrescent; leaves 2 per node, 3.5–6.5 × 2–3.5 cm; petioles 3–5 mm long; stipules with the interpetiolar portion broadly emarginate, 0.5–1 mm long in middle, the intrapetiolar portion 1–2 mm long; inflorescence purple, peduncle 2.5–3 cm long, branched portion 2– 2.5 × 3–3.5 cm, rounded-corymbiform; floral bracts reduced or to 0.8 mm long; calyx limb 0.5–1 mm long; corolla white to purple, tube 7–9.5 mm long, lobes 2.5–3.5 mm long; fruit 6–7 × 7 mm; pyrenes 2. Shrub formations on mountain summits, 1800–2100 m; Bolívar (Sierra de Maigualida), Amazonas (Cerro Yutajé). Guyana. Kirkbride reported that Palicourea wurdackiana is similar to P. jahnii Standl., a yellow-flowered species of the Venezuelan Andes, and noted that the unusual stipules of P. wurdackiana distinguish it from all other species of Palicourea. However, another species, Psychotria everardii Wernham, also found widely in the Guayana Highlands, shares these unusual stipules and is very similar in general morphology to Palicourea wurdackiana. These species appear to be separable only by the arrangement of their flowers, with most or all of the flowers sessile in Psychotria everardii versus most of them pedicellate in Palicourea wurdackiana. Psychotria everardii is variable morphologically, and Palicourea wurdackiana may eventually be considered part

of this complex. Kirkbride described the floral bracts of Palicourea wurdackiana as lacking, but they are sometimes present (e.g., Huber 12633, US!, paratype). Palicourea sp. A Shrub or small tree to 5 m tall, glabrescent; leaves 2 per node, 8–28 × 25–12.5 cm; petioles 5–45 mm long; stipules united around stem, 0.5–2.5 mm long, lobes deltoid, 2–4 mm long; inflorescence yellow, peduncles 6–13 cm long, branched portion corymbiform to broadly pyramidal, 2.5–6 × 4–7 cm; floral bracts 0.5–1.2 mm long; calyx limb 0.5–0.8 mm long; corolla yellow, tube 17–18 mm long, lobes ca. 2.5 mm long; fruit unknown. Evergreen lowland forests, 50–300 m; Amazonas (region of San Carlos de Río Negro). Endemic. Palicourea sp. A is similar to and has been confused with P. guianensis, but is apparently distinct and will be described soon (Taylor, in preparation). It appears to be endemic to the San Carlos region.

Fig. 542. Palicourea obtusata

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R UBIACEAE

Fig. 543. Palicourea corymbifera

Fig. 544. Palicourea guianensis subsp. guianensis

Palicourea 693

Fig. 545. Palicourea crocea

Fig. 546. Palicourea grandifolia

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R UBIACEAE

Fig. 547. Palicourea tepuicola

Fig. 548. Palicourea triphylla

Fig. 549. Palicourea longistipulata subsp. longistipulata

Perama 695

Fig. 550. Palicourea grandiflora

64. PERAMA Aubl., Hist. Pl. Guiane 54, t. 18. 1775. by Charlotte M. Taylor and Julian A. Steyermark Perennial or annual herbs or subshrubs, terrestrial, unarmed. Leaves cauline or sometimes basal, opposite or verticillate, sessile or shortly petiolate, venation not lineolate, sometimes palmate; stipules reduced, glandular, or apparently sometimes obsolete. Inflorescences terminal and axillary, several- to multiflowered, pedunculate, capitate or glomerulate, the heads or glomerules solitary to numerous and paniculate or aggregated in spikes, bracts reduced or none. Flowers sessile, distylous, small, frequently subtended by groups of linear-setose, often dimorphic scales. Hypanthium obconic to cylindrical. Calyx limb deeply lobed, lobes 2, without calycophylls; corolla funnelform or salverform, yellow, blue, purple, roseate, or white, pubescent in the throat, lobes (3)4(5), valvate in bud. Stamens (3)4(5), inserted in corolla throat; anthers dorsifixed near base, included or exserted, narrowly ellipsoid. Ovary 2–4-locular; ovules solitary in each locule, axile. Fruits capsular, obpyramidal, membranaceous, circumscissile above the equator. Seeds ovoid to trigonous, small. Martinique, Colombia, Venezuela, Trinidad, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia; 12 species, 4 in Venezuela, all in the flora area. The highly reduced or obsolete stipules make these plants sometimes difficult to recognize as Rubiaceae. Some specimens of Limnosipanea are similar vegetatively to Perama; Limnosipanea can be distinguished by its numerous ovules and seeds and its corolla lobes that are convolute in bud. Perama is also similar to Oldenlandia; their separation is discussed under Oldenlandia. Key to the Species of Perama 1. 1.

Leaves all in basal rosettes ........................................................ P. dichotoma Leaves all cauline ...................................................................................... 2

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R UBIACEAE

2(1). 2. 3(2). 3.

Leaves 25–90 × 10–25 mm ....................................................... P. plantaginea Leaves 2–13 × 0.2–4.5 mm ........................................................................ 3 Leaves 3- or 4-verticillate .............................................................. P. galioides Leaves opposite or sometimes ternate at a few nodes ................... P. hirsuta

Perama dichotoma Poepp., Nov. Gen. Sp. Pl. 3: 29, pl. 235. [1841] 1845. Herbaceous annual; leaves opposite or congested to apparently verticillate, 5–50 × 4–22 mm, all borne at base of plant, hispid to hirsute or glabrescent; secondary veins 1 or 2 pairs, palmate; inflorescences paniculate or rarely capitate, 5–40 × 3–30 cm, heads or glomerules 1–3(–5) mm diameter; calyx lobes 0.5–1.2 mm long, triangular; corollas blue, blue-lavender, or white, tube 1–1.5 mm long, lobes 1–1.5 mm long; seeds 0.6–0.7 mm long. Moist spots on bare sandstone or igneous substrate, shaded slopes along streams, crevices of rock outcrops, sandy openings bordering forests, margins of dwarf shrub vegetation, rocky terrain in open scrub savannas, white-sand savannas, 100–2100 m; common in Bolívar and Amazonas. Colombia, Guyana, Brazil (upper Rio Negro); 2 varieties, both in the flora area. The leaves of Perama dichotoma are sometimes purple on their undersides; this character appears to have no taxonomic significance but is said to give the species an elegant appearance.

Duida, Cerro Marahuaka, Cerro Parú, Cerro Yapacana, Río Autana, Sierra de la Neblina). Guyana, Suriname, French Guiana, Brazil (Amazonas, Pará, Roraima). ◆Fig. 552. Steyermark distinguished three varieties of Perama dichotoma that here are treated as synonyms of var. dichotoma. These were distinguished by details of the density, type, and distribution of the pubescence on the lower surface of leaves. The upper leaf surfaces are similarly moderately to densely hirsute to hispid in all of these varieties. Steyermark (1963) noted that the pubescence on the lower surface of leaves ranged from glabrescent on the blade and strigose on the principal veins (var. scaposa), through glabrescent on the blade and hirsute to strigose along the principal veins (var. dichotoma), to hirsute or strigose throughout (var. hirsutula). No geographic patterns are evident in this variation, and as admitted by Steyermark (1963, 237), specimens from the flora region with all the intermediate pubescence conditions are in ample supply.

1. Inflorescence of several to numerous heads or glomerules 1–3 mm diameter borne in a branched inflorescence ....... .................................... var. dichotoma 1. Inflorescence a solitary head 4–5 mm diameter borne on a simple peduncle ................................ var. monocephala

P. dichotoma var. monocephala Steyerm., Ann. Missouri Bot. Gard. 74: 658. 1987. Sandstone slopes, 100–200 m; Amazonas (base of Cerro Huachamacari). Endemic. This specimen (Delascio 12358, MO, VEN) is highly different in inflorescence arrangement from all other specimens of this species. It may represent a reduced plant or deserve separate specific status; this variety is here provisionally recognized pending further study.

P. dichotoma var. dichotoma Perama scaposa Gleason & Standl., Publ. Field Columbian Mus., Bot. Ser. 7: 475. 1931. —Perama dichotoma var. scaposa (Gleason & Standl.) Steyerm., Mem. New York Bot. Gard. 10(5): 236, fig. 78D. 1963. Perama dichotoma var. hirsutula Steyerm., Mem. New York Bot. Gard. 10(5): 236. 1963. Damp sandstone faces of cliffs and along streams of upper tepui slopes and summits, savannas, 100–2300 m; Bolívar (widespread), Amazonas (Cerro Aracamuni, Cerro

Perama galioides (Kunth) Poir. in F. Cuvier, Dict. Sci. Nat. ed. 2, 38: 414. 1825. —Mattuschkea galioides Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 272. 1817 [1818]. Mattuschkea hispida Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 271. 1817 [1818]. —Perama galioides var. hispida (Kunth) Steyerm., Mem. New York Bot. Gard. 10(5): fig. 80A–C. 1963. Perama galioides var. densipila Steyerm., Mem. New York Bot. Gard. 10(5): 244, fig. 79E, F. 1963.

Key to the Varieties of P. dichotoma

Perama 697

Perama galioides var. galioides f. quaternata Steyerm., Mem. New York Bot. Gard. 10(5): 239. 1963. Perama galioides var. galioides f. macrocephala Steyerm., Mem. New York Bot. Gard. 10(5): 239. 1963. Perama galioides var. hispida f. cinera Steyerm., Mem. New York Bot. Gard. 10(5): 243. 1963. Perama galioides var. intermedia Steyerm., Mem. New York Bot. Gard. 10(5): 243. 1963. Perama galioides var. longiflora Steyerm., Mem. New York Bot. Gard. 10(5): 241. 1963. Herb or subshrub to 40 cm tall, glabrescent to densely strigose or hispid; leaves 3- or 4-verticillate, 3–13 × 0.2–4 cm, borne along stems; secondary veins 1 or 2 pairs, palmate and strongly ascending; inflorescences unbranched to paniculate, 5–15 × 3–12 cm, heads or glomerules 1–10, 3–15 mm long, 3– 6 mm diameter; calyx lobes 2–4.5 mm long, narrowly triangular; corolla yellow, tube 2.5– 6.5 mm long, lobes 1–2 mm long; seeds 0.5– 0.7 mm long. Wet savannas, white-sand savannas, shaded stream banks in gallery forests, thin woodland by dry sandstone outcrops, among igneous or sandstone outcrops, 50–1600 m; Bolívar (Auyán-tepui, Cerro Chirikayén, Cerro Guaiquinima, Cerro Perro in upper Río Paragua basin, Gran Sabana, Macizo del Chimantá, Ptari-tepui, Río Caroní), Amazonas (widespread). Anzoátegui, Apure, Cojedes; Colombia, Guyana, Brazil (Amazonas). ◆Fig. 554. Perama galioides is variable in habit, inflorescence size, stem pubescence, leaf shape, size, and pubescence, corolla length, and arrangement of the calyx lobes. Steyermark (1963; 1974) considered this species well circumscribed by its verticillate leaves and minutely scrobiculate seeds. He recognized five varieties and five forms of P. galioides, noting (1963, 245–246) that this large number of infraspecific taxa was necessary to accommodate the numerous specimens that are intermediate between “the various infraspecific taxa,” and that he chose the varietal level because these were so poorly differentiated. The leaves of Perama galioides are palmately veined with the secondary veins well marked and strongly ascending, to the point that the leaves may appear parallel-veined. The verticillate leaves are shortly united

around the stem at their bases but stipules are not evident. Thus, this species is not easily recognized as Rubiaceae, and may also be confused with Limnosipanea; Limnosipanea can be distinguished by its flowers that are borne singly along the dichasial inflorescence axes and have 5 calyx lobes (versus densely grouped in heads and glomerule and with 2 calyx lobes in Perama). Perama hirsuta Aubl., Hist. Pl. Guiane 54, pl. 18. 1775. Perama stricta Benth., J. Bot. (Hooker) 3: 239. 1841. —Perama hirsuta var. stricta (Benth.) Bremek., Recueil Trav. Bot. Neérl. 31: 308. 1934. Herb to 40 cm tall, annual or perhaps sometimes perennial, hirsute or strigose to glabrescent; leaves paired or sometimes ternate on some nodes, 3–12 × 1–5 mm, borne along stems; secondary veins 1–3 pairs, palmate, ascending; inflorescences unbranched or branched 2–4 times, 6–14 × 1–6 cm, heads 1–6, 4–10 × 4–10 mm; calyx lobes 1–2 mm long, narrowly triangular; corolla yellow, tube 1.5–2 mm long, lobes 0.5–1 mm long; seeds 0.5–0.7 mm long. Dry or moist savannas, wet savannas bordering Mauritia palm swamps, wet open igneous outcrops, sandy openings bordering forests, 100–1300 m; Bolívar (Altiplanicie de Nuria, Gran Sabana, Los Pijiguaos, Río Canaracuni, Río Paragua, Río Parguaza), Amazonas (La Esmeralda, Río Autana, basin of Río Casiquiare, Río Manapiare, Río Yutajé). Anzoátegui, Guárico, Monagas; Martinique, Colombia, Trinidad, Guyana, Suriname, French Guiana, central to eastern Brazil, Bolivia. ◆Fig. 553. Steyermark (1963, 248–250) recognized two varieties of Perama hirsuta that he noted were connected by “various intergradations,” and even cited a number of intermediate specimens. The range of his var. stricta was given as “rare and scattered in the range of var. hirsuta.” Consequently these varieties are not recognized here. Perama plantaginea (Kunth) Hook. f. in Benth. & Hook., Gen. Pl. 2: 148. 1873. —Buchia plantaginea Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 2: 270, pl. 132. 1817 [1818]. Suffrutescent perennial to 0.4 m tall, densely silky-sericeous; leaves opposite, 25– 90 × 10–25 mm, borne along stems; second-

698

R UBIACEAE

ary veins 2–5 pairs, subpalmate to palmate; inflorescences densely spiciform; peduncles (absent or) 4–13 cm long; spikes 1–5 per stem, simple or with 2–6 fasciculate segments, spikes or segments 3–20 × 3–8 mm; calyx lobes 1–2 mm long, ovate to triangular; corolla white, tube ca. 1.5 mm long, lobes ca. 1 mm long; seeds 0.5–1 mm long. Scrubby forests on white sand, flooded or semi-flooded white-sand forests, 100–200 m; southwestern Amazonas (Río Guainía, basin of Río Negro, Río Orinoco, Río Pasimoni, north to Río Sipapo). Amazonian Colombia and Brazil. ◆Fig. 551.

Fig. 552. Perama dichotoma var. dichotoma

Fig. 553. Perama hirsuta

Fig. 551. Perama plantaginea

Fig. 554. Perama galioides

Platycarpum 699

65. PLATYCARPUM Bonpl. in Humb. & Bonpl., Pl. Aequinoct. 2: 81, pl. 104. 1811. by Julian A. Steyermark and Charlotte M. Taylor Trees or shrubs, terrestrial, unarmed. Leaves opposite or in whorls of 3 or 4, petiolate, venation not lineolate, the petioles usually with 1–3(–5) glands at the abaxial base; stipules calyptrate (i.e., fused into a conical cap), caducous. Inflorescences terminal, multiflowered, pedunculate to subsessile, cymose-paniculate with dichasial axes, bracteate. Flowers medium to large, showy, homostylous, protandrous. Hypanthium turbinate. Calyx limb deeply lobed, lobes 4 or 5, without calycophylls; corolla funnelform or campanulate-funnelform, rather zygomorphic, externally white to rose-colored, internally streaked reddish, internally glabrous except with a yellow beard in the throat and a pubescent ring at the base of the tube, lobes 5, in bud imbricate with one lobe external. Stamens 5, inserted at varied levels in lower portion of corolla tube; anthers narrowly elliptic, dorsifixed, exserted. Ovary partially inferior, 2-locular; ovules 2 in each locule, axile. Fruit partially inferior to apparently superior, capsular, oblong to orbicular and laterally strongly flattened parallel to septum, chartaceous, with a horizontal scar across the equator, loculicidally dehiscent along margins, after dehiscence the valves reflexed and separated in a distinctive arrangement. Seeds oblong-reniform, elliptic, or Dshaped, flattened, large, margins winged. Amazonian Colombia, Venezuela, Guyana, Peru, Amazonian Brazil; 12 species, 8 in Venezuela, all in the flora area. Platycarpum is notable for its zygomorphic corollas and its ovaries that are partially inferior in flower and in fruit become nearly to apparently fully superior due to the enlargement of the top part of the ovary. Henriquezia is similar to Platycarpum in fruit morphology, and Coutarea, Henriquezia, and Gleasonia are similar to it in flower size and shape; the distinctions among these are outlined under Henriquezia. Key to the Species of Platycarpum 1. 1. 2(1).

2.

3(1). 3. 4(3).

4.

5(4).

Leaves in whorls of 3 or 4 .......................................................................... 2 Leaves in pairs ........................................................................................... 3 Calyx lobes 4; glands at petiole bases solitary, rounded; lower surfaces of leaves glaucous or silvery green, the costa and secondary veins glabrous ........................................................................... P. rhododactylum Calyx lobes 5; glands at petiole bases solitary or 3–5, U-shaped; lower surfaces of leaves green, the costa and secondary veins strigose to hirsute ............................................................................................. P. rugosum Petioles and young stems glabrous or appressed-puberulous; lower surfaces of leaves puberulous, silvery or glaucous .................................... 4 Petioles and young stems densely villous or hirsute; lower surfaces of leaves pubescent and green ................................................................... 6 Corolla 10–16 mm long, the lobes 6–9 mm long; stipules glabrous; lower surfaces of leaves with the costa glabrous or puberulous; trees or shrubs 10 m tall or shorter; in savanna habitats ..................... P. negrense Corolla 15–26 mm long, the lobes 8–16 mm long; stipules strigose or hirsute at least on basal portion; lower surfaces of leaves with the costa strigose; trees 25–60 m tall; in montane or lowland forests ................ 5 Corolla lobes > 1/2 as long as the corolla; inflorescence with the higher-

700

5. 6(3).

6.

7(6). 7.

R UBIACEAE

order axes and pedicels sparsely to moderately strigulose or puberulous ............................................................................................ P. decipiens Corolla lobes 1/3–1/2 as long as the corolla; inflorescence with the higherorder axes and pedicels densely rufous-pilose to -hirsute ....... P. maguirei Corolla 10–12 mm long, the lobes < 3 mm wide; calyx lobes externally with conspicuous hoary-frosted appearance; capsule 16–22 mm long; seeds 15–20 mm long; petioles 5 mm long or shorter ............. P. schultesii Corolla 12–22 mm long, the lobes 3–10 mm wide; calyx lobes externally cinereous- to ferruginous-villous; capsule 25–50 mm long; seeds 25– 50 mm long; petioles 5–25 mm long...................................................... 7 Leaves oblanceolate, acute to shortly acuminate at apex; capsules 4.5– 5 cm long ........................................................................................ P. duckei Leave elliptic-oblong to elliptic, obtuse to rounded at apex; capsules 2.5– 4.5 cm long .............................................................................. P. orinocense

Platycarpum decipiens Woodson & Steyerm., Amer. J. Bot. 39: 422. 1952. —Picatón. Large tree 25–60 m tall, with dark scaly bark; leaves 7.5–17 × 2–7 cm, stipules 1–6 cm long; inflorescences 8–17 × 5–16 cm; calyx lobes 5, 6–12 mm long; corolla 1.3–2.5 cm long, white or pink externally, red in throat; capsule 4–6 × 3.5–5.5 cm. Evergreen lowland forests, mostly on white-sand soils, 100–200 m; Amazonas (Caño San Miguel, San Carlos de Río Negro to Solano, Yavita to Maroa). Endemic. Platycarpum duckei Steyerm., Amer. J. Bot. 39: 422. 1952. Large tree; leaves 7–35 × 2.8–11 cm; stipules 0.5–5 cm long; inflorescences 8–18 × 7–13 cm; calyx lobes 5, 4–9 mm long; corolla 1.2–2 cm long, externally white, interally with red markings; capsule 4.5–5 × 3.5–4 cm. Río Negro caatinga forests on white sand, 100–200 m; Amazonas (near Yavita). Northern Brazil. Platycarpum duckei is included here based on 2 sterile specimens from the flora region (Berry 6298, MO; Aymard 11220, MO). The combination of pubescence, stipule morphology, and leaf arrangement, size, and shape is distinctive for this species. This is a new record for Venezuela. The measurements of reproductive structures are based entirely on Brazilian plants. Platycarpum maguirei Steyerm., Mem. New York Bot. Gard. 10: 255, fig. 76I–J. 1963. Tree to 25 m tall; leaves 10–25 × 2–8 cm; stipules not seen; inflorescences 10–15 × 4–

10 cm; calyx lobes 5, 7–11 mm long; corolla 1.5–2.6 cm long, white with tawny hairs externally, internally streaked red; capsules not seen. Lower montane forests, 100–700 m; Amazonas (Sierra de la Neblina). Endemic. Platycarpum negrense Ducke, Arq. Inst. Biol. Veg. 1: 212. 1935. Venezuela, Brazil; 2 varieties, 1 in Venezuela. P. negrense var. glaucum G.K. Rogers, Fl. Neotrop. Monogr. 39: 114, fig. 208. 1984. Shrub or small tree to 10 m; young stems red-brown to dark purple; leaves 6–19 × 2.5– 8.5 cm, with lower surface glaucous; stipules 1.5–3 cm long; inflorescences and flowers unknown (in var. negrense, inflorescences 12– 16 × 6–9 cm; calyx lobes 5, 6–9 mm long; corolla 0.8–1.6 cm long, externally white to pink, internally spotted); capsules 2.5–3.3 × 2.3–3 cm. Seasonally flooded white-sand savannas, 100–200 m; Amazonas (Santa Cruz del Atabapo). Endemic. Platycarpum orinocense Bonpl. in Humb. & Bonpl., Pl. Aequinoct. 2: 81, pl. 104. 1811, “orenocense.” —Chaparro de sabana, Picatón, Platanote. Tree with somewhat irregular, slightly crooked trunk 4–20 m tall; bark gray, rough, with rectangular segments; leaves 10–28 × 4–13 cm; stipules 2–9 cm long; inflorescences 8.5–25 × 7–32 cm; flowers fragrant; calyx lobes 5, 6–12 mm long; corolla 1.2–2.2 cm long, externally white to pink, internally with white lobes, throat red-streaked, tube red at base; capsule 2.5–4.5 cm. Tree savannas and neighboring gallery forests, whitesand savannas, granitic outcrops, shrub is-

Platycarpum 701

lands in savannas, montane savannas, 100– 500(–900) m; Amazonas (Cerro Calentura in upper Río Parucito basin near border with Bolívar state, lower slopes of Cerro Moriche, western base of Cerro Parú, slopes of Cerro Yapacana, from ca. 30 km north of Puerto Ayacucho south to Samariapo, upper Río Asisa, Río Ventuari 15 km below mouth of Río Manapiare, from Santa Bárbara del Orinoco to Carmelitas). Adjacent eastern Colombia, Amazonian Peru; 2 varieties, both in the flora area. The wood is employed for fuel and supports. Key to the Varieties of P. orinocense 1. Corolla 1.9–2.2 cm long; calyx lobes 9–12 mm long; axes of the basal portion of the inflorescence with pubescence usually in a single layer, ferruginous-hispid or pilose, the trichomes shorter than or rarely as long as 1.3 mm long; lower surface of leaves pilose or villous with trichomes to 2.5(–3) mm long ...................... ............................... var. grandiflorum 1. Corolla 1.2–2.2 cm long; calyx lobes 6–9 mm long; axes of the basal portion of the inflorescence with pubescence in 2 distinct layers, the shorter layer often cinerous-tomentose, the longer layer consisting of scattered, light to ferruginous trichomes often > 1.3 mm long .................................... var. orinocense P. orinocense var. grandiflorum Steyerm., Mem. New York Bot. Gard. 10: 254. 1963. Northeastern Amazonas (near Bolívar border, Cerro Calentura in upper Río Parucito basin, Río Parucito headwaters). Endemic. Variety grandiflorum was recognized by Rogers (1984), but re-measurement of some of the original material suggests that it may represent an extreme of continuous variation rather than a clearly differentiated taxon. P. orinocense var. orinocense Sickingia orinocensis Spreng., Syst. Veg. 1: 622. 1825 [1824]. Henriquezia aturensis Standl., Publ. Field Columbian Mus., Bot. Ser. 7: 368. 1931. The typical variety is found throughout the range of the Platycarpum orinocense, and is the most commonly collected taxon of this genus. ◆Fig. 555.

Platycarpum rhododactylum Woodson & Steyerm., Amer. J. Bot. 39: 423. 1952. Shrub or small tree to 9(–15) m tall; bark dark gray, with rectangular segments; leaves 7–21 × 3–11 cm; stipules 1.2–2 cm long; inflorescences 4–9 × 4–10 cm; calyx lobes 4, 4–8 mm long; corolla 1–2 cm long, externally pink with dense white to tawny pubescence, internally pink with red spots; capsule 2–3.7 × 2.5–4.7 cm. Dry rocky sandstone outcrops, shrubby formations in rocky terrain or savanna habitats, 300–1300 m; Bolívar (area around Canaima, Cerro Guaiquinima, Cerro Tonoro in upper Paragua basin, Kurún-tepui near Cerro Venado, upper Río Caroní). Endemic. Platycarpum rugosum Steyerm., Mem. New York Bot. Gard. 10: 257. 1963. Tree to 15(–20) m tall; bark gray, with rectangular segments; leaves 8–23 × 3.5–9 cm; stipules 1.5–6.6 cm long; inflorescences 3–7 × 4–7 cm; calyx lobes 5, 4–7 mm long; corolla 1–1.5 cm long, externally pink with white- to rusty-sericeous pubescence, internally pink with red markings; capsule 3.5–5 × 3.5–6.5 cm. Tepui slope and summit forests, 1100–1800 m; Bolívar (Ilú-tepui, plateau above La Escalera, Mukuripá, Ueitepui). Endemic. Platycarpum schultesii Steyerm., Bot. Mus. Leafl. 17: 96. 1955. Shrub or small tree to 9 m tall. Venezuela, eastern Colombia; 2 varieties, 1 in Venezuela. Var. schultesii is known from eastern Colombia, and is a larger plant than var. zarucchii. Its leaves are ternate instead of paired as in var. zarucchii. Both varieties are known from relatively few collections, and although floral and fruit size differences have been used to separate them, the size ranges overlap significantly. P. schultesii var. zarucchii G.K. Rogers, Fl. Neotrop. Monogr. 39: 126. 1984. Shrub ca. 1 m tall; leaves 6–13 × 2.5–8 cm; stipules 0.5–4.5 cm long; inflorescences 10– 15 × 4–7 cm; calyx lobes 5, 6–8 mm long; corolla 1–1.5 cm long, externally pink, densely white-sericeous, internally pink with red marks; capsules 1.5–2.2 × 2.2–3 cm. Whitesand savannas and Río Negro caatinga, 50– 100 m; Amazonas (Canaripó, region of Cerro Yapacana, Río Guasacavi, in basin of Río Guainía, Río Orinoco, and Río Ventuari, Yavita). Endemic.

702

R UBIACEAE

Fig. 555. Platycarpum orinocense var. orinocense

66. POSOQUERIA Aubl., Hist. Pl. Guiane 133. 1775. by Charlotte M. Taylor and Julian A. Steyermark Trees or shrubs, terrestrial, unarmed. Leaves opposite, petiolate, venation not lineolate; stipules interpetiolar, persistent or deciduous, triangular to lanceolate, generally erect and flatly appressed in bud. Inflorescence terminal, cymose-corymbose to subumbellate, several- to multiflowered, subsessile to pedunculate, bracts usually present but reduced. Flowers large, showy, pedicellate, homostylous, protandrous, apparently nocturnal. Hypanthium obconic to cylindrical. Calyx limb lobed, lobes 5, without calycophylls; corolla salverform with the tube slender and prolonged, white becoming yellow when old, internally glabrous to papillose or pilosulous at least in throat, lobes 5, in bud imbricated and the whole group of lobes often tilted to one side. Stamens 5, inserted in the mouth of the corolla; anthers dorsifixed near base, narrowly oblong, exserted, with the connective usually prolonged on one or both ends, often pilosulous to hispidulous or pilose. Ovary 2-locular or incompletely 1-locular; ovules numerous in each locule, on axile placentas. Fruit baccate, subglobose to ellipsoid or ovoid, leathery to carnose, orange to brown, endocarp coriaceous to woody. Seeds angled to subglobose, smooth, 5–15 mm long, embedded in pulp or perhaps arillate. Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Bolivia, Brazil; ca. 20 species, 5 in Venezuela, 4 of these in the flora area.

Posoqueria 703

Posoqueria is similar to Tocoyena, and these genera are often confused. Tocoyena can be separated by its corolla lobes that are convolute in bud. Dried specimens of Tocoyena can often be recognized also by a blackened color, while those of Posoqueria usually remain green. Posoqueria is also similar to Sphinctanthus; their separation is discussed under Sphinctanthus. This neotropical genus has long been included in the tribe Gardenieae (e.g., Steyermark 1974; Robbrecht 1993), but more recent work (e.g., Persson 1996) suggests that it does not belong to this tribe and that in general its affinities in the family are not clear. The taxonomy of Posoqueria has been poorly understood. The treatment here and in the Flora Mesoamericana (Taylor, in preparation) differs significantly in several species delimitations from that of Steyermark (1967, 322–333). The flowers of at least some species of Posoqueria show an unusual pollination mechanism in the Rubiaceae: the filaments of recently open flowers are strongly reflexed under tension until they are disturbed by the touch of an insect. Then they spring forward rapidly and with force, and push the dehiscing anthers onto the insect. In some species of Posoqueria the stamens are inserted symmetrically into the corolla tube but the anthers are grouped on one or two sides of the flowers, giving the flowers a zygomorphic aspect. Key to the Species of Posoqueria 1. 1. 2(1). 2. 3(2).

3.

Lower surface of leaves densely pilosulous to velutinous-hirtellous .................................................................................................. P. williamsii Lower surface of leaves glabrous .............................................................. 2 Corolla tubes 8.5–12 cm long; calyx limb 1.2–2 cm long ............... P. latifolia Corolla tubes 18–29.5 cm long; calyx limb 2–4 mm long, sometimes elongating to 10 mm long in fruit ................................................................ 3 Corolla with the tube 28–29.5 cm long, the lobes 3.8–4.5 cm long; inflorescences (not counting the corollas and fruits) ca. 4 × 3.5 cm; fruit 4.5– 56 cm diameter ........................................................................ P. longiflora Corolla with the tube 18–19 cm long, the lobes 1.7–3 cm long; inflorescences (not counting the corollas and fruits) 5–7.5 × 4–8 cm; fruit ca. 6 cm diameter ....................................................................... P. panamensis

Posoqueria latifolia (Rudge) Roem. & Schult., Syst. Veg. 5: 227. 1819. —Solena latifolia Rudge, Pl. Guian. 1: 26, pl. 40. 1806. Shrub or tree to 20 m tall; leaves 5–24 × 3–12 cm; stipules 5–10 mm long; inflorescences (not including the corollas and fruits) 3.5–6 × 3.5–5 cm; calyx limb 1.5–2 mm long; corolla tube 8.5–12 cm long, lobes 1.2–2 cm long; fruit 3.5–4 cm diameter. Riparian forests, rain forests, montane forests, forested talus slopes, 100–1100 m; Mexico, Central America, Antilles, Venezuela, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 2 subspecies, both in the flora area.

Key to the Subspecies of P. latifolia 1. Leaf blades elliptic-oblong to lance-oblong, 5–16 × 3–5 cm wide, acuminate at apex ............................. subsp. gracilis 1. Leaf blades ovate to elliptic, elliptic-oblong, or ovate-oblong, 7.5–24 × 5–12 cm, obtuse to very shortly acute at apex .................................... subsp. latifolia P.

latifolia subsp. gracilis (Rudge) Steyerm., Mem. New York Bot. Gard. 17(1): 327. 1967. —Solena gracilis Rudge, Pl. Guian. 27, t. 41. 1806. —Posoqueria gracilis (Rudge) Roem. & Schult., Syst. Veg. 5: 227. 1819.

704

R UBIACEAE

Bolívar (Canaima), Amazonas (basin of Río Guainía, Río Negro and Río Orinoco). Guyana, Suriname, French Guiana, Amazonian Brazil. The subspecies status of these plants probably deserves re-evaluation: although flower size does not appear to differ between this and subsp. latifolia, the leaf, inflorescence, and fruit morphology appear to be consistently different and these may be better treated as species. P. latifolia subsp. latifolia Delta Amacuro (Curiapo), Bolívar (Río Caura, Río Cuyuní), Amazonas (Río Casiquiare, Río Guainía, Río Mawarinuma, Río Orinoco). Zulia; Mexico, Central America, Antilles, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Bolivia, Brazil. ◆Fig. 558. Posoqueria longiflora Aubl., Hist. Pl. Guiane 134, t. 51. 1775. —Aguacatillo, Cafecillo, Jasmín de estrella, Manzanillo, Toronjillo. Shrub or small tree to 10 m tall; leaves 7.5–20 × 3–10 cm; stipules 4–10 mm long; inflorescences (not including corollas and fruits) ca. 4 × 3.5 cm; calyx limb 2–4 mm long; corolla tube 28–29.5 cm long, lobes 3.8– 4.5 cm long; fruit 4.5–5 × 3–4.5 cm, with calyx limb to 10 mm long. Periodically flooded riparian forests, borders of Mauritia palm swamps, 50–400 m; Delta Amacuro (Caño Güiniquina, Río Orocoima), Bolívar (Río Caroní, Río Caura, Río Paragua, Río Suapure), Amazonas (Río Casiquiare, Río Orinoco, Río Siapa). Apure; Guyana, Suriname, French Guiana, Peru, Amazonian Brazil. ◆Fig. 557. Steyermark distinguished Posoqueria longiflora from other species of Posoqueria, in particular P. panamensis, by its “dorsal base of anther conspicuously setose-hispid with elongate hairs 0.7–1 mm long which are conspicuously longer than the other indument of the anther,” versus “with indument similar to rest of anther” in the other species. However, this character is not informative for separating species: the anthers of P. panamensis and P. longiflora appear to be similarly pubescent, with both species having anther pubescence that varies widely in length and density and occasional specimens of both species having anthers similar to

those described by Steyermark for P. panamensis. Flower size does seem to be consistently different, and consequently as treated here both P. panamensis and P. longiflora comprise a much smaller range of flower sizes than in Steyermark’s view. Posoqueria panamensis (Walp. & Duchass.) Walp., Ann. Bot. Syst. 2: 797. 1852. —Stannia panamensis Walp. & Duchass., Linnaea 23: 744. 1850. —Café montañero, Maspara, Mazano, Paraparo. Posoqueria panamensis subsp. grandiflora (H. Karst.) Steyerm., Mem. New York Bot. Gard. 17(1): 325. 1967. —Stannia grandiflora H. Karst., Fl. Columb. 1: 31, t. 16. 1859, not Posoqueria grandiflora Standl. 1928. Shrub or tree to 10 m tall; leaves 10.5– 20.5 × 4.5–9.5 cm; stipules 8–15 mm long; inflorescences (not including corollas and fruit) 5–7.5 × 4–8 cm; calyx limb ca. 3 mm long; corolla tube 18–19 cm long, lobes 1.7–3 cm long; fruit ca. 6 cm diameter. Riparian and evergreen lowland to montane forests, 100– 1500 m; Delta Amacuro (Caño Arature), Bolívar (Río Caura, basin of Río Paragua), Amazonas (basin of Río Guainía and Río Orinoco). Apure, Portuguesa; southern Central America, Colombia, Ecuador, Peru, Brazil, Bolivia. Steyermark recognized two subspecies of Posoqueria panamensis: subsp. panamensis, supposed to be found from Mexico to Colombia and to have flowers relatively shorter on average, and subsp. grandiflora (H. Karst.) Steyerm., found in South America and with its flowers relatively larger. Steyermark noted that the flower sizes and geographic ranges of these subspecies overlap, and these subspecies do not seem taxonomically meaningful and are not recognized here. Steyermark classified the plants of the flora area in subsp. grandiflora, whose type is from Antioquia in northwestern Colombia. Posoqueria williamsii Steyerm., Mem. New York Bot. Gard. 17(1): 331, fig. 37. 1967. —Hoja de danto. Tree to 6 m tall; leaves 9–39 × 9–20 cm; stipules 5–8 cm long; inflorescences (not including corollas and fruits) 2–3 × 3–3.5 cm; calyx limb 5–6 mm long; corolla tube 22–24 cm long, lobes 2–2.3 cm long; immature fruit to 5.5 × 3 cm. Evergreen lowland and ripar-

Posoqueria 705

ian forests, 100–200 m; Amazonas (basin of Río Casiquiare, Río Negro, and Río Orinoco). Endemic. ◆Fig. 556. Posoqueria williamsii is similar to P. maxima Standl. of Central America through central Colombia, and these may not be distinct but collections are currently too few to

evaluate this question. H. Clark et al. (2000) reported P. maxima from the area of San Carlos de Río Negro in the flora area. However, the source of this report is unknown, as is the distinction they draw between P. maxima and P. williamsii, which they also report from that region.

Fig. 556. Posoqueria williamsii

706

R UBIACEAE

Fig. 557. Posoqueria longiflora

Fig. 558. Posoqueria latifolia subsp. latifolia

67. PSYCHOTRIA L., Syst. Nat. ed. 10, 929. 1759, nom. cons. by Charlotte M. Taylor and Julian A. Steyermark Terrestrial shrubs, small trees, or trailing to erect herbs, unarmed. Leaves opposite, subsessile to petiolate, the venation not lineolate, sometimes with foveolate domatia in the abaxial axils of the secondary veins; stipules caducous or persistent, generally valvate to imbricate in bud, of various forms: interpetiolar and triangular to bilobed, calyptrate (i.e., united into a conical cap), interpetiolar and emarginate

Psychotria 707

to bilobed with the intrapetiolar portion reduced (i.e., laminar), or united around the stem into a continuous sheath that is truncate to bilobed on each interpetiolar side. Inflorescences terminal or infrequently pseudoaxillary (i.e., present in only one axil at each node) or truly axillary (i.e., present in both axils at each node), several- to multiflowered, of various forms including capitate, subcapitate, congestedcymose, corymbiform, or paniculate, sessile to pedunculate, ebracteate to bracteate, the bracts often well developed and sometimes involucral, green to brightly colored. Flowers sessile or pedicellate, usually distylous, usually small. Hypanthium turbinate to subglobose. Calyx limb reduced to well developed, lobes (4)5, without calycophylls; corolla tubular to funnelform or salverform, white to yellow or sometimes flushed with pink or purple, internally glabrous to variously pubescent, tube straight to occasionally curved and/or gibbous at base, lobes (4)5, valvate in bud. Stamens (4)5, inserted near or above middle of corolla tube; anthers narrowly oblong, dorsifixed near middle, included in long-styled flowers, usually partially or wholly exserted in short-styled flowers. Ovary 2(–5)-locular; ovules solitary in each locule, basal. Fruit subglobose to ovoid, drupaceous, carnose or spongy, red, blue, purple-black, white, yellow, or orange; pyrenes 2(–5), bony, hemispherical, subglobose, or when > 2 then triangular, dorsally longitudinally ridged or sometimes smooth. Seeds ellipsoid to subglobose, medium-sized for the family, smooth. Southeastern U.S.A., Mexico, Central America, the Antilles, Colombia, Venezuela, Trinidad-Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina, Uruguay, Africa, Madagascar, Asia, Pacific Islands; in its broad sense ca. 1400 species, perhaps 800 in the New World, ca. 162 in Venezuela, 120 of these in the flora area. Psychotria is widespread in both the New World and Old World tropics and subtropics. It is similar to and often confused with Ixora, Palicourea, Coussarea, and Rudgea. Its separation from these is discussed under each of those other genera. Psychotria has long been circumscribed broadly, to include essentially all the species of its tribe with small white insect-pollinated flowers that lack other distinctive features. Steyermark (1972; 1974) notably expanded Psychotria’s traditional circumscription by including in it Cephaelis Sw. More recent studies have concluded that this group, both in the Neotropics (Taylor 1996) and worldwide (Nepokroeff et al. 1999; Andersson 2002), comprises several distinct lineages that are better separated, though Steyermark’s synonymization of Cephaelis is supported by all of these studies. Two of these lineages, Notopleura (Benth. & Hook. f.) Bremek. and Ronabea Aubl., are treated separately in this flora. Andersson has also proposed separating Carapichea Aubl. (2002) and Margaritopsis Griseb. (2001) but the overall limits of these groups are not yet clear and their species are here included for now in Psychotria subgenus Heteropsychotria. Subgenus Heteropsychotria will eventually be combined with Palicourea (Taylor 1996; see the discussion under Palicourea above). The two subgenera are keyed separately here, and all species of Psychotria are presented alphabetically after the keys. Key to the Subgenera of Psychotria 1.

Stipules persistent with the leaves, when fallen leaving no persistent trichomes or these clear and ≤ 0.5 mm long; fruit red, blue, purple-black, yellow, orange, or white at maturity; leaves without foveolate domatia in the abaxial vein axils (if the plant otherwise keys out here and these structures are present, go to Rudgea); plants usually drying green or occasionally reddish brown ............. subg. Heteropsychotria, page 708

708

1.

R UBIACEAE

Stipules caducous or sometimes persistent on the distalmost 1 or 2 nodes, when fallen leaving a ring of persistent reddish brown trichomes 0.5– 1 mm long; fruit red at maturity; leaves with foveolate domatia in the abaxial vein axils; plants drying gray-green or reddish brown ............. ...................................................................... subg. Psychotria, page 719

Psychotria subg. Heteropsychotria Steyerm., Mem. New York Bot. Gard. 23: 484. 1972. Carapichea Aubl., Hist. Pl. Guiane 167, pl. 64. 1775. Nonatelia Aubl., Hist. Pl. Guiane 182, pl. 70–75. 1775. Tapogomea Aubl., Hist. Pl. Guiane 357, pl. 60, 63. 1775. Cephaelis Sw., Prodr. 45. 1788. Grumilea Gaertn., Fruct. Sem. 1: 138, pl. 1, fig. 7. 1788. Callicocca Schreb., Gen. Pl. 1: 126. 1789. Ipecacuanha Arruda, Diss. Pl. Brazil 44. 1810. Uragoga Baill., Adansonia 12: 323. 1879. Chytropsia Bremek., Meded. Bot. Mus. Herb. Rijks Univ. Utrecht 11: 289. 1934. Naletonia Bremek., Meded. Bot. Mus. Herb. Rijks Univ. Utrecht 11: 284. 1934. Mexico, Central America, the Antilles, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Argentina, Paraguay; perhaps 600 species, at least 133 in Venezuela, 105 in the flora area. Steyermark (1972) incorrectly considered subgenus Heteropsychotria to be pantropical, but it is restricted to the Neotropics (Taylor 1996). Subgenus Heteropsychotria is very similar to Palicourea. For comments regarding their separation see under Palicourea. The corollas of many species of subgenus Heteropsychotria range from pure white to cream and sometimes pale yellow. The inflorescences of the species of this group are generally green, pale green, white, or flushed with pink or pink-purple, but the infructescences of most species become purple or violet as the fruits mature no matter what color the inflorescences are. Steyermark (1974) considered the number and arrangement of the inflorescence bracts taxonomically informative, in particular the degree of fusion among them, whether each bract subtends one or more than one flower, and how many bracts there are per individual flower. The significance of these characters and their variation at a population level have not been carefully studied. Measurements given here for fruits are those for the dried fruits found on specimens. In many species of Psychotria, the fruits enlarge markedly in size (i.e., to 2–4 times the immature or dry diameters) on the day they become mature, apparently largely by water uptake, and then they are rapidly removed from the infructescence. Steyermark (1974) in particular in his descriptions occasionally mixes dry fruit size with the living mature fruit size, sometimes indicating the respective states and sometimes not. Key to the Species of Psychotria subg. Heteropsychotria and Similar Taxa 1. 1.

Inflorescences consistently axillary (i.e., produced in both axils at each node) ....................................................................................................... 2 Inflorescence terminal and/or pseudoaxillary (i.e., produced in only one axil at each node) ................................................................................... 6

Psychotria 709

2(1). 2. 3(2). 3.

4(3). 4. 5(4). 5. 6(1). 6. 7(6). 7. 8(7). 8. 9(6). 9. 10(9). 10. 11(10). 11. 12(10). 12. 13(12). 13. 14(9). 14. 15(14). 15. 16(15).

Stipules bilobed on each interpetiolar side, the lobes linear, 1–2.5 mm long ........................................................................................... P. vellosiana Stipules triangular or with only 1 lobe on each interpetiolar side, lobes to 4 mm long ............................................................................................... 3 Stipules triangular on each interpetiolar side; inflorescences sessile or with peduncles to 0.5 cm long ......................................... Ronabea latifolia Stipules with a narrowly triangular lobe 1–4 mm long on each interpetiolar side; inflorescences pedunculate, peduncles 0.5–5 cm long ......................................................................................................... 4 Leaves broadly elliptic to suborbicular, rounded to cordulate at base .................................................................................................... P. durifolia Leaves narrowly elliptic to oblanceolate or spathulate, acute to obtuse at base ......................................................................................................... 5 Calyx limb 2–3 mm long, lobed for ca. 3/4 its length or more ........ P. carrenoi Calyx limb 1–1.2 mm long, lobed for ca. 1/2 its length .............. P. phelpsiana Creeping herbs, regularly rooting at nodes, including at nodes with leaves ...................................................................................................... 7 Erect herbs, subshrubs, shrubs, or trees, not rooting at nodes along stem (except anomalously when plant is damaged) ...................................... 9 Leaves cordate or cordulate at base; fruit red ........................... 37. Geophila Leaves acute to rounded or cordulate at base; fruit blue or black .......... 8 Leaves 2–8.5 cm wide, rounded to cordulate at base, rounded to obtuse at apex, petioles 4–32 mm long .................................................. P. ulviformis Leaves 1–3 cm wide, truncate to acute at base, acute at apex, petioles 3– 10 mm long ................................................................................ P. variegata Inflorescences consistently pseudoaxillary (i.e., produced in only one axil at each node), produced at several nodes along each stem ................ 10 Inflorescences terminal, occasionally some of them displaced to pseudoaxillary but this arrangement not regular ......................................... 14 Inflorescences sessile, capitate, 0.7–1.5 × 0.8–2.5 cm, with bracts 6– 10 mm long ........................................................................................... 11 Inflorescences with peduncles 0.5–5 cm long, capitate, subcapitate, or congested-cymose, 0.5–2 × 0.5–2 cm, with bracts 1–2.5 mm long ..... 12 Leaves abaxially glabrous to strigillose, with the tertiary veins plane ................................................................................................. P. aubletiana Leaves abaxially hirtellous, with the tertiary veins markedly raised ......................................................................................................... P. celiae Leaves broadly elliptic to suborbicular, rounded to cordulate at base .................................................................................................... P. durifolia Leaves narrowly elliptic to oblanceolate, acute to obtuse at base ......... 13 Calyx limb 2–3 mm long, lobed for 3/4 or more its length .............. P. carrenoi Calyx limb 1–1.2 mm long, lobed for ca. 1/2 its length .............. P. phelpsiana Leaves with most or all of the secondary veins extending unbranched to unite with the margins, these cartilaginous to markedly thickened.... 15 Leaves with the secondary veins all reticulating before reaching the margins, these thin to thinly cartilaginous ............................................... 25 Leaves cordulate to cordate at base, with petioles 0–1 mm long .......... 16 Leaves acute to obtuse at base, with petioles 1–10 mm long ................ 18 Peduncle 7–10 cm long and leaves 10–16 × 5.5–12 cm ............ P. sipapoensis

710

R UBIACEAE

16. Peduncles 0.8–8 cm long, leaves 2.5–10.5 × 1.5–7 cm ............................ 17 17(16). Calyx limb 0.3–0.5 mm long; inflorescences green or white perhaps flushed with pink; corolla tube 2–3 mm long .................. P. cardiomorpha 17. Calyx limb 1–1.5 mm long; inflorescence vivid pink, orange, or red; corolla tube 6–6.5 mm long ......... Palicourea foldatsii, Palicourea lancigera 18(15). Petioles ca. 1 mm long; leaves 3–4 × 0.5–1.5 cm .......................... P. vareschii 18. Petioles 3–20 mm long; leaves 3–21 × 1–9 cm ........................................ 19 19(18). Inflorescences cymose with axes developed to tertiary or higher orders, the flowers borne in branched cymes .................................................. 20 19. Inflorescences capitate or with secondary axes developed, the flowers borne in capitula .................................................................................. 21 20(19). Leaves chartaceous with the margins markedly thickened; stipule lobes 3.5–17 mm long; fruit 3.5–4 × 4–5 mm, becoming orange then black ................................................................................................ P. longicuspis 20. Leaves papyraceous with the margins rather thin; stipule lobes 0.5– 1 mm long; fruit ca. 2 × 3 mm, becoming white ................ P. subundulata 21(19). Inflorescence corymbiform, with secondary axes developed and equal to or longer than the primary axis; leaves lanceolate; fruit orange ............................................................................................ P. schomburgkii 21. Inflorescences capitate, subcapitate, spiciform, or paniculate and pyramidal, the secondary axes not developed or these developed but shorter than the primary axis; leaves elliptic to narrowly so; fruit white, blue, or purple ............................................................................................... 22 22(21). Inflorescences with the bracts of each capitulum all about equal in length ............................................................................................. P. phaneroloma 22. Inflorescences with each capitulum subtended by a single bract that is 2 or more times as long as the remaining bracts ............................... 23 23(22). Inflorescences spiciform to pyramidal, with 9–17 capitula, these each ca. 3 × 3 mm ................................................................................ P. polycephala 23. Inflorescences capitate, subcapitate, or paniculate, with 1–5 capitula, these each 4–10 × 4–10 mm ................................................................ 24 24(23). Inflorescences subcapitate to paniculate, with the head or branched portion 1–3 × 1.5–2.5 cm; leaves 2–7 cm wide .................................. P. egensis 24. Inflorescences capitate to subcapitate or congested-cymose, with the head or branched portion 0.5–1 × 1–2 cm; leaves 1–2.5 cm wide .... .............................................................................................. P. lourteigiana 25(14). Leaves sessile to subsessile and cordate to cordulate at base ............... 26 25. Leaves sessile to petiolate and obtuse to acute at base ......................... 29 26(25). Stipules on all nodes with the lobes arising from the top of the sheath; floral bracts 0.5–1.5 mm long ............................................. P. imthurniana 26. Stipules on older nodes with the lobes inserted below the top of the sheath; floral bracts 3–11 mm long ..................................................... 27 27(26). Corolla tube 3–4 mm long ......................................................... P. amplectans 27. Corolla tube 6–8.5 mm long ..................................................................... 28 28(27). Leaves chartaceous to subcoriaceous, with a clearly developed continuous submarginal vein ........................................................................... P. blakei 28. Leaves papyraceous, without a continuous submarginal vein .... P. lupulina 29(25). Stipules significantly lacerate, divided for 1/4 or more their length into at least 4 segments on each interpetiolar side .................. P. sphaerocephala

Psychotria 711

29. 30(29).

30.

31(30). 31. 32(31). 32. 33(32). 33. 34(33). 34. 35(32). 35. 36(35). 36. 37(36).

37.

38(30). 38. 39(38). 39. 40(39). 40.

41(40). 41.

Stipules truncate to triangular, ovate, emarginate, or shortly to deeply bilobed .................................................................................................. 30 Stipules interpetiolar and sometimes shortly intrapetiolar, bilobed for 1/4 or more their lengths, and 3–22 mm long, or acute and > 10 mm long .............................................................................................................. 31 Stipules interpetiolar to united around the stem, truncate to rounded, obtuse, acute, emarginate, or shortly to deeply bilobed, 1–30 mm long, if unlobed then < 10 mm long ................................................................. 38 Plants pilose to hirsute ....................................................................... P. pilosa Plants glabrous to strigillose or puberulous ........................................... 32 Inflorescence bracts generally reduced, the floral bracts to 1.5 mm long .............................................................................................................. 33 Inflorescence bracts well developed, the floral bracts 2–20 mm long [P. capitata complex] ............................................................................ 35 Stipules 11–20 mm long ............................................................ P. microbotrys Stipules 3–8 mm long .............................................................................. 34 Inflorescence with branched portion 8–28 cm long; calyx limb ca. 0.5 mm long ................................................................................ P. bostrychothyrsus Inflorescence with branched portion 5–9 cm long; calyx limb ca. 0.2 mm long .......................................................................................... P. paniculata Stipules 10–20 × 10–12 mm, ovate in outline, acute or bilobed to 1/4 their length; corolla tube 8–11 mm long ........................................... P. stipulosa Stipules 3–19 × 2–6 mm, lanceolate to lingulate in outline, bilobed for 1/2 to nearly completely their length; corolla tube 2.2–9 mm long ......... 36 Corolla tube 2.2–3.5 mm long; inflorescence with branched portion 1– 1.8 × 1–2 cm; floral bracts 2.5–3 mm long ............................. P. guanchezii Corolla tube 3.5–9 mm long; inflorescence with branched portion 1.5–8 × 2–7.5 cm; floral bracts 1.5–15 mm long .............................................. 37 Inflorescence pyramidal, with the basalmost secondary axes less well developed than the primary axis; inflorescence axes, bracts, and corollas generally green to white or yellowish white or sometimes flushed with pink; corolla tubes 4.5–7 mm long ............................................. P. capitata Inflorescence congested-cymose to corymbiform, with the basalmost secondary axes usually about as well developed as the primary axis; inflorescence axes, bracts, and corollas variously greenish white, white, pink, purple, or orange-brown; corolla tube 7–9 mm long ................ ............................................................................................ P. maguireorum Inflorescence capitate or subcapitate, with no axes evident .................. 39 Inflorescence with axes developed at least shortly, at least the primary axis ....................................................................................................... 77 Inflorescence sessile or subsessile, with peduncle to 3 mm long ........... 40 Inflorescence with peduncle 4–70 mm long ............................................ 55 Stipules with the interpetiolar portion triangular to ovate, 3–10 mm long, entire to shortly 2-apiculate; fruit 10–15 × 8–11 mm ........ P. podocephala Stipules with the interpetiolar portion truncate to broadly rounded and to 4.5 mm long, or shortly to deeply bilobed and 1–20 mm long; fruit 3– 8 × 3–6.5 mm ........................................................................................ 41 Inflorescence bracts none or relatively small, to 2 mm long .................. 42 Inflorescence bracts well developed, the outermost 3–20 mm long ....... 44

712

R UBIACEAE

42(41). Stipules 2–4 mm long, truncate; calyx limb 2–3 mm long; corolla tube 3.5–5 mm long ......................................................................... P. deinocalyx 42. Stipules 0.5–1.5 mm long, bilobed or with 2 caducous aristas; calyx limb 0.5–1.5 mm long; corolla tube ca. 2 mm long (unknown in P. nana) .............................................................................................................. 43 43(42). Leaves 1.8–7 cm wide; inflorescence subcapitate, especially with the fruits often loosely grouped; calyx limb 0.5–0.8 mm long .............. P. nana 43. Leaves 2–3 cm wide; inflorescence and infructescence densely capitate; calyx limb 1–1.5 mm long ................................................... P. pallidinervia 44(41). Inflorescence bracts not involucral, not enclosing flower groups or the inflorescence but loosely subtending them ............................................ 45 44. At least some inflorescence bracts involucral, with 1–3 pairs enclosing the entire inflorescence and/or individual flower groups .................. 46 45(44). Petioles 1–5 mm long; calyx limb 3.5–8 mm long ...................... P. iodotricha 45. Petioles 5–12 mm long; calyx limb 0.5–2 mm long .................... P. medusula 46(44). External inflorescence bracts 4–7 mm long (at least some of them < 7 mm long) ...................................................................................................... 47 46. External inflorescence bracts 7–20 mm long (at least some of them > 7 mm long) ...................................................................................................... 51 47(46). Stipules truncate, sometimes with 1 caducous arista on each interpetiolar side; fruit red ................................................................. P. kappleri 47. Stipules bilobed on each interpetiolar side; fruit purple, blue, or black .............................................................................................................. 48 48(47). Plants slender and small; leaves 1–5.5 × 0.3–2 cm; inflorescence 0.3–0.5 × 0.5 cm .................................................................................... P. vichadensis 48. Plants small to large, generally robust; leaves 2–13 × 0.5–5 cm; inflorescence 0.5–1.5 × 0.5–2 cm ..................................................................... 49 49(48). Inflorescence bracts markedly unequal in size, the most external ones > 2 times as long as the floral bracts ........................... P. hoffmannseggiana 49. Inflorescence bracts subequal, the most external ones about as long as the floral bracts .................................................................................... 50 50(49). Leaves 2.5–5 cm wide; stipule sheath 1–2 mm long; inflorescence 0.8– 1.5 × 1–2 cm; calyx limb 0.3–0.5 mm long ............................ P. casiquiaria 50. Leaves 1–3.5 cm wide; stipule sheath ca. 0.5 mm long; inflorescence 0.5– 0.8 × 0.5–0.8 cm; calyx limb ca. 1 mm long .............................. P. spadicea 51(46). Stipules truncate, sometimes with 2 short glandular projections or groups of projections on each interpetiolar side ........................... P. apoda 51. Stipules bilobed on each interpetiolar side ............................................. 52 52(51). Inflorescence lanceoloid, the external bracts 2–10 mm long, tightly appressed; calyx limb 1–1.5 mm long ............................................ P. oblonga 52. Inflorescence subglobose to ovoid or oblate, the external bracts 10–30 mm long, loosely ascending to spreading; calyx limb 2.5–11 mm long ..... 53 53(52). Calyx limb 10–11 mm long .......................................................... P. fanshawei 53. Calyx limb 2.5–4 mm long ....................................................................... 54 54(53). Inflorescence subglobose to oblate, the external bracts lanceolate to ovate or elliptic; stipule sheath 2–3 mm long; calyx limb 3.5–4 mm long ............................................................................................... P. bolivarensis 54. Inflorescence ovoid, the external bracts pandurate; stipule sheath ca. 0.5 mm long; calyx limb 2.5–3 mm long ................................. P. prunifolia

Psychotria 713

55(39). Stipules truncate to triangular or ovate, 3–10 mm long, shortly biapiculate or with 1 or 2 caducous aristas on each side ................... 56 55. Stipules bilobed for 1/3 or more their length, 1–20 mm long (the sinus between the lobes may be quite broad and shallow in P. nana) ............ 58 56(55). Peduncle 3–6 cm long; inflorescence bracts well developed, the external ones 15–45 mm long .............................................................. P. pacimonica 56. Peduncle 0.4–0.9 cm long; inflorescence bracts none or reduced, to 1 mm long ....................................................................................................... 57 57(56). Stipules truncate; corolla tube ca. 3.5 mm long; fruit 17–25 × 15–17 mm ................................................................................................ P. paupertina 57. Stipules triangular to ovate; corolla tube ca. 5 mm long; fruit 10–15 × 8– 11 mm ................................................................................... P. podocephala 58(55). Inflorescence bracts none or relatively reduced, to 2 mm long .............. 59 58. Inflorescence bracts well developed, the outermost bracts 3–70 mm long .............................................................................................................. 60 59(58). Peduncle 1–1.8 cm long; corolla tube 12–15 mm long; fruit blue ... P. concinna 59. Peduncle 0.4–0.5 cm long; corolla tube in mature bud ca. 2 mm long (length at anthesis unknown); fruit red ......................................... P. nana 60(58). External inflorescence bracts broadly rounded to truncate ................... 61 60. External inflorescence bracts obtusely angled to acute ......................... 64 61(60). Peduncle 2.8–3.3 cm long; external bracts subtending individual flowers or axes, not involucral .......................................................... P. adenophora 61. Peduncle 0.5–4 cm long; external bracts involucral, enclosing the inflorescence ................................................................................................ 62 62(61). Peduncle 1–4 cm long; calyx limb ca. 1.5 mm long; corolla tube 7.5–8 mm long .................................................................................... P. hemicephaelis 62. Peduncle 0.5–1.7 cm long; calyx limb 0.8–1 mm long; corolla tube 5– 6.5 mm long .......................................................................................... 63 63(62). Inflorescence nodding; peduncle 0.4–1 cm long .......................... P. platypoda 63. Inflorescence erect; peduncle 0.9–1.6 cm long ............................ P. turbinella 64(60). Inflorescence with the most external bracts 6–20 mm long, these subtending flowers and flower groups but not involucral ....................... 65 64. Inflorescence with the external bracts 2.5–70 mm long, involucral, enclosing the head and flower groups .................................................... 66 65(64). Stipule lobes 3–10 mm long; corolla tube 10–14 mm long; plants of lowlands, 100–200 m ..................................................................... P. medusula 65. Stipule lobes 1–3.5 mm long; corolla tube 18–21 mm long; plants of montane areas, 2300–2600 m ............................................................... P. oblita 66(64). Inflorescence with external bracts 2.5–12 mm long, narrowly elliptic to ligulate or narrowly oblong, green to pale green or white ................. 67 66. Inflorescence with external bracts 9–70 mm long with at least some > 12 mm long, elliptic to broadly so or ovate, white, orange, red, pink, or purple ............................................................................................... 71 67(66). Inflorescence with outermost or basalmost bracts > 2 times as long as the floral bracts, the bracts situated between them not gradually grading in length ..................................................................... P. hoffmannseggiana 67. Inflorescence with outermost or basalmost bracts < 2 times as long as the floral bracts, with the bracts situated between them also intermediate in length ............................................................................................... 68

714

R UBIACEAE

68(67). Peduncle ca. 5 cm long .................................................................... P. cerronis 68. Peduncle 0.2–3.5 cm long ........................................................................ 69 69(68). Peduncle 0.7–3.5 cm long; calyx limb ca. 0.2 mm long; plants of montane areas, 1100–2200 m ............................................................ P. campylopoda 69. Peduncle 0.2–2.2 cm long; calyx limb 0.3–0.5 mm long; plants of lowland areas, 50–800 m ................................................................................... 70 70(69). Stipule lobes 1.5–2 mm long; corolla lobes 1.5–2 mm long; fruit ca. 3 mm diameter ................................................................................. P. casiquiaria 70. Stipule lobes 2–6 mm long; corolla lobes ca. 1 mm long; fruit 4–5 mm diameter ................................................................................... P. gracilenta 71(66). Calyx limb 2.5–4 mm long ....................................................................... 72 71. Calyx limb 0.8–2 mm long ....................................................................... 73 72(71). Stems, leaves, stipules, and inflorescences hirtellous or pilosulous, sometimes becoming glabrescent with age; peduncle 1–1.5 cm long ............................................................................................... P. bolivarensis 72. Stems, leaves, stipules, and inflorescences glabrous; peduncle 1–3.5 cm long .................................................................................... P. humboldtiana 73(71). Leaves sessile; stems and upper surface of leaves hirsute ...... P. parimensis 73. Leaves with petioles 2–50 mm long; stems and upper surface of leaves hirsute to strigose, puberulous, pilosulous, strigillose, hirtellous, or glabrous ................................................................................................ 74 74(73). Floral bracts obtuse to rounded, pilosulous uniformly and similarly on both surfaces; corolla tube 3.5–4.5 mm long .................... P. bracteocardia 74. Floral bracts acute, glabrous to strigillose, strigose, pilosulous, hirtellous, or pilose but the pubescence not uniform on each surface and not the same on both surfaces; corolla tube 8.5–14.5 mm long ...................... 75 75(74). Corolla yellow, with tube 13–14.5 mm long; inflorescence bracts orange to red, the largest 30–70 mm long .......................................... P. poeppigiana 75. Corolla white to purple, with tube 8.5–12 mm long; inflorescence bracts white to pink or purple, the largest 15–35 mm long .......................... 76 76(75). Plants glabrous; reported from Bolívar ......................................... P. colorata 76. Plants glabrous to puberulous, strigillose, strigose, pilosulous, or glabrescent; reported from Amazonas ........................................................ P. rosea 77(38). Inflorescences spiciform, with the flowers borne directly from the primary axis or in congested cymules that are borne directly from this ................................................................................................... P. spiciflora 77. Inflorescences congested-cymose to corymbiform or paniculate, with the flowers borne on secondary or higher-order axes ............................... 78 78(77). Inflorescence with the bracts enclosing flowers, cymules, and/or capitula 3–25 mm long, at least some of them > 4 mm long ............................ 79 78. Inflorescence with the bracts enclosing or subtending flowers, cymules, and/or capitula none or relatively reduced, to 3 mm long ............... 103 79(78). Stipules ovate, 3–13 mm long; petioles 20–75 mm long; peduncle 5.5– 11.5 cm long ......................................................................................... 80 79. Stipules truncate to emarginate or broadly to deeply bilobed on each interpetiolar side, 1–8 mm long; petioles 1–22 mm long; peduncle 0.4– 9.5 cm long ........................................................................................... 82 80(79). Stipules 10–13 mm long; corolla tube 15–16 mm long ......... P. maturacensis 80. Stipules 3–11 mm long; corolla tube 5–6 mm long (length at anthesis un-

Psychotria 715

81(80). 81. 82(79). 82. 83(82). 83. 84(83). 84. 85(82). 85. 86(85). 86. 87(86). 87.

88(87).

88.

89(85). 89. 90(89). 90. 91(90). 91.

92(91). 92.

93(89). 93. 94(93). 94.

known in P. franquevilleana) .............................................................. 81 Calyx limb 0.7–2 mm long ................................................. P. franquevilleana Calyx limb 3–3.5 mm long ........................................................... P. vasivensis Calyx limb 2.5–7.5 mm long .................................................................... 83 Calyx limb 0.3–1.5 mm long .................................................................... 85 Calyx limb 6–7.5 mm long ........................................................... P. tepuiensis Calyx limb 2.5–4 mm long ....................................................................... 84 Inflorescence bracts 9–20 mm long, pink to magenta; corolla tube 18– 21 mm long ........................................................................ P. humboldtiana Inflorescence bracts 3.5–6 mm long, green to perhaps white; corolla tube 7–8 mm long .............................................................................. P. transiens Petioles 10–20 mm long, at least some of them > 10 mm long .............. 86 Petioles 1–10 mm long ............................................................................. 89 Inflorescence bracts 12–23 mm long ..................................... P. crocochlamys Inflorescence bracts 4.5–11 mm long ...................................................... 87 Stipule lobes narrowly triangular, acute; corolla pale blue; plants of lowland forests, 100–200 m ................................................... P. tapirapecoana Stipule lobes none, ligulate to narrowly triangular, rounded to obtuse or acute; corolla white; plants of premontane and montane forests, 1000– 1800 m .................................................................................................. 88 Floral bracts 1–few per cymule, fewer than 1 per flower; corolla with tube ca. 8 mm long, lobes ca. 5 mm long; stipules lobes triangular and acute ...................................................................................................... P. brazaoi Floral bracts 1 or more per flower; corolla with tube 6.5–7 mm long, lobes ca. 3 mm long; stipule lobes ligulate and broadly rounded to obtuse .................................................................................................. P. wurdackii Stipule lobes 0.2–2 mm long, equal to sheath and at least some of them shorter than sheath ............................................................................. 90 Stipule lobes 1–7 mm long, equal to or longer than sheath ................... 93 Inflorescence bracts strongly narrowed at base and usually shortly stipitate, 8–13 mm long; petioles 2–4 mm long ................................ P. leiantha Inflorescence bracts straight to a little narrowed at base, sessile, 2–9 mm long ....................................................................................................... 91 Inflorescence bracts numerous, involucral, enclosing capitula and/or flower groups; stipule lobes ca. 0.2 mm long ............................. P. cerronis Inflorescence bracts several to few, not involucral, subtending 1–several flowers and cymules per inflorescence axis; stipule lobes 0.5–2 mm long ....................................................................................................... 92 Inflorescence axes expanded, the flowers separated along them; pyrenes with a rather deep excavation on the ventral face .................. P. adderleyi Inflorescence axes mostly short and congested, the flowers grouped in glomerules or congested cymules; pyrenes with the ventral face flat with a central longitudinal fissure ......................................... P. officinalis Bracts enclosing flowers and/or cymules and capitula 3–9 mm long, or bracts subtending but not enclosing these ......................................... 94 Largest bracts enclosing flowers, cymules, and/or capitula 7–25 mm long .............................................................................................................. 97 Calyx limb 1–1.5 mm long .................................................................. P. berryi Calyx limb 0.3–0.5 mm long .................................................................... 95

716

R UBIACEAE

95(94). Bracts subtending flowers but not enclosing them; inflorescence with secondary axes well developed, longer than the primary axis, and verticillate at basalmost inflorescence node ...................................... P. officinalis 95. Bracts enclosing flowers; inflorescence with the secondary axes short, about equal to primary axis, and paired to verticillate ..................... 96 96(95). Lobes 1.5–2 mm long; corolla lobes 1.5–2 mm long; fruit ca. 3 mm diameter ................................................................................. P. casiquiaria 96. Stipule lobes 2–6 mm long; corolla lobes ca. 1 mm long; fruit 4–5 mm diameter ................................................................................... P. gracilenta 97(93). Corolla tube 18–21 mm long; plants of montane areas, 2300–2600 m ......................................................................................................... P. oblita 97. Corolla tube 3–11 mm long; plants of lowland, premontane, and montane areas, 90–1800 m ................................................................................. 98 98(97). Corolla tube 3–3.5 mm long; at least some of the bracts enclosing flowers and cymules < 7 mm long ............................................................... P. berryi 98. Corolla tube 5–11 mm long; bracts enclosing flowers and cymules all > 7 mm long ......................................................................................... 99 99(98). Plants of lowland areas, 90–350 m ....................................................... 100 99. Plants of premontane and montane areas, 800–1800 m ...................... 101 100(99). Leaves chartaceous to subcoriaceous, rounded to cordate at base .................................................................................................... P. blakei 100. Leaves papyraceous, rounded to shortly cordate, obtuse, cuneate, or acute at base .......................................................................... P. lupulina 101(99). Calyx limb ca. 1.5 mm long; inflorescence bracts maroon to carmine red ................................................................................. P. hemicephaelis 101. Calyx limb ca. 0.8 mm long; inflorescence bracts green to white, yellow, or orange ........................................................................................ 102 102(101). Inflorescence bracts 12–23 mm long, orange to yellow; corolla lobes 1.5–2 mm long ............................................................... P. crocochlamys 102. Inflorescence bracts 9–12 mm long, green to perhaps white; corolla lobes 3–3.5 mm long .............................................................. P. duidana 103(78). Calyx limb 1.5–4 mm long ................................................................. 104 103. Calyx limb 0.2–1.5 mm long, on at least some flowers < 1.5 mm long ........................................................................................................ 109 104(103). Stipules bilobed, with sheath 1–3 mm long and lobes 1–4.5 mm long; plants of premontane and montane areas, 1000–2300 m ........... 105 104. Stipules triangular, 1.5–6 mm long, obtuse to truncate or emarginate, sometimes with 1 or 2 caducous aristas or groups of caducous glands; plants of lowland areas, 120–600 m ................................ 106 105(104). Stipule lobes 1–2 mm long; leaves elliptic to broadly so or ovate, with intersecondary veins few to several, half as long as or shorter than the secondary veins; peduncle 2–7.5 cm long ...................... P. duricoria 105. Stipule lobes 1.5–4.5 mm long; leaves narrowly elliptic to narrowly elliptic-oblong, with intersecondary veins several and well developed, nearly as long as the secondary veins; peduncle 1–3 cm long .............................................................................................. P. speluncae 106(104). Stipules 4.5–6 mm long; floral bracts none or to 0.5 mm long ............................................................................................ P. paupertina 106. Stipules 1.5–2.5 mm long; floral bracts 0.3–1.5 mm long ................ 107

Psychotria 717

107(106). Calyx limb 2–2.5 m long ........................................................... P. microdon 107. Calyx limb 0.5–1 mm long ................................................................. 108 108(107). Stipules with a group of caducous glands inserted medially or near base on each side ........................................................... P. coussareoides 108. Stipules with caducous glands at apex ............................... P. ventuariana 109(103). Stipules interpetiolar or united around the stem, the interpetiolar portion deltoid to obtuse, rounded, truncate, emarginate, or bilobed to 1/2 their length with the lobes closely set and their sinus narrow, or with 1 or 2 caducous aristas ......................................................... 110 109. Stipules united around stem, the intrapetiolar portion sometimes shorter than the intrapetiolar but still developed and continuous, in the interpetiolar portion bilobed with the lobes separated by a concave to truncate sheath, the lobes shorter than to longer than the sheath ............................................................................................. 115 110(109). Peduncle 0.1–0.5 cm long, terminating in a cyme 0.5–0.8 cm long ...................................................................................................... P. nana 110. Peduncle 0.5–9.5 cm long, terminating in a branched cyme or paniculate portion 2–28 cm long .............................................................. 111 111(110). Inflorescence with peduncle 5–9.5 cm long and branched portion 8– 28 cm long; stipules 3–8 mm long ............................ P. bostrychothyrsus 111. Inflorescence with peduncle 0.5–7 cm long and branched portion 2– 9 cm long; stipules 1–4 mm long ................................................... 112 112(111). Inflorescence with branched portion 4–7 cm wide; calyx limb ca. 0.2 mm long ...................................................................................... P. paniculata 112. Inflorescence with branched portion 2–4 cm wide; calyx limb 0.5– 1.3 mm long ................................................................................... 113 113(112). Floral bracts ca. 1 mm long; corolla tube 6.5–7 mm long; plants from 1600 to 2000 m .............................................................. P. venezuelensis 113. Floral bracts absent or reduced, to 0.5 mm long; corolla tube 1.2– 3.5 mm long; plants from 50 to 1100 m ........................................ 114 114(113). Corolla tube 1.2–2 mm long; inflorescence with secondary axes paired or usually verticllate at basalmost node ........................ P. astrellantha 114. Corolla tube 3–3.5 mm long; inflorescence with secondary axes paired at basalmost node ................................................................. P. boliviana 115(109). Stipule lobes 5–12 mm long, at least some of them > 5 mm long .... 116 115. Stipule lobes 0.5–5 mm long, at least some of them < 5 mm long ... 118 116(115). Stigmas, ovary locules, and pyrenes of fully developed fruits 5 ............................................................................................... P. racemosa 116. Stigmas, ovary locules, and pyrenes of fully developed fruits 2 ...... 117 117(116). Leaves chartaceous to subcoriaceous, stiff-textured; floral bracts 1– 1.2 mm long; calyx limb ca. 0.5 mm long ....................... P. anartiothrix 117. Leaves papyraceous, thin-textured; floral bracts none or reduced, to 0.5 mm long; calyx limb ca. 0.2 mm long ................................ P. deflexa 118(115). Inflorescence sessile ......................................................................... P. tatei 118. Inflorescence with peduncle 0.5–10.5 cm long ................................. 119 119(118). Floral bracts 2–3.5 mm long, involucral, enclosing the glomerules and flowers .............................................................................. P. parvibractea 119. Floral bracts none, reduced, or to 4 mm long but not involucral, only loosely subtending individual flowers or cymules ....................... 120

718

R UBIACEAE

120(119). Calyx limb very reduced, ca. 0.2 mm long, truncate to undulate [P. deflexa group] ........................................................................... 121 120. Calyx limb reduced to developed, 0.3–1 mm long, truncate to undulate, dentate, or lobed ............................................................................ 123 121(120). Inflorescence with peduncle 5–7 cm long and branched portion 12– 18 cm long; petiole 6–20 mm long; corolla tube 3–3.5 mm long ....................................................................................... P. campyloneura 121. Inflorescence with peduncle 1–5.5 cm long and branched portion 1– 8 cm long; petiole 2–8 mm long; corolla tube 2–3 mm long ......... 122 122(121). Stipule with sheath and lobes developing together, the lobes continuous with the top of the sheath ................................................. P. deflexa 122. Stipule with sheath elongating on older nodes adaxially to the lobes, the lobes thus displaced to appear inserted below the top of the sheath ..................................................................................... P. venulosa 123(120). Floral bracts none or reduced, to 0.5 mm long; pyrenes smooth dorsally [Didymocarpos group] ................................................................... 124 123. At least some floral bracts present and 0.5–4 mm long; pyrenes dorsally smooth to ridged ................................................................... 127 124(123). Inflorescence with branched portion 1–2 × 1–2 cm; plants of montane areas, 2100–2185 m .............................................................. P. jauaensis 124. Inflorescence with branched portion 1–6 × 2–10 cm; plants of low to montane areas, near sea level to 2400 m ..................................... 125 125(124). Pedicels 0.5–3 mm long; calyx limb 1–1.5 mm long; corolla tube 6– 9 mm long ........................................................................... P. ceratantha 125. Pedicels none or to 1.5 mm long; calyx limb 0.3–1 mm long; corolla tube 3–6 mm long .................................................................................. 126 126(125). Corolla with tube 3–4 mm long, lobes 1–1.5 mm long; fruit 4.5–5 × 5– 6 mm ................................................................................... P. acuminata 126. Corolla with tube 4–6 mm long, lobes 1.5–3 mm long; fruit 3–4 × 4– 6 mm ..................................................................................... P. cornigera 127(123). Inflorescence with branched portion 0.5–1 × 0.5–1 cm (not including corollas); corolla tube 12–15 mm long; plants of montane areas, 2000–2500 m .......................................................................... P. concinna 127. Inflorescence with branched portion 1–14 × 1–14 cm (including corollas or not); corolla tube 2–10 mm long; plants of various areas, near sea level to 2500 m ........................................................................ 128 128(127). Calyx limb 1–1.5 mm long, on at least some flowers > 1 mm long ............................................................................................. P. ceratantha 128. Calyx limb 0.3–1 mm long ................................................................. 129 129(128). Petiole 12–45 mm long ...................................................................... 130 129. Petiole 1–11 mm long ........................................................................ 131 130(129). Inflorescence with branched portion 7–14 cm long; corolla tube ca. 3 mm long ........................................................................... P. berteroana 130. Inflorescence with branched portion 1–6 cm long; corolla tube 5– 10 mm long ............................................................................ P. everardii 131(129). Plants of premontane and montane areas, 1000–2500 m ............... 132 131. Plants of lowland areas, 50–500 m ................................................... 134 132(131). Inflorescence with branched portion 6–8 cm wide; floral bracts 2–4 mm long .................................................................................... P. heteroneura

Psychotria 719

132. 133(132). 133. 134(131). 134. 135(134). 135. 136(134). 136.

Inflorescence with branched portion 1–6 cm wide; floral bracts 0.5– 1.5 mm long ................................................................................... 133 Inflorescence with branched portion 1–6 × 1–6 cm, paniculate; fruit purple to black ....................................................................... P. everardii Inflorescence with branched portion 1–2.5 × 1–2 cm, congestedcymose; fruit deep yellow to yellow-orange .................. P. glandulicalyx Corolla tube 4–4.5 mm long; peduncle 1.5–4.3 cm long ................... 135 Corolla tube 2–3.5 mm long; peduncle 0.5–3.5 cm long ................... 136 Petiole 6–10 mm long; stipule lobes 1–1.5 mm long; peduncle ca. 4.3 cm long .................................................................................... P. barnebyana Petiole 1–3 mm long; peduncle ca. 1.5 cm long; stipule lobes 4.5–5 mm long ................................................................................. P. yapacanensis Inflorescence corymbiform, the branched portion 1–3 × 1.5–5 cm and the axes generally straight; stipule sheath ca. 1 mm long ....... P. adderleyi Inflorescence pyramidal, the branched portion 2–4 × 1.5–6 cm and the axes generally flexuous; stipule sheath ca. 0.5 mm long ...................................................................................... P. nematostachya

Psychotria subg. Psychotria Myristiphyllum P. Browne, Civ. Nat. Hist. Jamaica. 152. 1756. Psychotrophum P. Browne, Civ. Nat. Hist. Jamaica. 160. 1756. Mapouria Aubl., Hist. Pl. Guiane 175, pl. 67. 1775. Southeastern U.S.A., Mexico, Central America, the Antilles, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Argentina, Uruguay, Paraguay, Africa, Madagascar, Asia, Pacific Islands; perhaps 800 species, ca. 200 of them Neotropical, at least 29 in Venezuela, perhaps 15 in the flora area. Steyermark (1972) incorrectly considered subgenus Psychotria to be restricted to the Neotropics, but this group is pantropical (Taylor 1996) and its type, P. asiatica L., has recently been lectotypified and shown to be from China (Davis et al. 2001). Steyermark (1972; 1974) included Psychotria microdon in subgenus Psychotria, but more recent work shows that this species belongs to the Margaritopsis group, which is here provisionally included in subgenus Heteropsychotria. However, Steyermark treated P. occidentalis in subgenus Heteropsychotria, but this species is here included in subgenus Psychotria. Key to the Species of Subgenus Psychotria 1. 1. 2(1).

2.

3(2).

Petioles 1–3 mm long; stipules bilobed for ca. 1/4 their length; inflorescence capitate ........................................................................ P. occidentalis Petioles 0.5–40 mm long; stipules entire to bilobed for > 1/2 their length; inflorescences subcapitate to extensively branched ............................ 2 Inflorescences spiciform or branched to one order, with the primary and secondary axes spiciform, the flowers borne in sessile glomerules along the axes .......................................................................................... P. viridis Inflorescences subcapitate or paniculate and branched to several orders, with the flowers all borne in one glomerule or at least some of them borne singly in cymules ......................................................................... 3 Stipules bilobed, glabrescent on lower part of sheath and hirtellous on upper part and lobes, the lobes linear, 1.5–6 mm long; inflorescence

720

R UBIACEAE

subcapitate to congested-cymose, peduncle 0.1–1.3 cm long, flowering portion ca. 0.5 × 1 cm ................................................................ P. pectinata 3. Stipules acute to rounded, uniformly densely puberulous, hirtellous, strigillose, or glabrous; inflorescence branched to several orders with well-developed axes, peduncle 0.5–10.5 cm long, flowering portion 1– 10 × 2–18 cm .......................................................................................... 4 4(3). Stipules calyptrate, united into a single cap-like covering, this single structure sharply acute; inflorescences, depending on interpretation, sessile or with 2 or 3 fasciculate peduncles .......................... P. trichotoma 4. Stipules interpetiolar, free or sometimes shortly united intrapetiolarly, acute to rounded on each side; inflorescence with 1 peduncle ............ 5 5(4). Calyx limb 1.5–2.5 mm long and lobed for > 1/2 its length ...... P. horizontalis 5. Calyx limb 0.2–2 mm long, subtruncate to denticulate or lobed to ca. 1/3 its length ..................................................................................................... 6 6(5). Inflorescences pyramidal in outline with the axes rather spiciform, the higher-order axes, flowers, and fruits spreading to ca. 90° from each other .............................................................................................. P. remota 6. Inflorescences corymbiform-rounded to pyramidal in outline with the axes generally cymose, the higher-order axes, flowers, and fruits ascending to irregularly spreading at angles noticeably < 90° from each other ....................................................................................................... 7 7(6). Inflorescences with the secondary axes occasionally 2 but usually 4 and arranged in unequal pairs at the basalmost node; calyx limb 0.3– 0.7 mm long; fruit 3.5–7 × 3–6 mm ....................................................... 8 7. Inflorescences with the secondary axes 2 or 4 and subequal at the basalmost node; calyx limb 0.2–2 mm long; fruit 5–7.5 × 3.5–7 mm .............................................................................................................. 11 8(7). Corolla tube 2.5–4 mm long; inflorescences and infructescences usually terminal ............................................................................ P. carthagenensis 8. Corolla tube 1.2–2 mm long; the infructescences usually displaced to pseudoaxillary by subsequent vegetative growth from one of the axils at the base of the peduncle, the inflorescences terminal or also often displaced to pseudoaxillary ................................................................... 9 9(8). Stipules 3.5–7.5 mm long; peduncles 0.5–2.5 cm long; inflorescence with the branched portion 1.5–2.5 cm long ....................................... P. borjensis 9. Stipules 5–20 mm long; peduncles 1–9 cm long; inflorescence with the branched portion 5–15 cm long ........................................................... 10 10(9). Corolla yellow; inflorescences pyramidal in outline; fruit 5–7 × 4–6 mm ........................................................................................................... P. alba 10. Corolla white; inflorescences rounded-corymbiform to broadly pyramidal in outline; fruit 4–4.5 × 3–4 mm ................................................. P. ernestii 11(7). Inflorescence and infructescence corymbiform-rounded in outline, usually equal to or exceeded by the leaves; corolla tube 1–3 mm long ... P. anceps 11. Inflorescence and infructescence pyramidal to subglobose in outline, exceeding the leaves; corolla tube 2.5–6.5 mm long .............................. 12 12(11). Leaves generally oblanceolate or obovate; inflorescences with secondary axes usually (2)4 at the basalmost node; stipules ligulate to usually elliptic or subcircular ................................................................ P. cupularis 12. Leaves generally elliptic to elliptic-oblong; inflorescences with secondary

Psychotria 721

axes usually 2(4) at the basalmost node; stipules ligulate to obovate .............................................................................................................. 13 13(12). Calyx limb 1.5–2 mm long; corolla tube 6–6.5 mm long ................. P. irwinii 13. Calyx limb 0.5–1.2 mm long; corolla tube 2.5–5.5 mm long .................. 14 14(13). Secondary leaf veins 6–15 pairs and < 20 mm apart; peduncles 2.8– 13 cm long; inflorescence with the branched portion 3–10 cm long ............................................................................................ P. mapourioides 14. Secondary leaf veins 5 or 6 pairs and 20–25 mm apart; peduncles 2.2– 4.5 cm long; inflorescence with the branched portion 6–8.5 cm long .................................................................................................. P. yavitensis Psychotria acuminata Benth., Bot. Voy. Sulphur 107. 1845. [Subg. Heteropsychotria]. Psychotria urophylla Schltdl., Linnaea 28: 522. 1856. Subshrub or shrub to 4 m tall, glabrous; leaves 9–20 × 3.5–8.5 cm; petioles 8–20 mm long; stipules persistent, united around stem, sheath 1–2 mm long, lobes 1–4 mm long; inflorescences terminal, paniculate, peduncle 1.5–3 cm long, branched portion 1–3 × 2–4 cm, floral bracts reduced; pedicels 0.5– 1.5 mm long; calyx limb 0.3–0.5 mm long, truncate to denticulate or dentate; corolla white to cream, tube 3–4 mm long, lobes 1– 1.5 mm long; fruit didymous, 4.5–5 × 5–6 mm, blue or white; pyrenes 2, ventrally with a longitudinal furrow, dorsally smooth. Lowland to upland forests, sea level to 800 m; Delta Amacuro (lower Río Orinoco, Serranía de Imataca), Bolívar (Altiplanicie de Nuria, Guyana border at Anacoco, basins of Río Caroní, Río Caura, Río Cuyuní, and Río Paragua), Amazonas (basins of upper Río Ventuari and Río Orinoco). Apure, Barinas, Portuguesa, Zulia; Central America, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. The relatively long (1–2.5 cm), slender, usually falcate leaf tips of Psychotria acuminata are distinctive. It is similar to P. cornigera, P. cuspidata Bredem. ex Roem. & Schult., and P. rhodothamna Standl. of Amazonian Peru; the latter may not be entirely distinct. As noted by Steyermark (1972), for a long time the name P. cuspidata was incorrectly used for P. acuminata. However, P. cuspidata is a separate species of premontane forests of the Coastal Cordillera of Venezuela and the Eastern Cordillera of the Andes. The circumscription here of Psychotria acuminata, P. cornigera, and P. bahiensis

DC. differs significantly from that of Steyermark (1972); the separation of these species is outlined under P. cornigera. The calyx limbs of P. acuminata in the flora area are generally 0.3–0.5 mm long and truncate to denticulate, as described by Steyermark. However, a few specimens from the easternmost part of the flora area (i.e., Gentry & Berry 15013, MO) and from Guyana (Pipoly & Lall 8357, MO) have calyx limbs ca. 0.8 mm long and lobed for ca. 1/2 their length, that is, similar to those of P. cornigera, although the corollas of these two species are clearly different. Some fruiting specimens of these two species may be difficult to separate. Psychotria adderleyi Steyerm., Mem. New York Bot. Gard. 23: 513, fig. 72. 1972. [Subg. Heteropsychotria]. —Carrisillo, Carriso morado, Carrizo de picure, Sortija. Subshrub or shrub to 1.5 m tall, puberulous to glabrous; leaves 6.5–12.5 × 1.5–4.5 cm; petioles 3–6 mm long; stipules persistent, united around the stem, sheath ca. 1 mm long, lobes narrowly triangular, 0.5–1 mm long; inflorescence terminal, corymbiform, peduncle 0.5–2.5 cm long, branched portion 1–3 × 1.5–5 cm, floral bracts 2–4 mm long, apparently green; flowers sessile; calyx limb ca. 0.5 mm long, dentate; corolla white, tube 2–3.5 mm long, lobes ca. 2 mm long; fruit subglobose, 2.5–4 × 3.5–4 mm, purpleblack; pyrenes 2, ventrally with a deep central excavation, dorsally ridged. Riparian forests, forests on white sand bordering savannas, 50–200 m; Amazonas (basins of Río Orinoco and Río Negro). Colombia, Peru, Amazonian Brazil. Psychotria adderleyi is similar to Psychotria officinalis; P. officinalis can be

722

R UBIACEAE

separated by its flowers arranged in a few densely bracteate glomerules or congested cymules and pyrenes with a flat ventral face, versus the flowers separated or in pairs along dichotomous axes that elongate and become secund with age, the bracts fewer than the flowers, and the pyrenes with an adaxial furrow in P. adderleyi. These species are difficult to separate in flower, however, because the inflorescences of both are quite congested. The axes elongate markedly in the infructescences of P. adderleyi, so the infructescences are much larger and have a different arrangement of the axes. Psychotria adderleyi is also very similar to P. pariensis Steyerm., which is apparently restricted to the Paria Peninsula in northeastern Venezuela, and to P. manausensis Steyerm. of the Manaus region of central Amazonian Brazil. Steyermark distinguished P. manausensis from P. adderleyi by its “more densely tomentose hypanthium, more numerously flowered ultimate forks of the inflorescence, strongly reflexed basal axes with straighter, more divergently horizontal ultimate forks, reduction of most of the inflorescence bracts to minute stubs or even lacking.” Plants from near Manaus lack floral bracts; however, the floral bracts of P. adderleyi that are so evident and distinctive on the younger inflorescences fall as the flowers and fruits develop, so that the infructescences generally appear to lack bracts (e.g., Liesner 7122, MO, all stages shown). Also similar is P. kaieturensis Sandwith from the Kaietur Plateau in Guyana, which can be separated by its larger leaves, larger pyramidal inflorescences, and separate geographic distribution. Psychotria adenophora Steyerm., Mem. New York Bot. Gard. 23: 599. 1972. [Subg. Heteropsychotria]. Subshrub or shrub to 1 m tall, glabrous; leaves 10–16 × 3.5–7 cm; petioles 10–15 m long; stipules persistent, united around the stem, sheath 1.5–2 mm long, lobes 2 per side, narrowly triangular, 1–3 mm long; inflorescence terminal, capitate or subcapitate, peduncle 2.8–3.3 cm long (to 6 cm long in Brazil), head 0.6–0.8 cm diameter (to 1.5 cm diameter in Brazil), external bracts suborbicular, 5–6 mm long, green; flowers sessile; calyx limb ca. 1 mm long, lobed; corolla not seen, reportedly white; fruit ellip-

soid, ca. 5 × 3 mm, color unknown; pyrenes 2, ventrally with a longitudinal central fissure, dorsally ridged. Rocky sandstone forested slopes in montane forests on tepuis, 1200– 1500 m; Amazonas (Cerro Huachamacari). Brazil. Psychotria adenophora is similar to P. lucentifolia (S.F. Blake) Steyerm. of northern to central Venezuela; P. lucentifolia can be separated by its peduncles 1–1.5 mm long and larger, red, ovate, involucral bracts. Psychotria alba Ruiz & Pav., Fl. Peruv. 2: 58, pl. 205, fig. a. 1799. —Mapouria alba (Ruiz & Pav.) Müll. Arg., Flora 59: 458. 1876. —Uragoga alba (Ruiz & Pav.) Kuntze, Revis. Gen. Pl. 1: 299. 1891. [Subg. Psychotria]. Shrub or small tree to 15 m tall, glabrous; leaves 7.5–17 × 3–9 cm; petioles 5–15 mm long; stipules caducous, interpetiolar, ligulate to triangular, 5–11 mm long, acute to obtuse; inflorescence terminal, paniculate, peduncles 5–9 cm long, branched portion 7–15 × 11–20 cm, floral bracts ca. 0.3 mm long, pedicels 1–2 mm long; calyx limb ca. 0.5 mm long, subtruncate; corolla yellow, tube 1.5–2 mm long, lobes 1.5–2 mm long; fruit ellipsoid, 5–7 × 4–6 mm, red; pyrenes 2, dorsally ridged. Riparian forests, rain forests, evergreen seasonally dry forests, 100–900 m; southeastern Bolívar (basin of Río Caroní), Amazonas (upper Río Orinoco basin). Colombia, Suriname (doubtful), Peru, Bolivia. Psychotria alba is generally recognizable by its interpetiolar, obtuse stipules, its relatively large inflorescences with the axes usually 4 per node in unequal pairs at least at the basalmost node, and its flowers borne on well-developed pedicels in umbelliform groups. Psychotria alba is a distinctive species found in wet lowland forests of the western Amazon basin. It has frequently been confused with P. carthagenensis; see additional comments under that species. Psychotria amplectans Benth., J. Bot. (Hooker) 3: 230. 1841. [Subg. Heteropsychotria]. —Cafecillo. Psychotria paradoxa Müll. Arg. in Mart., Fl. Bras. 6(5): 313. 1881. Psychotria romboutsii Bremek., Recueil Trav. Bot. Néerl. 33. 712. 1936. Subshrub or shrub to 2 m tall, glabrous; leaves 5–13 × 1–3.5 cm, sessile; stipules per-

Psychotria 723

sistent, united around stem, sheath 1–1.5 mm long, lobes 2 per side, linear, 1–5 mm long, inserted near base of sheath; inflorescences terminal, paniculate, peduncles 0.5–4 cm long, branched portion 1–2 × 2–5 cm, floral bracts 3–11 mm long; flowers sessile; calyx limb ca. 0.3 mm long, undulate to dentate; corolla white, tube 3–4 mm long, lobes 1–1.5 mm long; fruit ellipsoid, ca. 3 mm diameter, blue; pyrenes 2, ventrally plane with a rather deep central cavity, dorsally ridged. Riparian and evergreen lowland forests, 100–200 m; southwestern Amazonas (basin of Río Casiquiare, upper Río Orinoco, and Río Negro). Suriname, Brazil, Bolivia. ◆Fig. 561. In Psychotria amplectans, the stipule lobes are inserted near the base rather than the top of the sheath. Steyermark considered this species related to several montane species of Psychotria (e.g., P. imthurmiana, P. amita Standl., P. eggersii Standl.), but those all have stipule lobes inserted on the top of the sheath and very reduced floral bracts. Thus P. amplectans seems misplaced here; it is probably related to P. lindenii. Psychotria amplectans is similar to Psychotria sipapoensis, Psychotria cardiomorpha, Palicourea lancigera, and Palicourea foldatsii, all of which can be distinguished by their stipule lobes that are inserted at the top of the stipule sheath. Psychotria amplectans is also often confused with Faramea sessilifolia. Psychotria anartiothrix Steyerm., Ann. Missouri Bot. Gard. 75: 340. 1988. [Subg. Heteropsychotria]. Shrub to 2 m tall, puberulous to glabrous; leaves 4–11 × 1–3 cm; petioles 2–6 mm long; stipules persistent, united around stem, sheath ca. 1 mm long, lobes 2 per side, linear, 5–9 mm long; inflorescence terminal, congested-cymose, peduncle 1.4–4 cm long, branched portion 0.8–2.5 × 1.5–3 cm, floral bracts 1–1.2 mm long, apparently green; flowers sessile; calyx limb ca. 0.5 mm long, dentate; corolla white, tube ca. 3.5 mm long, lobes ca. 1 mm long, with abaxial projection 0.5–1.5 mm long; fruit subglobose, ca. 5 mm diameter, orange; pyrenes 2, ventrally with a central longitudinal furrow, dorsally ridged. Flooded riparian forests, ca. 100 m; Amazonas (lower Río Baría). Southeastern Colombia.

The lanceolate, stiff-textured leaves, relatively long narrow stipule lobes, short congested inflorescences, and corolla lobes with abaxial projections are distinctive for Psychotria anartiothrix. It is known from very few collections; the fruits are described as orange, but they may be similar to those of Psychotria racemosa, which pass through an orange stage but are purple-black when mature. Psychotria anartiothrix is similar in general aspect to P. racemosa; P. racemosa can be distinguished by its stigmas and pyrenes 5. Psychotria anartiothrix also resembles Palicourea angustifolia Kunth; Palicourea angustifolia is restricted to montane forests of the Andes. Psychotria anceps Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 3: 360. 1818 [1819]. [Subg. Psychotria]. —Coloradito. Psychotria viburnoides Kunth in H.B.K., Nov. Gen. Sp. 3: 361. 1818 [1819]. Psychotria chlorantha Benth., J. Bot. (Hooker) 3: 226. 1841. Shrub or small tree to 10 m tall, glabrous; leaves 8–18 × 2.5–6 cm; petioles 0.5–3 cm long; stipules caducous, interpetiolar, triangular to ovate, 5–10 mm long, acute to obtuse or rounded; inflorescence terminal, corymbiform, peduncle 1–6 cm long, branched portion 2–12 × 4–15 cm, floral bracts 0.5–1 mm long; pedicels 0–0.5 mm long; calyx limb 0.2– 1 mm long, shortly dentate; corolla white, tube 1–3 mm long, lobes 1–3 mm long; fruit subglobose to ellipsoid, 5–6 × 4.5–6 mm, red; pyrenes 2, dorsally ridged to smooth. Riparian forests, rain forests, forested slopes over igneous substrate, dry rocky sandstone slopes in evergreen seasonally dry forests, 100–1300 m; Bolívar (basin of Río Cuyuní, Río Caroní, Río Paragua), Amazonas (basin of Río Orinoco and Río Ventuari). Anzoátegui, Apure, Carabobo, Distrito Federal, Falcón, Guárico, Monagas; Colombia, Guyana, Suriname, French Guiana, northern Brazil. The new leaves of Psychotria anceps are used to induce vomiting. A boiled infusion with brown sugar is prepared for this purpose. Psychotria anceps is generally recognizable by its relatively narrow, usually very shiny leaves that equal or usually exceed the corymbiform inflorescences, and its rela-

724

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tively large fruits. It is similar to P. mapourioides. Steyermark recognized two varieties of Psychotria anceps in Venezuela, with the plants of the Coastal Cordillera separated into var. robustior Steyerm.; var. robustior was distinguished by its markedly larger inflorescences. However, these plants are linked to his var. anceps, which is quite variable, by a number of specimens from throughout the country with inflorescences of intermediate size. Steyermark’s illustration of Psychotria anceps (1974, 1221, fig. 177) is inaccurate in its portrayal of the stipule. The stipule is shown there as shortly bilobed, a condition found in a few plants from western South America, but as noted in Steyermark’s textual description, the Venezuelan plants of this species have only acute (i.e., entire) stipules. Psychotria apoda Steyerm., Mem. New York Bot. Gard. 23: 668. 1972, not Psychotria violacea Aubl. 1775. —Tapogomea violacea Aubl., Hist. Pl. Guiane 157, pl. 60. 1775. —Cephaelis violacea (Aubl.) Sw., Fl. Ind. Occid. 1: 439. 1797. —Uragoga violacea (Aubl.) Pulle, Enum. Vasc. Pl. Surin. 446. 1906. [Subg. Heteropsychotria]. Shrub or subshrub to 3 m tall, puberulous to glabrous; leaves 8–20 × 2–9 cm; petioles 7– 15 mm long; stipules persistent, united around the stem, sheath 2–4.5 mm long, truncate to rounded, margin often hyaline or cartilaginous and to 1 mm wide, sometimes 1 or 2 subterminal groups of glands present on each side; inflorescences terminal, capitate, peduncle 0–3 mm long, heads 1–2.5 × 1.5–2 cm, external bracts 10–20 mm long, green; flowers sessile; calyx limb 1–1.5 mm long, lobed; corolla white, tube 12–15 mm long, lobes 1.5–2 mm long; fruit ellipsoid, ca. 6 × 4 mm, blue; pyrenes 2, ventrally with a longitudinal central fissure, dorsally angled to ridged. Riparian and lowland to lower montane forests, 50–500 m; Delta Amacuro (Serranía de Imataca), Bolívar (Altiplanicie de Nuria, basins of Río Caroní and Río Cuyuní, Serranía de Imataca). Eastern Colombia, Guyana, Suriname, French Guiana, Brazil. Psychotria apoda is similar to P. muscosa

(Jacq.) Steyerm. of northern Venezuela and the Guianas, and to P. hyalina Steyerm. of northern Venezuela. Psychotria muscosa differs in its inflorescences 0.8–1 cm diameter with the external bracts ca. 6 mm long; P. hyalina differs in its peduncles 4–30 cm long. Although Steyermark in his description of P. hyalina compared his new species only with P. urbaniana Steyerm. of the Antilles, P. hyalina is most similar in general aspect and details to P. apoda. Psychotria astrellantha Wernham, J. Bot. 52: 316. 1914. —Chytropsia astrellantha (Wernham) Bremek., Recueil Trav. Bot. Néerl. 31: 292. 1934. —Margaritopsis astrellantha (Wernham) L. Andersson, Syst. Geogr. Pl. 72: 230. 2002. [Subg. Heteropsychotria]. —Carrizo de picure, Yejama (Yekwana). Shrub to 2 m tall, puberulous to glabrescent; leaves 9.5–20 × 4–8 cm; petioles 3–10 mm long; stipules interpetiolar and shortly intrapetiolar, fragmenting below apical nodes, ovate to broadly rounded, 1.5–5 mm long, obtuse to rounded, shortly 1- or 2-apiculate; inflorescence terminal, paniculate, peduncles 1–2 cm long, branched portion 2–3.5 × 3–4 cm, bracts reduced; flowers sessile; calyx limb 0.5–0.8 mm long, denticulate; corolla white, tube 1.2–2 mm long, lobes 1–1.5 mm long; fruit ellipsoid to subglobose, 7–8 × 5–6 mm, deep red; pyrenes 2, ventrally plane, dorsally shallowly ridged. Evergreen lowland to lower montane forests, 50–800 m; Amazonas (San Carlos de Río Negro, near San Fernando de Atabapo). Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil. Psychotria astrellantha will belong to the segregate genus Margaritopsis Griseb. when that is eventually separated from Psychotria (Andersson 2001). Psychotria astrellantha is similar to P. boliviana, and some of the specimens treated as P. boliviana by Steyermark (1974) are here treated as P. astrellantha. The name P. astrellantha is applied somewhat provisionally here to the Venezuelan plants, because the limits of the South American species of this group are not well understood. Psychotria aubletiana Steyerm., Mem. New York Bot. Gard. 23: 694. 1972. —Cephaelis axillaris Sw., Prodr. 45.

Psychotria 725

1788, not Psychotria axillaris Willd. 1798. [Subg. Heteropsychotria]. Psychotria cacuminis Steyerm., Acta Bot. Venez. 2: 335. 1967. —Psychotria aubletiana var. cacuminis (Steyerm.) Steyerm., Mem. New York Bot. Gard. 72: 697. 1972. Psychotria aubletiana var. andina Steyerm., Mem. New York Bot. Gard. 23: 697. 1972. Psychotria aubletiana var. cacuminis f. tomentella Steyerm., Mem. New York Bot. Gard. 23: 697. 1972. Subshrub or low shrub to 1.5 m tall, hirtellous or strigillose to glabrescent; leaves 5.5–18 × 1.5–6.5 cm; petioles 3–30 mm long; stipules persistent, shortly united around stem, sheath 4–8.5 mm long, lobes ligulate to deltoid, 1–3.5 mm long, obtuse; inflorescences pseudoaxillary, capitate, sessile, heads 0.7–1.5 × 0.8–2.5 cm, bracts 6–8 mm long, green or flushed with purple; flowers sessile; calyx limb 1.5–2.2 mm long, lobed to 1/2; corolla white, tube 5–9 mm long, lobes 2– 3.5 mm long; fruit subglobose to ellipsoid, ca. 4 × 2.5 mm, blue; pyrenes 2, ventrally with 1 longitudinal furrow, dorsally ridged. Tepui slope and summit forests, 1600–2500 m; Bolívar (Aparamán-tepui, Auyán-tepui, Ilútepui, Macizo del Chimantá, Ptari-tepui, Roraima-tepui), Amazonas (Cerro Marahuaka, Sierra de la Neblina). Widespread in premontane and montane forests in Venezuela; Central America, West Indies, Trinidad, Colombia, Ecuador, Peru. Psychotria aubletiana is similar to P. celiae of the Andes, and these may not be distinct. Psychotria aubletiana varies widely in length and shape of the stipule teeth, leaf and stem pubescence, and number of secondary leaf veins. Steyermark recognized five varieties: var. aubletiana of the Antilles, var. centro-americana Steyerm. of that region, var. producta Steyerm. of the Venezuelan Andes, var. andina of southern Central America (sympatric there with var. centroamericana) to Trinidad and the flora area, and var. cacuminis of northwestern Venezuela to the flora area. He separated these by overlapping ranges of stipule size, number of secondary leaf veins, and pubescence details. Because these varieties cannot be clearly distinguished and have no evident geographic

component, they are not recognized here. Psychotria barnebyana Steyerm., Mem. New York Bot. Gard. 23: 542. 1972. [Subg. Heteropsychotria]. Shrub ca. 2 m tall, puberulous to glabrous; leaves 8–10 × 3–5 cm; petioles 6–10 mm long; stipules persistent, united around the stem, sheath ca. 2 mm long, lobes 2 per side, ligulate, 1–1.5 mm long; inflorescences terminal, paniculate, peduncle ca. 4.3 cm long, branched portion ca. 3.7 × 5.5 cm, floral bracts 1.6–1.8 mm long, apparently green; flowers apparently sessile; calyx limb ca. 0.3 mm long, undulate; corolla white, tube 4–4.5 mm long, lobes 2–3 mm long; fruit unknown. Riparian forests on igneous strata, 200–300 m; Amazonas (Piedra Tururumeri in basin of Río Pasimoni and Río Yutua). Endemic. Psychotria barnebyana is known only from the type. Based on the description and photo of the type at NY, this species is generally similar to P. berteroana but has shorter stipule lobes and petioles. Psychotria berryi Wingf., Ann. Missouri Bot. Gard. 77: 597. 1990. —Psychotria davidsei Steyerm., Ann. Missouri Bot. Gard. 71: 1177. 1984, “davidsae,” nom. illeg., not Psychotria davidsei Dwyer 1980. [Subg. Heteropsychotria]. Subshrub ca. 0.5 m tall, puberulous or hirtellous to glabrescent; leaves 6–26 × 4–13 cm; petioles 4–12 mm long; stipules persistent, united around stem, sheath 1–1.5 mm long, lobes 2 per side, triangular, ca. 1.5 mm long; inflorescence terminal, corymbiform to congested-cymose, peduncle 0.5–2.5 cm long, branched portion 1–2 × 3–5 cm, bracts 4–10 mm long, green to white; flowers sessile; calyx limb 1–1.5 mm long, lobed; corolla white, tube 3–3.5 mm long, lobes 1–1.5 mm long; fruit not seen. Rain forests, 200–1200 m; Bolívar (Sierra Pakaraima), Amazonas (basin of Río Orinoco southeast of Puerto Ayacucho). Endemic. Psychotria berryi is similar to P. buchtienii (H. Winkler) Standl. of western South America and Central America. These may in fact not be distinct, but P. berryi is too poorly known to evaluate this. Psychotria berteroana DC., Prodr. 4: 515. 1830. [Subg. Heteropsychotria].

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Shrub or small tree to 10 m tall, pilosulous or puberulous to glabrous; leaves 10–27 × 4– 12.5 cm; petioles 12–45 mm long; stipules persistent, united around stem, sheath 1.5–3 mm long, lobes 2 per side, narrowly triangular, 1–3.5 mm long; inflorescences terminal, paniculate, peduncle 4–10.5 cm long, branched portion 7–14 × 7–14 cm, floral bracts 1–2 mm long, green; flowers sessile; calyx limb 0.8–1 mm long, lobed; corolla white, tube ca. 3 mm long, lobes ca. 1.5 mm long; fruit subglobose, ca. 3 mm diameter, black; pyrenes 2, ventrally with a central invagination, dorsally ridged. Wet forests, montane forests, 200–1400 m; Bolívar (Macizo del Chimantá, basins of Río Aponguao and Río Cuyuní), Amazonas (upper Río Orinoco, Sierra de la Neblina). Mexico, Central America, Antilles, Colombia, Guyana, Ecuador, Peru, Brazil, Bolivia. This name was published and has been widely cited as “berteriana,” but recent changes to the International Code of Botanical Nomenclature (Greuter et al. 2000) require this to be corrected to “berteroana.” This species is here circumscribed differently than was done by Steyermark (1972; 1974). He treated this species as wide-ranging and variable, especially in stipule form, bract size, pubescence, and sizes of inflorescences, corollas, and fruits. He recognized three varieties: var. berteroana of the Antilles; var. venezuelica Steyerm. of Guyana and montane Venezuela; and var. luxurians (Rusby) Steyerm. of western South America and lowland Venezuela including the flora area. Steyermark did not mention the Central American and Mexican plants in his treatment. More recently, Taylor (e.g., 2001a) has recognized two distinct species, P. berteroana and P. luxurians Rusby. In this circumscription, followed here, P. luxurians is found from Costa Rica to Bolivia and differs from P. berteroana in its stipules that usually only persist on the apical nodes and are interpetiolar and rounded to truncate or emarginate (except at the node subtending the peduncle, which frequently has deeply bilobed stipules with obtuse lobes); its floral bracts 1–2.5 mm long; its calyx limb 0.5–1 mm long; its corolla tubes 2.5–4 mm long and lobes 1.2–1.5 mm long; and its fruits 2.5–3 mm diameter (where these species are sympatric, P. berteroana differs clearly from P. luxurians in the sizes of these structures,

though it approaches these sizes outside the range of P. luxurians). Psychotria luxurians has not yet been seen from the flora area. Steyermark’s P. berteroana subsp. venezuelica was distinguished only by details of the pubescence and a generally shorter though overlapping inflorescence size, and is not recognized here. Psychotria blakei Standl. & Steyerm., Fieldiana, Bot. 28: 596, fig. 133. 1953. [Subg. Heteropsychotria]. Subshrub to 1.5 m tall, glabrous; leaves 8.5–15 × 3–6 cm; petioles 2–5 mm long; stipules apparently persistent, united around stem, sheath ca. 2 mm long, lobes 2 per side, narrowly triangular, 3–4 mm long; inflorescence terminal, corymbiform to congested-cymose, peduncle 2.5–5 cm long, branched portion 1.5–3 × 3–4.5 cm, bracts 9– 15 mm long, red to red-orange; flowers sessile; calyx limb ca. 0.5 mm long, dentate; corolla white, tube ca. 7 mm long, lobes ca. 2.5 mm long; fruit subglobose, ca. 4.5 × 4 mm, white; pyrenes 2, ventrally with a central longitudinal fissue, dorsally ridged. Riparian forest bordering savanna, dwarf forest on quartzite outcrops, 100–200 m; Amazonas (base of Cerro Yapacana, Sabanita Morocoto below mouth of Río Atabapo, Río Orinoco basin). Northern Brazil. Psychotria blakei is a poorly known species similar to P. lupulina (see comments there) and P. maguireorum. The figure in the original description of P. blakei shows the interior of the opened calyx having groups of well-developed colleters borne below some of the sinuses between the calyx lobes. Psychotria bolivarensis (Standl. & Steyerm.) Steyerm., Mem. New York Bot. Gard. 23: 651. 1972. —Cephaelis bolivarensis Standl. & Steyerm., Fieldiana, Bot. 28: 567. 1953. [Subg. Heteropsychotria]. Shrub to 3 m tall, densely hirtellous often becoming glabrescent; leaves 5.5–9 × 1.5–2.5 cm; petioles 3–10 mm long; stipules persistent, united around the stem, sheath 2–3 mm long, lobes 2 per side, narrowly triangular, 3– 6 mm long; inflorescences terminal, capitate, peduncles 0.1–1.5 cm long, heads 0.5–1 cm diameter, external bracts 10–20 mm long, apparently green, floral bracts 5–8 mm long; flowers sessile; calyx limb 3.5–4 mm long,

Psychotria 727

deeply lobed; corolla unknown; fruit ellipsoid to ovoid, ca. 7 × 5 mm, purple; pyrenes 2, ventrally with 1 slender longitudinal furrow, dorsally smooth. Upper montane tepui forests, 2200–2300 m; Bolívar (Sororopántepui), Amazonas (base of Cerro Duida, Cerro Huachamacari, south of Coromoto, Río Cunucunuma). Colombia, Ecuador, Peru. Psychotria bolivarensis is similar to P. rosea in general aspect; P. rosea can be separated by its shorter calyx limb and colored inflorescence bracts. Psychotria bolivarensis is also similar to P. tetramera Steyerm. of Guyana, P. caquetensis Steyerm. of Colombia, and Cephaelis florentina Standl. ex Steyerm.; in fact, these last three may not actually be distinct but they are poorly known. Psychotria boliviana Standl., Publ. Field Columbian Mus., Bot. Ser. 7: 302. 1931. [Subg. Heteropsychotria]. Psychotria foetidiflora Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 22: 20. 1940. Psychotria kukenanensis Steyerm., Fieldiana, Bot. 28: 601. 1953. Psychotria ronaldii Steyerm., Ann. Missouri Bot. Gard. 74: 111, fig. 10. 1987. Shrub or small tree to 4 m tall, strigillose or hirtellous to glabrous; leaves 10–22(30) × 2.5–8(11) cm; petioles 3–25 mm long; stipules fragmenting below apical nodes, interpetiolar and shortly intrapetiolar, 1.5–4 mm long, ovate to broadly rounded, truncate to obtuse, shallowly emarginate, minutely bidentate or with 1 or 2 usually caducous aristas to 4 mm long; inflorescences terminal, paniculate, peduncles 0.5–4 cm long, branched portion 2–6 × 2–4 cm, bracts reduced; flowers sessile; calyx limb 0.8–1.3 mm long; corolla white to cream, tube 3–3.5 mm long, lobes 2.5–3 mm long; fruit ovoid to ellipsoid, 6–9 × 4–5 mm, deep red; pyrenes 2, ventrally plane, dorsally shallowly ridged. Forested slopes on igneous substrate and along stream in gallery forests, forested tepui slopes, 100–1100 m; Bolívar (Gran Sabana at San Ignacio de Yuruaní, north of Santa Elena de Uairén), Amazonas (below Cerro Arauicaua, Cerro Huachamacari, Río Yatúa, Serranía Batata 55 km southeast of Puerto Ayacucho). Guyana, French Guiana, Colombia, Amazonian Ecuador, Peru, Brazil, Bolivia. Psychotria boliviana will belong to the segregate genus Margaritopsis when that is separated from Psychotria. Psychotria bo-

liviana is similar to P. astrellantha, and some of the specimens that would have been included by Steyermark (1974) in P. boliviana are here treated as P. astrellantha. Psychotria boliviana is here treated as a relatively widespread species that varies in pubescence pattern, inflorescence arrangement (i.e., many specimens from western Amazonia have the secondary inflorescence axes 4 in unequal pairs, versus paired in eastern Amazonia), and stipule form. Steyermark named Psychotria ronaldii as an endemic species of Cerro Huachamacari, but it differs from P. boliviana only in the relatively large leaves on the type collection. Another collection from the same locality (Liesner 25592, MO!) has leaves on a single stem with the entire size range known from P. boliviana together with P. ronaldii. Steyermark also distinguished P. ronaldii from P. boliviana in his protologue by its supposedly smaller corollas, but these fall within the size range found in P. boliviana in Venezuela. Steyermark distinguished Psychotria puberulenta Steyerm. of Guyana from P. boliviana only by its puberulous hypanthium and smaller inflorescences. However, these supposedly smaller inflorescences fall within the size range found in P. boliviana in Venezuela (cf. Steyermark 1974), and specimens from Venezuela that were determined by Steyermark as P. puberulenta (Liesner 24239, 24466, MO) have the hypanthia glabrous or sparsely pubescent on their lower portions. Thus, P. puberulenta does not seem to be present in Venezuela, if in fact it is distinct from P. boliviana. Psychotria borjensis Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 3: 357. 1818 [1819]. —Mapouria borjensis (Kunth) Müll. Arg. in Mart., Fl. Bras. 6(5): 404. 1881. [Subg. Psychotria]. Mapouria umbrosa Müll. Arg., Flora 59: 459, 466. 1876. Shrub to 3 m tall, densely puberulous to glabrescent; leaves 5–14 × 1.3–3 cm; petioles 0.5–10 mm long; stipules caducous, interpetiolar, triangular to ovate, 3.5–7.5 mm long, acute; inflorescences terminal, paniculate, peduncle 0.5–2.5 cm long, branched portion 1.5–2.5 × 5–7 cm, floral bracts ca. 1 mm long; flowers sessile; calyx limb 0.3–0.7 mm long, shallowly dentate; corolla white or

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cream, tube 1.2–2 mm long, lobes ca. 1.2 mm long; fruit ellipsoid, 3.5–5 × 3 mm, red; pyrenes 2, dorsally ridged. Riparian forests, bases of igneous outcrops, borders of savannas, savannas, temporarily flooded savannas, 100–200 m; Bolívar (Río Erebato), Amazonas (basins of Río Orinoco, Río Cuyuní, and Río Ventuari east to San Juan de Manapiare and Río Cunucunuma). Colombia, Guyana, Suriname, French Guiana, Brazil, Bolivia, and possibly Ecuador and Peru. Psychotria borjensis is generally recognizable by its interpetiolar acute stipules, its quite slender inflorescences axes, its inflorescences with the secondary axes usually 4 in unequal pairs at least at the basalmost node, its relatively small corollas, and its relatively small fruits. It is similar to Psychotria carthagenensis; see additional comments under that species. Psychotria bostrychothyrsus Sandwith, Kew Bull. 1939: 553. 1940. [Subg. Heteropsychotria]. Subshrub or shrub to 4 m tall, glabrous; leaves 14–24 × 3.5–11 cm; petioles 6–35 mm long; stipules persistent, interpetiolar and shortly intrapetiolar, ovate to triangular, 3–8 mm long, emarginate to shortly bilobed; inflorescence terminal, paniculate, peduncle 5– 9.5 cm long, branched portion 8–28 × 4–23 cm, floral bracts 1–3.5 mm long, green; flowers sessile; calyx limb ca. 0.3 mm long, lobed; corolla white sometimes tinged with lavender, tube 3–3.5 mm long, lobes 2.5–3 mm long; fruit subglobose to ovoid, 4 × 4–4.5 mm long, blue; pyrenes 2, ventrally with a rather deep invagination arising from a central longitudinal fissure, dorsally ridged. Upland forests, 400–1000 m; Bolívar (basins of Río Cuyuní and upper Río Caroní west to Macizo del Chimantá and Cerro Abismo). Guyana, Suriname, French Guiana, northeastern Brazil. Psychotria bostrychothyrsus can be recognized by its relatively large inflorescences and infructescences with slender, usually flexuous axes, its rather large leaves, and its flowers in regularly dichotomous cymules. Psychotria paniculata, P. berteroana, P. ramiflora Rusby of Bolivia and Peru, and P. angustiflora K. Krause of Central America and northwestern South America also have rather large, much-branched inflorescences, but these species have shorter inflorescence axes and usually rather irregular branching

in the cymules. Psychotria bostrychothyrsus is also similar to and often confused with P. deflexa and P. subundulata. Psychotria bracteocardia (DC.) Müll. Arg. in Mart., Fl. Bras. 6(5): 362. 1881. —Cephaelis bracteocardia DC., Prodr. 4: 534. 1830. [Subg. Heteropsychotria]. —Tulipán montañero morado, Tulipán morado, Saquí ya rojo. Cephaelis pubescens Hoffmanns. ex Roem. & Schult., Syst. Veg. 5: 213. 1819, not Psychotria pubescens Sw. 1797. Shrub or subshrub to 4 m tall, puberulous to pilosulous; leaves 9–25 × 2–9 cm; petiole 4–10 mm long; stipules persistent, united around the stem, sheath 0.5–2 mm long, lobes 2 per side, narrowly triangular, 4–9 mm long; inflorescences terminal, capitate, peduncle 2–7 cm long, head 1–1.5 × 1.5–4 cm, external bracts 1.5–3.5 cm, pale green to purple or cream; flowers sessile; calyx limb 0.8–1 mm long, dentate; corolla white to usually purple, tube 3.5–4.5 mm long, lobes 1–2 mm long; fruit ellipsoid, 3–4 × 2–3 mm, blue; pyrenes 2, ventrally with a slender central longitudinal fissure, dorsally ridged to nearly smooth. Riparian and evergreen lowland to montane forests, 100–1100 m; Bolívar (common), Amazonas (basin of Río Orinoco). Apure, Monagas, Portuguesa, Sucre, Zulia; Colombia, Trinidad, Guyana, Suriname, French Guiana, Brazil. Psychotria bracteocardia is similar to P. colorata, P. rosea, and P. ctenophora Steyerm. Psychotria bracteocardia can be recognized by its inflorescences with the floral bracts obtuse to rounded, all of the bracts pilosulous throughout, and the corollas with relatively short tubes. Typically only the corolla lobes and anthers are visible above the floral bracts, though the old corollas frequently remain attached to the developing fruits. Psychotria ctenophora of Anzoátegui, French Guiana, and Suriname was separated by Steyermark based on its calyx limb 1–2 mm long, its calyx and hypanthium externally sericeous (versus puberulous in P. bracteocardia), and its corolla puberulous over most of the external surface (versus only in the upper 1/3 in P. bracteocardia); its status as a separate species may deserve re-evaluation. Psychotria brazaoi Steyerm., Mem. New York Bot. Gard. 23: 610, fig. 83. 1972, “brazoi.” [Subg. Heteropsychotria].

Psychotria 729

Tree ca. 5 m tall, glabrous; leaves 10–19 × 4–8 cm; petioles 10–20 mm long; stipules persistent, united around stem, sheath ca. 1.5 mm long, lobes 2 per side, triangular, ca. 1.5 mm long; inflorescence terminal, paniculate, peduncle 2.2–5.5 cm long, branched portion 1.7–2 × 3–4.5 cm, floral bracts 5–9 mm long, apparently green; flowers sessile; calyx limb 0.8–1 mm long, dentate; corolla cream, tube ca. 8 mm long, lobes ca. 5 mm long; fruit unknown. Tepui slope forests, 1400–1800 m; expected in Amazonas. Adjacent Brazil (Amazonas: Serra da Neblina). Psychotria brazaoi can be recognized by its relatively long corollas borne in several congested groups, its few but relatively large floral bracts, and its nodes and stipules that become rather discolored when dry. This species was named for its collector, Umbelino Brazão, but was mistakenly published as “brazoi.” Steyermark reported this species also from the eastern slopes of the eastern Colombian Andes, but that one specimen has not been seen. Such a geographic pattern would be unusual. Another specimen from the Sierra de Pakaraima that was determined by Steyermark as P. brazaoi (Steyermark et al. 107206, MO) is here included in P. berryi. Psychotria campyloneura Müll. Arg. in Mart., Fl. Bras. 6(5): 302. 1881. —Psychotria deflexa subsp. campyloneura (Müll. Arg.) Steyerm., Mem. New York Bot. Gard. 23: 504. 1972. [Subg. Heteropsychotria]. Shrub or small tree to 6 m tall, glabrous; leaves 14–24 × 3.5–9 cm; petioles 6–20 mm long; stipules persistent, united around stem, sheath 1–2.5 mm long, lobes 2 per side, triangular, 1–3 mm long; inflorescences terminal, paniculate, peduncle 5–7 cm long, branched portion 12–18 × 3–8 cm, floral bracts reduced, pedicels 0–2 mm long; calyx limb ca. 0.2 mm long, undulate; corolla white, tube 3–3.5 mm long, lobes 1–1.2 mm long; fruit subglobose, 3–4 mm diameter, purple-black; pyrenes 2, ventrally smooth and concave, often rather deeply so, dorsally ridged and shallowly alveolate. Riparian and flooded forests, talus slope forests, 100–600 m; Delta Amacuro (Serranía de Imataca), Bolívar (Amaruay-tepui, Río Samay near Cerro Abismo), Amazonas (Río Negro basin, upper Río Orinoco, Río Siapa). Central

America, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil. Psychotria campyloneura was included by Steyermark in his circumscription of Psychotria deflexa; see the discussion under that species. The name used here for this species follows Steyermark’s usage; however, the older name P. paniculata may apply to this species; see comments under that species. Psychotria campylopoda Standl., Publ. Field Columbian Mus., Bot. Ser. 7: 437. 1931. [Subg. Heteropsychotria]. Rather delicate shrub or subshrub to 4 m tall, hirtellous or puberulous to glabrescent; leaves 2.5–7 × 1–3.5 cm; petioles 1–5 mm long; stipules persistent, united around the stem, sheath 0.3–1 mm long, lobes 2 per side, triangular, 1–2 mm long; inflorescences terminal or sometimes later displaced to pseudoaxillary, capitate, peduncle 0.7–3.5 cm long, heads 0.5–0.8 × 0.5 cm, bracts 3–7 mm long; flowers sessile; calyx limb ca. 0.2 mm long, dentate; corolla white, 3–4 mm long, lobes 1–2 mm long; fruit subglobose, 4–5 × 3 mm, deep blue to blue-black; pyrenes 2, ventrally plane with a longitudinal slit in the basal half, dorsally smooth. Tepui slope and summit forests, 1100–2200 m; Bolívar (Auyán-tepui, Cerro Guaiquinima, Macizo de Chimantá), Amazonas (Cerro Duida, Cerro Huachamacari, Cerro Parú, Sierra Parima, Yutajé). Adjacent Guyana, Brazil. ◆Fig. 569. In Psychotria campylopoda the leaf margins are typically revolute, and at first glance the secondary veins may appear united to a thickened margin, which is not actually the case. This species is similar to P. hoffmannseggiana, P. vichadensis, and P. concinna (when corollas are not present). Psychotria capitata Ruiz & Pav., Fl. Peruv. 59, pl. 206, fig a. 1709. [Subg. Heteropsychotria]. A common understory plant of upland, flooded, and gallery forests; Central America, Colombia, Venezuela, Trinidad, Tobago, Ecuador, Peru, Brazil, Bolivia; 2 or possibly 3 subspecies, 1 in Venezuela. Psychotria capitata is similar to P. maguireorum and P. stipulosa (one or both of which may not be distinct from P. capitata), and for no apparent reason is often confused with P. lupulina and P. lindenii. The characteristic stipules help identify P. capitata.

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This species or complex of species is widely variable in stipule size, shape, and degree of lobing; inflorescence size and degree of branching; size, color, and arrangement of the inflorescence bracts; corolla size; and leaf size. Steyermark attempted to segregate the variation in this group in the flora area into several species, subspecies, varieties, and forms. Within this complex, Psychotria stipulosa is separated by its relatively large, entire to shallowly lobed stipules; in the flora area this character is correlated with relatively broad inflorescence bracts (1.5–4 mm wide) and relatively long corolla tubes (8–11 mm long). Psychotria stipulosa is recognized here provisionally as a species of eastern and central Amazonia, although plants with similar distinctive stipules but narrower bracts and shorter corollas are found in western Amazonia. Psychotria maguireorum is also provisionally recognized here, and distinguished by its relatively longer corolla tubes (11–13 mm long) and pink, purple, or orangebrown inflorescence axes, bracts, and/or corollas. Additionally, Steyermark distinguished P. maguireorum by the width of the basalmost inflorescence bracts, 2–4 mm wide versus 0.5–1.5 mm wide in P. capitata, but with more specimens now available this character clearly varies continuously across these two groups. Steyermark recognized three subspecies of Psychotria capitata: subsp. capitata, subsp. inundata, and subsp. amplifolia (Raeusch.) Steyerm., but they are all linked by intermediate characters. P.

capitata subsp. inundata (Benth.) Steyerm., Mem. New York Bot. Gard. 23: 618. 1972. —Psychotria inundata Benth., J. Bot. (Hooker) 3: 229. 1841. [Subg. Heteropsychotria]. —Tulipán. Psychotria arcuata Benth., J. Bot. (Hooker) 3: 229. 1841. Psychotria setifera Benth., J. Bot. (Hooker) 3: 228. 1841. Declieuxia roraimensis Wernham, J. Bot. 52: 325. 1914. —Psychotria capitata subsp. inundata var. roraimensis (Wernh.) Steyerm., Mem. New York Bot. Gard. 23: 619. 1972. Psychotria cubitalis Standl. & Steyerm., Fieldiana, Bot. 28: 598. 1953. Psychotria heterocarpa Standl. & Steyerm., Fieldiana, Bot. 28: 600. 1953.

Psychotria capitata subsp. inundata var. septentrionalis Steyerm., Mem. New York Bot. Gard. 23: 621. 1972. Subshrub or shrub to 3(–6) m tall, hirtellous to glabrous; leaves 6–25 × 1.5–9 cm; petioles 2–30 mm long; stipules usually deciduous after distalmost nodes, interpetiolar and sometimes shortly intrapetiolar, sheath 0.5–2 mm long, lobes 2 per side, narrowly triangular, 2–15 mm long; inflorescences terminal, paniculate, peduncles 1.5–11.5 cm long, branched portion 1.5–8 × 2–7.5 cm, bracts 2.5–15 mm long; flowers sessile; calyx limb 0.5–1 mm long, dentate; corolla white to yellow or cream, tube 3.5–7.5 mm long, lobes 1.5–4 mm long; fruit ellipsoid, 5–6 × 5–6 mm, purple-black; pyrenes 2, ventrally with a longitudinal fissure, dorsally ridged. Riparian forests, gallery forests, forests on igneous or sandstone formations, 100–1400 m; common in Bolívar and Amazonas. Widespread elsewhere in lowland Venezuela; Central America, Trinidad, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 559. Psychotria cardiomorpha C.M. Taylor & A. Pool, Novon 4: 303. 1994. —Psychotria cordifolia Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 3: 365. 1818 [1819], nom. illeg., not Psychotria cordifolia F. Dietr. 1807. [Subg. Heteropsychotria]. Psychotria cordifolia subsp. perpusilla Steyerm., Mem. New York Bot. Gard. 23: 708. 1972, nom. illeg. —Psychotria cardiomorpha subsp. perpusilla (Steyerm.) C.M. Taylor & A. Pool, Novon 4: 303. 1994. Subshrub or shrub to 4(–5) m tall, glabrous; leaves 2.5–9.5 × 1.5–5.5 cm, sessile or subsessile; stipules persistent, united around stem, sheath 0.5–1 mm long, lobes 2 per side, narrowly triangular, 1–3.5 mm long; inflorescences terminal, paniculate, peduncles 2–8 cm long, branched portion 2.5–8 × 2–5 cm, floral bracts 0.5–2 mm long; flowers sessile; calyx limb 0.3–0.5 mm long, dentate; corolla white sometimes flushed with pink, tube 2–3 mm long, lobes 1–2 mm long; fruit ellipsoid to subglobose, 2–3 × 2–3 mm, purple-blue; pyrenes 2, ventrally shallowly concave, dorsally shallowly ridged and with surface often shallowly foveolate. White-sand savannas, borders of forests, forested rocky igneous slopes, 100–1000 m; Amazonas (basins of Río

Psychotria 731

Orinoco and Río Ventuari). Colombia, Guyana, Suriname, French Guiana, Brazil. Psychotria cardiomorpha is similar to Psychotria sipapoensis, Palicourea lancigera, and Palicourea foldatsii. Steyermark recognized two subspecies of Psychotria cardiomorpha (as Psychotria cordigera), the typical one and subsp. perpusilla of the Guianas. He distinguished these by details of the pubescence, flower size, details of the inflorescence branching, and degree of development of the bracts. However, in addition to plants that fall clearly in subsp. cordifolia (e.g., Gröger 940, MO) or subsp. perpusilla (e.g., Stergios 112568, MO; Fernández & Delgado 5828, MO), there are a number of plants from the flora area that are fully intermediate between these two subspecies (e.g., Gröger 874, MO); consequently, these subspecies are not recognized here. Psychotria carrenoi Steyerm. in Lasser & Steyerm., Fl. Venez. 9(3): 1658. 1974. [Subg. Heteropsychotria]. Shrub to 2 m tall, glabrous; leaves 2–4.5 × 0.5–2 cm, sessile or subsessile; stipules persistent, united around the stem, sheath 1.5–2 mm long, on each side with 1 projection, this narrowly triangular, 2.5–4 mm long; inflorescences pseudoaxillary and/or axillary, subcapitate to congested-cymose, peduncles 0.5– 2 cm long, branched portion or capitulum ca. 0.5 × 0.5 cm, bracts 1.5–2.5 mm long, pedicels 0–1 mm long; calyx limb 2–3 mm long, deeply lobed; corolla white sometimes flushed with purple, tube 2.5–4 mm long, lobes 1.5–3 mm long; fruit unknown. Tepui meadows with Brocchinia hechtioides, tepui meadows, 1800–2300 m; Bolívar (summit of Cerro Jaua). Endemic. Psychotria carrenoi is similar to P. phelpsiana. These species differ basically in their calyx length, in spite of the long list of continuously varying characters that Steyermark listed to separate them (1974, 1660). Both of these species are similar to P. durifolia, and all three of these probably do not belong to Psychotria. However their affinities otherwise are not clear (perhaps with Coccochondra), so they are provisionally treated in Psychotria here. Psychotria carthagenensis Jacq., Enum. Syst. Pl. 16. 1760. [Subg. Psychotria]. Psychotria foveolata Ruiz & Pav., Fl.

Peruv. 2: 59, pl. 207, fig. b. 1799. Psychotria fockeana Miq., Linnaea 18: 296. 1844. —Mapouria fockeana (Miq.) Bremek., Recueil Trav. Bot. Néerl. 31: 286. 1934. Shrub or small tree to 5 m tall, puberulous to glabrous; leaves 6–15 × 2–6.5 cm; petioles 3–10 mm long; stipules caducous, interpetiolar and often coherent intrapetiolarly, ligulate to ovate, 3–10 mm long, obtuse to rounded; inflorescences terminal, paniculate, peduncle 3.5–6 cm long, branched portion 3– 6 × 2.5–6 cm, floral bracts 0.5–1 mm long, pedicels 0–0.5 mm long; calyx limb ca. 0.5 mm long, lobed to 1/2 its length; corolla white to cream, tube 2.5–4 mm long, lobes 1.5–2 mm long; fruit ellipsoid, 4.5–5.5 × 3.5–4 mm, red; pyrenes 2, dorsally ridged. Gallery forests, forested igneous outcrops, secondary growth, rain forests, 100–200 m; Bolívar (Río Cuyuní), Amazonas (near San Carlos and Puerto Ayacucho). Common and widespread elsewhere in Venezuela; Mexico, Central America, Antilles, Colombia, Trinidad, Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina, Uruguay. Psychotria carthagenensis is usually recognizable by its stipules that remain coherent and loosely encircling the stem for a while after their abscission, its inflorescences with the secondary axes usually 4 in unequal pairs at several nodes, and its relatively welldeveloped corolla tubes. Psychotria carthagenensis is quite similar to P. alba, P. borjensis, and P. ernestii of the flora area, and care must be taken to separate these species. Psychotria casiquiaria Müll. Arg. in Mart., Fl. Bras. 6(5): 324. 1881. [Subg. Heteropsychotria]. Shrub to 3 m tall, pilosulous or puberulous to glabrous; leaves 7.4–13 × 2.5–5 cm; petioles 3–10 mm long; stipules persistent, united around the stem, sheath 1–2 mm long, lobes 2 per side, triangular, 1.5–2 mm long; inflorescences terminal, capitate to subcapitate or congested-cymose, peduncles 0.2–2.2 cm long, capitulum or branched portion 0.8–1.5 × 1–2 cm, bracts 3–6 mm long; flowers sessile; calyx limb 0.3–0.5 mm long, dentate; corolla white, tube ca. 3.5 mm long, lobes 1.5–2 mm long; fruit ellipsoid, ca. 3 × 3 mm, purple or blue; pyrenes 2, ventrally with a longitudinal central fissure that is widened

732

R UBIACEAE

in the distal 1/2, dorsally ridged. Rain forests, montane forests, forests on white sand, 100– 800 m; Bolívar (El Abismo south of El Paují), Amazonas (foot of Cerro Huachamacari, Río Mawarinuma, basins of Río Orinoco and Río Negro). Colombia, Peru, Amazonian Brazil, Bolivia. ◆Fig. 570. Psychotria casiquiaria is similar to P. spadicea, P. gracilenta, P. platypoda, and P. hoffmannseggiana. Like P. gracilenta and P. platypoda, P. casiquiaria has inflorescences that are capitate and peduncles that are only partially elongated when the flowering begins, but then as the inflorescences develop the peduncles elongate and in some cases the inflorescence axes also begin to elongate. Then typically by the time the fruits begin to mature, the inflorescences are notably larger with evident to rather well-developed axes (e.g., Liesner 7560, MO; this one specimen shows all the developmental stages). Plants of the flora area vary in degree of expansion of the infructescence, from quite compact (e.g., Stergios et al. 4383, ca. 0.8 × 1 cm) to relatively expanded (e.g., Liesner 3720, 1.5–3 × 3–3.5 cm). This relatively expanded form matches the type of P. casiquiaria (Spruce 3436, P, G-DC, isotypes, photos MO!). Psychotria celiae Steyerm., Mem. New York Bot. Gard. 23: 700, fig. 88. 1972. [Subg. Heteropsychotria]. Subshrub to 1 m tall, hirtellous to glabrescent; leaves 8–12 × 3.8–5.5 cm; petioles 10– 17 mm long; stipules persistent, shortly united around stem, sheath 7–8 mm long, lobes 2 per side, deltoid, 2.5–3 mm long, obtuse; inflorescences pseudoaxillary, capitate, sessile, capitulum 1.3–1.5 × 2.5 cm, bracts 9– 10 mm long, apparently green; flowers sessile; calyx limb 2–2.2 mm long, lobed; corolla white, tube ca. 8 mm long, lobes ca. 2 mm long; fruit unknown. Upper montane tepui forests, ca. 2400 m; expected in Amazonas. Brazil (Amazonas: Serra da Neblina). Psychotria celiae is quite similar to P. aubletiana, and it may eventually be shown to be only a form of that variable and wideranging species. Psychotria ceratantha Standl., Publ. Field Columbian Mus., Bot. Ser. 7: 439. 1931. [Subg. Heteropsychotria]. Psychotria arenaria Standl. & Steyerm., Fieldiana, Bot. 28: 594. 1953.

Psychotria yutajensis Steyerm., Ann. Missouri Bot. Gard. 75: 348. 1988. Shrub or small tree to 4(–8) m tall, glabrous; leaves 6–21 × 2–8 cm; petioles 6–15 mm long; stipules persistent, united around stem, 1–3 mm long, lobes 2 per side, 1–3 mm long; inflorescences terminal, congested-cymose to paniculate, peduncles 0.7–4.5 cm long, branched portion 2–6 × 2–10 cm, floral bracts reduced or few and 0.5–1 mm long; pedicels 0.5–3 mm long; calyx limb 1–1.5 mm long, lobed to 1/2 its length; corolla greenish white to white, cream, or pink-purple, tube 6–9 mm long, lobes 3–4 mm long, with abaxial appendages 0.3–1 mm long; fruit didymous, 5–6 × 6–10 mm, blue to purpleblack; pyrenes 2, ventrally with a slender central furrow, dorsally smooth. Tepui summit and slope forests and savannas, 1000– 2400 m; Bolívar (Aprada-tepui, Auyán-tepui, Cerro Jaua, Macizo del Chimantá, Ptaritepui, Roraima-tepui, Sororopán-tepui), Amazonas (Cerro Aracamuni, Cerro Duida, Cerro Huachamacari, Cerro Marahuaka, Cerro Yaví, Cerro Yutajé, Sierra de la Neblina). Guyana. Psychotria ceratantha is similar to Psychotria jauaensis, P. acuminata, and P. bahiensis. It is here treated as a rather variable complex of forms found at higher elevations throughout most of the Guayana Highland. In general, plants from different mountains cannot be separated morphologically, and the flower sizes are similar throughout the region. There is some variation in inflorescence size and arrangement, which is correlated with developmental stage: the inflorescences initially have relatively short, congested axes, which become longer and more spreading as the flowers mature and the fruits develop. This developmental variation has been confused with geographic variation, probably due to the small number of specimens available from most sites. Psychotria cerronis Steyerm., Mem. New York Bot. Gard. 23: 886, fig. 17. 1972. [Subg. Heteropsychotria]. Shrub to 1.5 m tall, glabrous; leaves 4.5–6 × 1.5–2.3 cm; petioles 6–9 mm long; stipules apparently persistent, united around stem, sheath ca. 3 mm long, lobes 2 per side, ligulate, ca. 0.2 mm long; inflorescence terminal, capitate to subcapitate or congested-cymose, peduncle ca. 5 cm long, head or branched por-

Psychotria 733

tion 1–2.5 × 1–2 cm, bracts 7–7.5 mm long, color unknown; flowers sessile; calyx limb ca. 1 mm long, dentate; corolla in bud with color unknown, tube to 3 mm long, lobes to 2 mm long; fruit unknown. Brushy growth along streams over rocky summits of tepuis, 1900– 2100 m; Bolívar (Cerro Jaua, Cerro Sarisariñama). Endemic. Psychotria colorata (Willd. ex Roem. & Schult.) Müll. Arg. in Mart., Fl. Bras. 6(5): 372. 1881. —Cephaelis colorata Willd. ex Roem. & Schult., Syst. Veg. 5: 214. 1819. [Subg. Heteropsychotria]. —Chi-ví-du (Yekwana). Cephaelis amoena Bremek., Recueil Trav. Bot. Néerl. 31: 297. 1934. Psychotria calviflora Steyerm., Mem. New York Bot. Gard. 23: 686. 1972. Subshrub or shrub to 1.5 m tall, puberulous or glabrous; leaves 7–20 × 2.5–8 cm; petioles 2–15 mm long; stipules persistent, united around the stem, sheath 1–3 mm long, lobes 2 per side, narrowly triangular, 1– 7 mm long; inflorescence terminal, capitate, peduncle 0.5–6 cm long, capitulum 1–2 × 2–6 cm, external bracts white to purple, 15–35 mm long; flowers sessile; calyx limb 1–2 mm long, lobed; corolla white in lower 1/2 and white or usually purple above, tube 8.5–12 mm long, lobes 2–3 mm long; fruit ellipsoid, 5–6 × 3–4 mm, blue or purple; pyrenes 2, ventrally with a central longitudinal fissure, dorsally ridged. Evergreen lowland forests, riparian forests, 100–500 m; Bolívar (Macizo del Chimantá, basin of middle and upper Río Orinoco, Río Paragua), Amazonas (basin of Río Orinoco north to Puerto Ayacucho). Guyana, Suriname, French Guiana, Brazil. Steyermark separated Psychotria colorata, P. calviflora, P. rosea, and P. glabrescens Müll. Arg. of eastern Brazil based on details of pubescence and degree of fusion of the external inflorescence bracts. However, these cannot be separated consistently based on any morphological distinctions, and Steyermark (1972, 685, 686) cited the collection Williams 12992 twice, once under his new species P. calviflora and again as P. colorata. Both Psychotria colorata and P. rosea are similar to P. bracteocardia, and these have frequently been confused; the first two differ from P. bracteocardia in their inflorescence bracts glabrous or not uniformly pubescent and their acute floral bracts, versus uni-

formly pubescent and obtuse to rounded, respectively, in P. bracteocardia. Psychotria concinna Oliv., Timehri 5: 196. 1886. [Subg. Heteropsychotria]. Slender shrub or subshrub to 1.2 m tall, glabrous; leaves 1–6 × 0.5–2.8 cm; petioles 2– 8 mm long; stipules persistent, united around the stem, sheath 0.3–1 mm long, lobes 2 per side, linear, 1–2 mm long; inflorescences terminal, capitate to subcapitate or congested-cymose, nodding, peduncle 1–1.8 cm long, capitulum or branched portion 0.5–1 × 0.5–1 cm (not including corollas), bracts 1– 2 mm long; pedicels 0–3.5 mm long; calyx limb 0.8–1 mm long, dentate; corolla purple or purple and white, tube 12–15 mm long, lobes 2.5–3 mm long; fruit ellipsoid, ca. 6 × 4 mm, blue; pyrenes 2, ventrally with a central longitudinal furrow, dorsally ridged. Tepui summit and slope forests, 2000–2500 m; Bolívar (Aprada-tepui, Auyán-tepui, Ilútepui, Kukenán-tepui, Macizo del Chimantá, Murisipán-tepui, Ptari-tepui, Roraima-tepui, Yuruaní-tepui), Amazonas (Cerro Sipapo). Guyana. Psychotria concinna is similar to P. oblita, and when corollas are not present P. concinna can also be confused with P. campylopoda. Psychotria cornigera Benth., J. Bot. (Hooker) 3: 227. 1841. —Psychotria bahiensis var. cornigera (Benth.) Steyerm., Mem. New York Bot. Gard. 72: 518. 1972. [Subg. Heteropsychotria]. —Cuentica. Psychotria subcuspidata Müll. Arg. in Mart., Fl. Bras. 6(5): 261. 1888. Subshrub or shrub to 3 m tall, puberulous to glabrescent; leaves 6–21 × 2–10.5 cm; petioles 2–15 mm long; stipules persistent, united around stem, sheath 1–2 mm long, lobes 2 per side, 0.5–4 mm long, narrowly triangular; inflorescences terminal, congestedcymose to paniculate, peduncle 1–3 cm long, branched portion 2–3 × 3–5 cm, floral bracts reduced, pedicels 0–1.5 mm long; calyx limb 0.3–1 mm long, denticulate to dentate; corolla white to yellow, tube 4–6 mm long, lobes 1.5–3 mm long, abaxially with appendage 0.3–1 mm long; fruit didymous, 3–4 × 4–6 mm, blue to purple or white; pyrenes 2, ventrally with a longitudinal furrow, dorsally smooth. Evergreen lowland to lower montane

734

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forests, 50–1000 m; Bolívar (base of Auyántepui, basins of Río Paragua and Río Caroní, Río Supamo, Serranía de Imataca), Amazonas (basins of Río Orinoco and Río Ventuari). Widespread elsewhere in lowland moist Venezuela. Colombia, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. Psychotria cornigera is similar to P. acuminata (see comments about fruit color there); to P. cuspidata (see comments under P. acuminata); to P. rhodothamna Standl. of western Amazonia, which has very congested to subcapitate inflorescences, longer corollas, and larger fruits borne on thickened infructescence axes; and to P. cuspidulata K. Krause of Amazonian Peru, which has relatively larger leaves and longer corolla tubes, 10–12 mm long. Fruiting collections of P. cornigera are sometimes difficult to separate from those of P. acuminata. The circumscription of Psychotria cornigera, P. bahiensis DC., and P. acuminata here differs significantly from that of Steyermark (1972). He treated P. bahiensis as a morphologically very variable species found widely in lowland South America, and recognized two varieties of it, var. bahiensis and var. cornigera. He separated these varieties by inflorescence size, corolla length, and presence versus absence of abaxial projections on the corolla lobes. However, with more material now available, these two varieties are clearly not conspecific, and P. bahiensis itself is distinct from both var. cornigera and the plants Steyermark places in his “P. bahiensis var. bahiensis.” True P. bahiensis (Salzmann s.n., isotype, MO!; Harley 17610, MO!) is apparently endemic to Bahia, Brazil, and is clearly not closely related to these other two species. It can be distinguished from them by its leaves with the secondary and tertiary veins notably raised on the shiny upper leaf surfaces (versus plane to sometimes a little raised in these other species) and its pyrenes that are hemispherical with the dorsal surface longitudinally ridged and also finely foveolate (versus subglobose and smooth in these other species). The plants from outside Bahia that Steyermark included in his “P. bahiensis” comprise two groups: those he treated in “var. bahiensis” are here included in P. acuminata, while those he treated in var. cornigera are here treated as P. cornigera.

Psychotria coussareoides Standl., Publ. Field Columbian Mus., Bot. Ser. 7: 441. 1931. [Subg. Heteropsychotria]. Rudgea tillettii Steyerm., Mem. New York Bot. Gard. 17(1): 416. 1967. Psychotria coussareoides var. ciliata Steyerm., Mem. New York Bot. Gard. 23: 496. 1972. Shrub or small tree to 5 m tall, glabrous; leaves 8–21 × 3–7 cm; petioles 8–25 mm long; stipules persistent, united around stem, 3–5 mm long, truncate, on each side with a medial or sub-basal group of caducous glands ca. 1 mm long; inflorescence terminal, paniculate, peduncle 2.5–4 cm long, branched portion 5–10 × 3–8 cm with secondary axes 2–4 at basal node, floral bracts ca. 1 mm long, green; flowers sessile or subsessile; calyx limb ca. 1 mm long, lobed; corolla white, tube ca. 4 mm long, lobes ca. 5 mm long; fruit ellipsoid, 6–7 × 6–6.5 mm, purple; pyrenes not seen, reportedly 2. Tepui slope forests, 900– 1500 m; Bolívar (Cerro Venamo, uppermost Río Caroní, Río Cuyuní basin), Amazonas (Cerro Aracamuni, Cerro Duida, Cerro Huachamacari, Cerro Sipapo). Guyana. ◆Fig. 563. Psychotria coussareoides probably belongs to Rudgea, as shown by its stipules with groups of caducous glands on each side. It is keyed in both the Psychotria and Rudgea treatments here, but provisionally treated in Psychotria pending further study. Psychotria crocochlamys Sandwith, Kew Bull. 1939: 555. 1939. [Subg. Heteropsychotria]. Cephaelis tatei Standl., Bull. Torrey Bot. Club 56: 406. 1929, not Psychotria tatei Standl., Publ. Field Columbian Mus., Bot. Ser. 7: 460. 1931. Subshrub or shrub to 3(–4) m tall, glabrous; leaves 7.5–21 × 2.5–9.2 cm; petioles 5– 27 mm long; stipules persistent or sometimes fragmenting below apical nodes, united around stem, sheath 1–4 mm long, lobes 2 per side, ligulate, 2–5 mm long; inflorescence terminal, corymbiform, peduncle 2–9.5 cm long, branched portion 1–4 × 4–6.5 cm, external bracts 12–23 mm long, orange to yellow or red; flowers sessile; calyx limb ca. 0.8 mm long, undulate to dentate; corolla white to yellow or orange, tube 5–9 mm long, lobes 1.5–2 mm long; fruit ellipsoid, ca. 6 × 5 mm, white; pyrenes 2, ventrally with a shallow slender central longitudinal fissure, dorsally

Psychotria 735

ridged. Upland and tepui slope forests, 800– 1600 m; Bolívar (common in basin of upper Río Caroní and Río Cuyuní). Guyana. ◆Fig. 571. Steyermark noted that Psychotria crocochlamys is a very handsome shrub. It is similar to P. hemicephaelis, P. maturacensis, P. vasivensis, and P. tepuiensis. Psychotria cupularis (Müll. Arg.) Standl., Publ. Field Columbian Mus., Bot. Ser. 8: 210. 1930. —Mapouria cupularis Müll. Arg., Flora 59: 459, 465. 1876. [Subg. Psychotria]. Shrub or tree to 5 m tall, glabrous; leaves 7.5–18 × 3–8.5 cm; petioles 0.5–2.5 cm long; stipules caducous, interpetiolar, ligulate to elliptic, obovate, or subcircular, 3–14 mm long, obtuse to rounded; inflorescence terminal, paniculate, peduncle 1.8–7 cm long, branched portion 1–5 × 2–7 cm, floral bracts reduced, pedicels 0.5–2 mm long; calyx limb 0.7–2 mm long, subtruncate; corolla white, tube 3–6.5 mm long, lobes 3–4 mm long; fruit ellipsoid, 5–7 × 3.5–5 mm, red; pyrenes 2, dorsally ridged to nearly smooth. Rain forests, riparian forests, mixed montane forests, 300–1500 m; Delta Amacuro (Serranía de Imataca), Bolívar (basin of Río Cuyuní and upper Río Caroní west locally to Río Caura), Amazonas (headwaters of Río Orinoco near Brazilian frontier). Guyana, Suriname, French Guiana, Brazil. Psychotria cupularis is recognizable by its usually oblanceolate to obovate leaves, its well-developed and usually rounded stipules, its inflorescences with the secondary axes usually 4 in subequal pairs at least at the basalmost node, and its relatively large fruits. It is similar to P. mapourioides, and these may eventually be shown to intergrade but are provisionally recognized here. Psychotria deflexa DC., Prodr. 4: 510. 1830. [Subg. Heteropsychotria]. —Alatrique peludo, Pata de tintín. Subshrub or low shrub to 1(–3) m tall, puberulous to glabrous; leaves 8–19 × 2.5–7 cm; petioles 4–8 mm long; stipules persistent, united around stem, sheath 1–1.5 mm long, lobes 2 per side, narrowly triangular, 2– 6 mm long; inflorescence terminal, paniculate, peduncle 2.5–5.5 cm long, branched portion 4–7 × 2.5–5 cm, floral bracts reduced; flowers sessile; calyx limb ca. 0.2 mm long,

undulate; corolla white, tube 2–3 mm long, lobes 1–1.2 mm long; fruit subglobose to oblate, ca. 3 × 3.5 mm, white or purple; pyrenes 2, ventrally with a deep central cavity, dorsally shallowly ridged and markedly alveolate. Riparian forests, evergreen lowland to lower montane forests, 100–900 m; Delta Amacuro (near El Palmar), Bolívar (upper Río Caroní, Río Ichún, basin of Río Paragua, Serranía de Imataca), Amazonas (basin of Río Negro, Río Orinoco, and Río Ventuari). Aragua, Distrito Federal, Falcón, Lara, Mérida, Miranda, Portuguesa, Sucre, Táchira, Trujillo, Yaracuy, Zulia; Mexico, Antilles, Central America, Colombia, Guyana, Suriname, French Guiana, Ecuador, Brazil, Bolivia, Paraguay. The plants of Psychotria deflexa in the flora area and the Guianas show a larger range of leaf and inflorescence sizes than plants in other regions. Psychotria deflexa has long been confused with P. patens Sw., a separate species from the Greater Antilles, and has also been incorrectly called P. flexuosa Willd. Psychotria deflexa is similar to and has been confused in the flora area with P. paniculata, which can be distinguished by its emarginate to shortly lobed, interpetiolar stipules; with P. subundulata, which can be distinguished by its secondary leaf veins that extend to unite with the margins; with P. berteroides Wernham of western Amazonia, which can be distinguished by its inflorescences with the secondary axes branching dichotomously once into two linear, unbranched axes; to P. argoviensis Standl. of Goiás; and to P. araguana Standl., a species of premontane forests at 800–1500 m in the Coastal Cordillera. (Psychotria berteroides has often been incorrectly called P. loretensis Standl.; however, this latter name is a synonym of P. longicuspis. Psychotria berteroides has also been confused by Dwyer with P. berteroana, probably because of the similarity in the names; P. berteroides has pyrenes with a markedly concave but smooth ventral face, versus plane with a central invagination in P. berteroana.) Steyermark recognized four subspecies of Psychotria deflexa, which are here treated as three distinct species; the fourth, subsp. cubensis Steyerm., is confined to Cuba and western Hispaniola and has not been studied here. These three species share pyrenes with an alveolate dorsal surface, inflorescences

736

R UBIACEAE

with the secondary axes branched dichotomously once into two usually unbranched axes, and corollas with tubes 2–3.5 mm long. However, P. campyloneura differs from the other species in its fruits that are usually a bit larger, its corollas on the larger end of this size range, its flowers frequently arranged in dichotomous cymules rather than on secund axes, and its pyrenes with the ventral face concave and smooth. Psychotria venulosa differs from the other species in its stipules with the sheath elongating with age, so that on older nodes the lobes are inserted below its top (as in P. lupulina), and its leaves with the secondary veins usually subparallel and raised on the upper leaf surface. The pyrenes of P. venulosa are similar to those of P. deflexa, or sometimes the central cavity is closed and appears as a longitudinal fissure, and occasional specimens have pyrenes with the ventral face smooth (e.g., Nee 30584, NY); more than one species may be included here, or there may be more infraspecific variation in pyrene morphology in these plants than has previously been appreciated. Psychotria deinocalyx Sandwith, Kew Bull. 1939: 555. 1939. [Subg. Heteropsychotria]. Shrub to 4 m tall, glabrous; leaves 7.5–20 × 2.5–6.5 cm; petioles 7–20 mm long; stipules fragmenting below apical nodes, united around stem, 2–4 mm long, truncate; inflorescences terminal, capitate, sessile or subsessile, capitulum subglobose, bracts reduced; flowers sessile; calyx limb 2–3 mm long, truncate to undulate; corolla white, tube 3.5–5 mm long, lobes 1.5–2 mm long; fruit ellipsoid to subglobose, 6–7 × 5.5–6.5 mm, red; pyrenes 2, ventrally plane, dorsally shallowly ridged. Semideciduous upland forests, 400–900 m; Bolívar (region of Río Icabarú and Río Hacha). Guyana, Suriname, Brazil. This species will be included in the segregate genus Margaritopsis when it is separated from Psychotria. Psychotria deinocalyx is similar to P. podocephala and P. pallidinervia. It is also similar to P. moroidea Steyerm. of French Guiana, which differs in its shortly pedunculate inflorescences, calyx limb ca. 1 mm long, and stipules with aristas 1–3 mm long. Plants of P. deinocalyx from Brazil may have peduncles to 4 mm long, but

the Venezuelan plants have peduncles to only about 1 mm long. Psychotria duidana Standl., Publ. Field Columbian Mus., Bot. Ser. 7: 443. 1931. [Subg. Heteropsychotria]. Shrub to 1.5 m tall, puberulous to glabrous; leaves 6.5–10 × 2–3.7 cm; petiole 4–10 mm long; stipules persistent, united around stem, sheath 0.5–2 mm long, lobes 2 per side, narrowly triangular, 1.5–7 mm long; inflorescence terminal, corymbiform, peduncle 5–7.5 cm long, branched portion 1.5–2.5 × 1.5–2.5 cm, bracts 9–12 mm long, apparently green or possibly white; flowers sessile; calyx limb ca. 0.8 mm long, dentate; corolla white, tube 9–11 mm long, lobes 3–3.5 mm long; fruit unknown. Tepui summit forests, ca. 1800 m; Amazonas (summit of Cerro Duida). Endemic Psychotria duidana is known only from the type. It is similar to P. hemicephaelis, P. oblita, and in particular P. ayangannensis Steyerm. of Guyana. In his original description, Steyermark contrasted P. ayangannensis with P. oblita, but it is more similar to P. duidana. Psychotria duricoria Standl. & Steyerm., Fieldiana, Bot. 28: 599. 1953. [Subg. Heteropsychotria]. Shrub to 3 m tall, glabrous; leaves 6–12 × 2.3–6 cm; petioles 5–16 mm long; stipules persistent, united around stem, sheath 2–3 mm long, lobes 2 per side, triangular to ligulate, 1–2 mm long; inflorescences terminal, paniculate, peduncle 2–7.5 cm long, branched portion 5–7 × 5–8.5 cm, floral bracts 1–4 mm long, pedicels 0–4 mm long; calyx limb 1.5–4 mm long, lobed shallowly to deeply; corolla white, tube 5–9 mm long, lobes 2–4 mm long; fruit subglobose, 3–4 mm diameter, blue; pyrenes 2, ventrally with a central narrow invagination, dorsally ridged. Montane forests and tepui meadows, 1000– 1800 m. Venezuela, adjacent northern Brazil; 2 varieties, both in the flora area. Psychotria duricoria is similar to and sympatric with Psychotria everardii and was confused with it by Steyermark (1972). Psychotria everardii can be separated by its calyx limb 0.5–1 mm long. Steyermark recognized two varieties, which appear to be separable and are provisionally recognized here.

Psychotria 737

Key to the Varieties of P. duricoria 1. Calyx limb 1.5–2 mm long, on at least some flowers < 2 mm long, lobed for 1/2 of to much of its length, the lobes subequal ...................................... var. duricoria 1. Calyx limb 2–4 mm long, on at least some flowers > 2 mm long, lobed deeply, the lobes subequal to strongly unequal ...................................... var. longiloba P. duricoria var. duricoria. [Subg. Heteropsychotria]. Psychotria everardii subsp. sipapoensis Steyerm., Mem. New York Bot. Gard. 23: 540. 1972. Amazonas (Cerro Aracamuni, Cerro Duida, Cerro Marahuaka, Cerro Parú, Cerro Sipapo, Sierra de la Neblina). Adjacent Brazil. P. duricoria var. longiloba Steyerm., Mem. New York Bot. Gard. 23: 533. 1972. [Subg. Heteropsychotria]. Amazonas (Cerro Marahuaka, Cerro Yutajé, Sierra de la Neblina). Adjacent Brazil. ◆Fig. 567. Psychotria durifolia Standl., Publ. Field Columbian Mus., Bot. Ser. 7: 444. 1931. [Subg. Heteropsychotria]. Shrub to 3 m tall, glabrous; leaves 1.5–4 × 1–3 cm, subsessile or sessile; stipules persistent, united around stem, 1.5–3 mm long, broadly triangular, with 1 lobe on each side, narrowly triangular, 1–2 mm long; inflorescence pseudoaxillary and/or axillary, cymose or congested-cymose, peduncle 1.5–5 cm long, branched portion 1–2 × 1.5–2 cm, floral bracts 1–2.5 mm long; flowers sessile; calyx limb 1–1.2 mm long, dentate; corolla white to greenish yellow, tube ca. 3 mm long, lobes ca. 1.5 mm long; fruit obpyriform, ca. 3 × 3.5 mm, color unknown; pyrenes 2, ventrally plane, dorsally ridged. Tepui slope and summit forests, 1200–2100 m; Amazonas (Cerro Duida). Endemic. ◆Fig. 566. Psychotria durifolia is similar to P. phelpsiana and P. carrenoi; see comments about these under P. carrenoi. Psychotria durifolia is separated from these by its distinctive leaves, which are broadly elliptic to suborbicular, stiff, thick-textured, and rounded to cordulate at the base, with the secondary leaf veins usually extending unbranched to unite with the margins.

Psychotria egensis Müll. Arg., Flora 59: 542, 545. 1876. [Subg. Heteropsychotria]. Shrub or small tree to 3 m tall, glabrous; leaves 8.5–16 × 2–7 cm; petioles 3–10 mm long; stipules persistent, united around the stem, sheath 0.3–1 mm long, lobes 2 per side, narrowly triangular, 1–3 mm long; inflorescence terminal, subcapitate to paniculate with capitula 3(5), peduncle 2–3 cm long, capitulum or branched portion 1–3 × 1.5–2.5 cm, bracts subtending the capitula 5–13 mm long, remaining bracts 2–6 mm long; flowers sessile; calyx limb 1–1.5 mm long, lobed; corolla white, tube ca. 3.5 mm long, lobes ca. 1.5 mm long; fruit subglobose, ca. 4 × 5 mm, white or purple; pyrenes 2, ventrally with a central longitudinal furrow, dorsally ridged. Río Negro caatinga, riparian, and lowland forests, 50–400 m; Amazonas (basin of Río Casiquiare, Río Negro, Río Orinoco, and Río Ventuari, Río Cunucunuma, San Simon de Cocuy northeast to Río Parú). Zulia; Colombia, Amazonian Brazil. Psychotria egensis can be recogized by its leaves with the secondary veins extending unbranched to unite with the thickened margins together with its congested inflorescences usually having each capitulum subtended by a single well-developed bract. This species is similar to Psychotria schomburgkii, P. lourteigiana, and P. phaneroloma. Psychotria ernestii K. Krause, Verh. Bot. Vereins Prov. Brandenburg 50: 109. 1908. [Subg. Psychotria]. Palicourea elliptica Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 3: 369. 1818 [1819]. Shrub or small tree to 4 m tall, densely strigillose or hirtellous to glabrescent; leaves 7–27 × 2–10 cm; petioles 3–10 mm long; stipules caducous, interpetiolar, ligulate to ovate or obovate, 6–20 mm long, obtuse to rounded; inflorescences terminal or often displaced to pseudoaxillary especially as fruits mature, paniculate, peduncles 1–7 cm long, branched portion 5–10 × 6–14 cm, floral bracts 0.3–0.5 mm long, pedicels 0–1 mm long; calyx limb ca. 0.5 mm long, subtruncate; corolla white, tube ca. 1.5 mm long, lobes ca. 1.5 mm long; fruit ellipsoid, 4–4.5 × 3–4 mm, red; pyrenes 2, dorsally ridged. Riparian forests, ca. 50 m; Bolívar (Ciudad Bolívar). Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia.

738

R UBIACEAE

Psychotria ernestii can generally be recognized by its relatively large stipules that are usually ridged in the basal portion, its inflorescences with the secondary axes usually 4 in unequal pairs at least at the basalmost node, and its relatively small flowers and fruits. This species was not well characterized by Steyermark (1972; 1974), and thus has often been confused with other Psychotria species. Psychotria everardii Wernham, J. Bot. 52: 316. 1914. [Subg. Heteropsychotria]. Psychotria cacuminis Standl., Publ. Field Columbian Mus., Bot. Ser. 7: 436. 1931. Psychotria vernicifolia Standl., Publ. Field Columbian Mus., Bot. Ser. 7: 462. 1931. Psychotria phaneroneura Standl., Publ. Field Columbian Mus., Bot. Ser. 7: 455. 1930. Psychotria everardii subsp. neblinensis Steyerm., Mem. New York Bot. Gard. 23: 540. 1972. Shrub to 2 m tall, puberulous or pilosulous to glabrous; leaves 4–15 × 1.5–8 cm; petioles 1–25 mm long; stipules persistent, united around stem, sheath 1–3 mm long, lobes 2 per side, triangular, 1–3 mm long; inflorescence terminal, paniculate, peduncle 1–4.5 cm long, branched portion 1–6 × 1–6 cm, floral bracts 0.5–1.5 mm long, apparently green, pedicels 0–3 mm long; calyx limb 0.5–1 mm long, dentate; corolla white or cream flushed with pink, tube 5–10 mm long, lobes 1.5–6 mm long; fruit ellipsoid to subglobose, 5–6 × 5–6 mm, purple to black; pyrenes 2, ventrally with a central longitudinal fissure, dorsally ridged. Upper slopes and summits of tepuis, savannas, Bonnetia roraimae thickets, 1200–2500 m; Bolívar (common on all the higher sandstone mountains), Amazonas (Cerro Autana, Cerro Duida, Cerro Huachamacari, Cerro Marahuaka, Cerro Sipapo, Sierra de la Neblina). Guyana. ◆Fig. 568. Psychotria everardii is locally common and morphologically variable, particularly in leaf and inflorescence sizes, similarly to many cloud forest Rubiaceae. It is similar to Psychotria duricoria, which can be separated by its secondary inflorescence axes that are dichotomously branched once producing two elongated, secund axes with the flowers borne singly or in pairs along them, versus

branched to several orders with the flowers borne in branched cymules in P. everardii. Psychotria everardii is also similar to Palicourea wurdackiana; see comments under that species. Steyermark (1967) synonymized several of Standley’s species, each of which was based on a different Tate collection. Standley noted in his descriptions of these that several were very similar to P. everardii, and also that all of these are arguably as well classified in Palicourea as Psychotria. Steyermark recognized four subspecies of P. everardii. His subsp. sipapoensis is here treated as a synonym of P. duricoria, and his subsp. tatei is here treated as a separate species. Steyermark considered shape of the calyx lobes (i.e., rounded to obtuse in subsp. everardii versus subacute to acute or acuminate in subsp. neblinensis) to be taxonomically informative, although this feature typically varies in Psychotria species. These two subspecies were reported as sympatric by Steyermark, and apart from calyx lobe shape subsp. neblinensis is separable from subsp. everardii only by details of the pubescence on the outer surface of the corolla lobes (Steyermark 1974, 541). This latter character also varies, and these subspecies are not recognized here. Psychotria fanshawei (Standl.) Steyerm., Mem. New York Bot. Gard. 23: 662. 1972. —Cephaelis fanshawei Standl., Bull. Torrey Bot. Club 75: 574. 1948. [Subg. Heteropsychotria]. Shrub or small tree to 4 m tall, hirsute; leaves 5.5–10 × 2.5–4.5 cm; petioles 5–10 mm long; stipules persistent, united around the stem, sheath 1.5–3 mm long, lobes 2 per side, narrowly triangular, 1.5–4 mm long; inflorescences terminal, capitate, peduncle 0–3 mm long, capitulum ca. 2 × 3 cm, external bracts orange to red, 17–30 mm long; flowers sessile; calyx limb 10–11 mm long, shortly dentate; corolla white, tube 25–31 mm long, lobes 10–13 mm long; fruit subglobose, 8–13 mm diameter, blue; pyrenes 2, ventrally with a shallow longitudinal invagination, this generally broadest near apex, dorsally ridged. Tepui slope forests, 1000–1300 m; Bolívar (Ilú-tepui). More common in adjacent Guyana. Psychotria fanshawei is a handsome species in flower and fruit. It is similar to several

Psychotria 739

species of the Guianas, notably P. aetantha (Sandwith) Steyerm. and P. potaroensis (Sandwith) Steyerm., but it resembles no other species known from the flora area. Psychotria franquevilleana Müll. Arg. in Mart., Fl. Bras. 6(5): 325. 1881. [Subg. Heteropsychotria]. Shrub to 2 m tall, glabrous; leaves 17–26 × 8–8.5 cm; petioles ca. 30 mm long; stipules apparently persistent, interpetiolar or perhaps shortly united around stem, ovate to ovate-oblong, 7–8 mm long, shallowly emarginate; inflorescence terminal, corymbiform to paniculate, peduncle 8.5–10 cm long, branched portion ca. 3 × 5 cm, bracts 4–7.5 mm long, white, pedicels ca. 0.5 mm long; calyx limb ca. 0.7 mm long, truncate to undulate; corolla white to cream in bud, no measurements available; fruit subglobose-ovoid, ca. 8 × 6 mm, color unknown; pyrenes 2, ventral face not described, dorsally ridged. Evergreen lowland forests, 100–200 m; Amazonas (Río Pasimoni). Amazonian Colombia. Psychotria franquevilleana probably will be included in the segregate genus Carapichea Aubl. when it is separated from Psychotria, though the limits of Carapichea will have are not yet entirely clear. Psychotria franquevilleana is apparently similar to P. vasivensis and P. maturacensis, but the identity of P. franquevilleana is also not clear. Psychotria glandulicalyx Steyerm., Mem. New York Bot. Gard. 23: 536. 1972. [Subg. Heteropsychotria]. Psychotria glandulicalyx subsp. glandulicalyx var. densipubens Steyerm., Mem. New York Bot. Gard. 23: 538. 1972. Slender shrub or subshrub to 3.5 m tall, puberulous to pilosulous, hirtellous, or glabrescent; leaves 2–7.5 × 0.5–3 cm; petioles 2– 9 mm long; stipules persistent, united around stem, sheath 1–2 mm long, lobes 2 per side, triangular, 0.5–2 mm long; inflorescence terminal, paniculate, peduncle 0.7–2.5 cm long, branched portion 1–2.5 × 1–2 cm, floral bracts 0.5–1 mm long, apparently green, pedicels 0–4 mm long; calyx limb 0.5– 1 mm long, dentate; corolla white or yellow sometimes flushed with pink, tube 5–9 mm long, lobes 1.5–3 mm long; fruit subglobose, ca. 3 mm diameter, golden yellow to yelloworange; pyrenes 2, ventrally with a central

longitudinal fissure, this sometimes widened in the middle, dorsally ridged. Tepui slopes and summits, granitic slopes, 1000–2100 m; Bolívar (Auyán-tepui, Cerro Sarisariñama, Macizo del Chimantá), Amazonas (Cerro Aratitiyope, Cerro Yaví, northwest of headwaters of Río Orinoco, Sierra de la Neblina). Guyana, adjacent Brazil. Psychotria glandulicalyx is similar to P. everardii in general aspect. The yellow fruit color is highly unusual, particularly in combination with the infructescence branches said by some collectors to be purple. Steyermark (1972) recognized several infraspecific taxa of Psychotria glandulicalyx, none of which are recognized here. Psychotria glandulicalyx subsp. camaniensis Steyerm. (1972, 538) is also excluded here from P. glandulicalyx. Psychotria gracilenta Müll. Arg., Flora 59: 545. 1876. [Subg. Heteropsychotria]. Psychotria brachybotrya Müll. Arg. in Mart., Fl. Bras. 6(5): 327. 1881. Shrub or subshrub to 1.5 m tall, glabrous; leaves 7–17 × 2.5–7.5 cm; petioles 2–10 mm long; stipules persistent, united around stem, sheath 0.5–1 mm long, lobes 2 per side, narrowly triangular, 2–6 mm long; inflorescences terminal, capitate or subcapitate to congested-cymose, peduncle 0.5–1 cm long, capitulum or branched portion 1–1.8 × 1–1.8 cm, bracts 2.5–6 mm long, green to pale green; flowers sessile; calyx limb 0.3–0.5 mm long, dentate; corolla white, tube 1.5–3.5 mm long, lobes ca. 1 mm long; infructescences cymose to paniculate, to 3.5 × 4.5 cm; fruit ellipsoid, 4–5 × 4–5 mm, purple-black; pyrenes 2, ventrally with a central longitudinal fissure, dorsally ridged. Semideciduous to evergreen lowland and lower montane forests, 50–600 m; Delta Amacuro (Caño Güiniquina), Bolívar (basins of Río Caroní, Río Caura, and Río Paragua), Amazonas (basins of Río Orinoco and Río Negro). Barinas, Falcón, Lara, Táchira, Zulia; Central America, Colombia, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. Psychotria gracilenta is similar to P. hoffmannseggiana (particularly when in flower), P. platypoda, and P. casiquiaria (to the latter particularly when in fruit); see comments under P. casiquiaria about the inflorescence and infructescence forms of these species.

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R UBIACEAE

Psychotria guanchezii Steyerm., Ann. Missouri Bot. Gard. 74: 109, fig. 9. 1987. [Subg. Heteropsychotria]. Subshrub or shrub to 1.5 m tall, puberulous to glabrous; leaves 4–9.5 × 0.8–2 cm; petioles 3–9 mm long; stipules deciduous after distalmost nodes, interpetiolar, sheath 2– 2.5 mm long, lobes 2 per side, narrowly triangular, ca. 3 mm long; inflorescences terminal, corymbiform, peduncle 4–4.3 cm long, branched portion 1–18 × 1–2 cm, floral bracts 2.5–3 mm long; flowers sessile; calyx limb ca. 1 mm long, dentate; corolla white, tube 2.2–3.5 mm long, lobes 1.7–2.5 mm long; fruit unknown. Riparian forests, 100–200 m; Amazonas (Río Sipapo). Endemic. Psychotria hemicephaelis Wernham, J. Bot. 52: 314. 1914. —Cephaelis hemicephaelis (Wernham) Steyerm., Mem. New York Bot. Gard. 17(1): 426, fig. 41. 1967. [Subg. Heteropsychotria]. Subshrub or shrub to 3 m tall, glabrous; leaves 3–11 × 1–4 cm; petioles 1–10 mm long; stipules persisting at least on apical nodes, interpetiolar or sometimes shortly united around stem, sheath 2–3 mm long, lobes 2 per side, narrowly triangular, closely set, 1.5–2.5 mm long; inflorescence terminal or sometimes displaced to pseudoaxillary, capitate to subcapitate, peduncle 1–4 cm long, capitulum or branched portion 1–2 × 2–2 cm, bracts maroon to bright carmine red, 8–13 mm long; flowers sessile; calyx limb ca. 1.5 mm long, dentate; corolla white to yellow, tube 7.5–8 mm long, lobes 1.5–2 mm long; fruit ellipsoid, ca. 4 × 4 mm, purple and/or white; pyrenes 2, ventrally with a central longitudinal fissure, dorsally ridged. Tepui slope forests, 900–1300 m; Bolívar (Amaruay-tepui, basins of Río Aponguao and Río Cuyuní). Adjacent Guayana. ◆Fig. 560. Psychotria heteroneura Steyerm., Mem. New York Bot. Gard. 23: 714, fig. 91. 1972. [Subg. Heteropsychotria]. Shrub ca. 3 m tall, glabrous; leaves 7.5–11 × 1–2 cm; petioles 5–6 mm long; stipules persistent, united around stem, sheath 1.5–2 mm long, lobes 2 per side, narrowly triangular, 1.2–1.5 mm long; inflorescence and flowers unknown; infructescence displaced to pseudoaxillary, paniculate to corymbiform, peduncle 1.9–3.1 cm long, branched portion 4–7 × 6–8 cm, floral bracts 2–4 mm long, ap-

parently green; flowers sessile; calyx limb ca. 1 mm long, lobed; fruit ellipsoid, ca. 6 × 6 mm, white; pyrenes 2, said to have a central invagination ventrally, dorsally ridged. Tepui slope forests, 1300–2000 m; expected in Amazonas. Brazil (Amazonas: Serra da Neblina). Psychotria heteroneura is similar to P. speluncae of the flora area and to P. hazenii Standl. of the Andes. Steyermark distinguished P. heteroneura from P. speluncae primarily by its secondary leaf veins that are better developed than the intersecondary veins (versus developed to almost similar degrees in P. speluncae); additionally, the leaves of P. heteroneura have quite well-developed tuft domatia in the abaxial vein axils (versus none or small in P. speluncae). Steyermark placed P. heteroneura in his section Oppositiflora, but unlike the other species he placed here, the infructescences of P. heteroneura are pseudoaxillary rather than truly axillary. Section Oppositiflorae is here treated as a separate genus, Ronabea, which includes only one of the species that Steyermark placed in that section; the other species he placed there (P. heteroneura, P. speluncae, and P. vellosiana) are all here still classified in Psychotria. Psychotria hoffmannseggiana (Willd. ex Roem. & Schult.) Müll. Arg. in Mart., Fl. Bras 6(5): 336. 1881. —Cephaelis hoffmannseggiana Willd. ex Roem. & Schult., Syst. Veg. 5: 214. 1819. [Subg. Heteropsychotria]. —Seréu-ka-yek (Pemón). Cephaelis dichotoma Willd. ex Roem. & Schult., Syst. Veg. 5: 214. 1819, nom. illeg., not Psychotria dichotoma Rudge 1805. Cephaelis rubra Willd. ex Roem. & Schult., Syst. Veg. 5: 214. 1819. Psychotria barbiflora DC., Prodr. 4: 509. 1830. Cephaelis (?) microcephala Miq., Linnaea 28: 748. 1844, nom. illeg., not Cephaelis microcephala Humb. & Bonpl. ex Roem. & Schult. 1819. —Psychotria microcephala Miq., Stirp. Surinam. Select. 180. 1850 [1851]. Psychotria erythrophylla Müll. Arg., Flora 59: 542, 546. 1876. —Psychotria hoffmannseggiana var. erythrophylla (Müll. Arg.) Steyerm., Mem. New York Bot. Gard. 23: 607. 1972. Subshrub or shrub to 2 m tall, puberulous to glabrous; leaves 3.5–13 × 1–6 cm; petioles

Psychotria 741

2–8 mm long; stipules persistent, united around stem, sheath 0.5–2 mm long, lobes 2 per side, narrowly triangular, 0.5–2.5 mm long; inflorescence terminal, capitate to subcapitate, peduncle 0.2–1.2 cm long, head 0.5–1 × 0.8–2 cm, bracts 3.5–12 mm long, green; flowers sessile; calyx limb 0.3–0.7 mm long, dentate; corolla white, tube 2–5 mm long, lobes 1.5–3 mm long; fruit subglobose, 4–5 × 4–5 mm, purple-black; pyrenes 2, ventrally with a central longitudinal fissure, dorsally ridged. Riparian forests, seasonally flooded forests, forested igneous and sandstone outcrops, tree islands in shrubby savannas, 50–1700 m; common throughout Delta Amacuro, Bolívar, and Amazonas. Widespread elsewhere in lowland Venezuela; Central America, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. Psychotria hoffmannseggiana is one of the most commonly collected species of Psychotria in the flora area. The circumscription here of P. hoffmannseggiana differs significantly from that of Steyermark (1972; 1974) in treating P. barbiflora as a synonym. Steyermark recognized three allopatric varieties of this variable species, var. celsa Steyerm. of east-central and eastern Brazil, var. tribracteata (Wright ex Griseb.) Steyerm. of Cuba, and var. hoffmannseggiana, which is found in the rest of the range of this species. Within var. hoffmannseggiana he recognized two forms, both reported from the flora area, and two forms, one reported from the flora area and f. pubescens Steyerm. from Guyana. These varieties and forms were separated by leaf shape, details of pubescence, and, in var. erythrophylla, peduncle length, all continuously variable features; consequently these are not recognized here. Steyermark (1972) reported the peduncle of var. erythrophylla as 15–40 mm long, which is quite long and it is questionable that this variety is even conspecific with P. hoffmannseggiana. Psychotria horizontalis Sw., Prodr. 44. 1788. [Subg. Psychotria]. —Cuentica, Fruta de pajarito, Guayabita. Psychotria glaucescens Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 3: 358. 1818 [1819]. —Psychotria horizontalis var. glaucescens (Kunth) Steyerm., Mem. New York Bot. Gard. 23: 472. 1972.

Psychotria horizontalis var. psilophylla Steyerm., Mem. New York Bot. Gard. 23: 472. 1972. Subshrub or shrub to 2 m tall, hirtellous to glabrous; leaves 4–9 × 1.5–4 cm; petioles 3–6 mm long; stipules caducous, interpetiolar, triangular to lanceolate, 3–6 mm long, acute; inflorescences terminal, paniculate, peduncles 2–5 cm long, branched portion 1.5–2.5 × 2–4 cm, floral bracts 0.5–1 mm long, pedicels 0–2 mm long; calyx limb 1.5– 2.5 mm long; corolla white, tube 2.5–3.5 mm long, lobes 1.5–2 mm long; fruit ellipsoid, 4–5 × 3–3.5 mm, red; pyrenes 2, dorsally ridged. Riparian forests, semideciduous to evergreen lowland forests, 50–300 m; Bolívar (Altiplanicie de Nuria, Río Caroní, Río Orinoco, Río Paragua, near Río Parhueña, Serranía de Imataca), Amazonas (upper Río Orinoco). Aragua, Carabobo, Distrito Federal, Falcón, Miranda, Sucre, Yaracuy; Mexico, Central America, the Antilles, Colombia, Guyana, Ecuador, Peru, Brazil. Psychotria horizontalis is generally recognizable by its triangular acute stipules, its inflorescences with the secondary axes usually 4 in unequal pairs at least at the basalmost node, and its relatively long, deeply lobed calyx limb. It is similar to Psychotria borjensis. Steyermark (1972) recognized three varieties of this species based on variations in distribution of the pubescence. However, pubescence varies widely throughout the range of this species and without apparent pattern, and does not seem taxonomically informative. Psychotria horizontalis is frequently collected in seasonally dry vegetation throughout Venezuela. Psychotria humboldtiana (Cham.) Müll. Arg. in Mart., Fl. Bras. 6(5): 333. 1881. —Cephaelis humboldtiana Cham., Linnaea 4: 136. 1829. [Subg. Heteropsychotria]. —Oreja de picure. Psychotria humboldtiana var. ornata Müll. Arg. in Mart., Fl. Bras. 6(5): 334. 1881. —Cephaelis humboldtiana var. ornata (Müll. Arg.) Standl., Publ. Field Columbian Mus., Bot. Ser. 7: 427. 1931. Cephaelis humboldtiana var. caudata Müll. Arg. in Mart., Fl. Bras. 6. (5): 334. 1881. Subshrub or shrub to 3 m tall, glabrous; leaves 6.5–17 × 1–5 cm; petioles 6–22 mm

742

R UBIACEAE

long; stipules persistent, united around stem, sheath 1.5–2 mm long, lobes 2 per side, narrowly triangular, 1–4 mm long; inflorescence terminal, subcapitate to corymbiform, peduncle 1–3.5 cm long, capitulum or branched portion 2.5–5 × 2.5–6 cm, bracts 9– 20 mm long, pink to magenta; flowers sessile; calyx limb 2.5–4 mm long, lobed; corolla white, tube 18–21 mm long, lobes 3.5–6 mm long; fruit ellipsoid, ca. 7 × 3 mm, blue; pyrenes 2, ventrally with a central longitudinal fissure, dorsally ridged. Scrub forests, low forests on white sand, flooded forests, borders of white-sand savannas, 100–800 m; Bolívar (basin of Río Paragua at Río Ichún and base of Cerro Guaiquinima), Amazonas (basins of Río Baría, Río Casiquiare, Río Guainía, Río Negro, and Río Orinoco). Amazonian Colombia and Brazil. ◆Fig. 573. Psychotria humboldtiana is one of the most handsome species of Psychotria. It is distinctive because of its leaves with the venation prominulous on both surfaces and the lower surface often shiny, its relatively large colorful inflorescence bracts, and the relatively long showy corollas. Psychotria imthurniana Oliv., Timehri 5(2): 196. 1886. [Subg. Heteropsychotria]. Psychotria auyantepuiensis Standl., Brittonia 3: 193. 1939. Psychotria ptariensis Standl. & Steyerm., Fieldiana, Bot. 28: 604. 1953. —Psychotria imthurniana f. ptariensis (Standl. & Steyerm.) Steyerm., Mem. New York Bot. Gard. 23: 526. 1972. Subshrub or shrub to 2.5 m tall, glabrous; leaves 3–9 × 0.5–3 cm, sessile or subsessile; stipules persistent, united around the stem, sheath 0.5–1 mm long, lobes 2 per side, narrowly triangular, 1–2 mm long; inflorescences terminal, paniculate, peduncle 1–3 cm long, branched portion 1.5–3 × 1.5–3 cm, floral bracts 0.5–1.5 mm long, pedicels 0–2 mm long; calyx limb ca. 0.5 mm long, dentate; corolla yellow, cream, white, and/or purple, tube 3–3.5 mm long, lobes 1.5–2 mm long; fruit ellipsoid, ca. 3.5 × 3.5 mm, purple to blue; pyrenes 2, ventrally with a longitudinal central fissure, dorsally ridged. Tepui slope and summit forests, 1300–2220 m; Bolívar (Auyán-tepui, Ptari-tepui, basins of Río Cuyuní and Río Caroní). Adjacent Guyana. ◆Fig. 574.

The subsessile leaves with their rounded to cordulate bases and the general slender habit of the plant are distinctive for Psychotria imthurniana. It is similar to P. amplectans. Psychotria iodotricha Müll. Arg. in Mart., Fl. Bras. 6(5): 375. 1881. [Subg. Heteropsychotria]. —Boyuyo, Pata morrocoy, Peludita. Psychotria atricapilla Bremek., Recueil Trav. Bot. Néerl. 31: 296. 1934. —Psychotria iodotricha subsp. atricapilla (Bremek.) Steyerm., Mem. New York Bot. Gard. 23: 646. 1972. Subshrub or low shrub to 2 m tall, hirsute to villous or occasionally glabrescent; leaves 4.5–11 × 1–4.5 cm; petioles 1–5 mm long; stipules persistent, united around stem, sheath 1–1.5 mm long, lobes 2 per side, narrowly triangular, 3–7 mm long; inflorescence terminal, capitate, sessile, capitula 0.5–1.5 cm diameter, bracts 4–20 mm long; flowers sessile; calyx limb 3.5–8 mm long, deeply lobed; corolla white, tube 9–12 mm long, lobes 3–3.5 mm long; fruit subglobose, 3–4 × 3–4 mm, purple to blue; pyrenes 2, ventrally with a longitudinal furrow, dorsally shallowly ridged. Evergreen lowland to montane forests, 100–1400 m; Delta Amacuro (Serranía de Imataca), Bolívar (Macizo del Chimantá, Ptari-tepui, Río Aponguao, upper Río Caroní, basin of Río Paramichi, Serranía de Imataca), Amazonas (basins of Río Casiquiare, Río Negro, and Río Orinoco). Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia. ◆Fig. 577. Psychotria iodotricha is similar to P. medusula, P. variegata, and P. trichophora Müll. Arg. of Brazil, and to P. trichophoroides Müll. Arg. if that is distinct from P. trichophora. Psychotria trichophora can be separated by its subsessile inflorescences with four lanceolate external bracts, versus bracts numerous and very narrow in P. iodotricha. Psychotria iodotricha is commonly collected in the flora area and rather variable morphologically, especially in pubescence, leaf size, and inflorescence size and arrangement, including number and size of the bracts and number of flowers. Steyermark recognized four subspecies of P. iodotricha based on variations in pubescence, number of flowers per head, and number of bracts per head, but the variation appears to be con-

Psychotria 743

tinuous, and these subspecies are not recognized here. Psychotria irwinii Steyerm., Mem. New York Bot. Gard. 23: 453, fig. 66. 1972. [Subg. Psychotria]. —Alatriquillo. Shrub or tree to 8 m tall, glabrous; leaves 9–21.5 × 3.5–13.5 cm; petioles 0.5–2.5 cm long; stipules caducous, interpetiolar, ligulate to ovate, 6–10 mm long, obtuse to acute; inflorescences terminal, paniculate, peduncle 3.5–9 cm long, branched portion 3.5–8.5 × 5.5–9 cm, floral bracts 0.5–1 mm long, pedicels 0–6 mm long; calyx limb 1.5–2 mm long, subtruncate; corolla white, tube 6–6.5 mm long, lobes 4–5 mm long; fruit ellipsoid to subglobose, 6–7.5 × 5–7 mm, red, with pedicels to 6 mm long; pyrenes 2, dorsally smooth to shallowly ridged. Semideciduous to evergreen lowland and lower montane forests, 200–800 m; Delta Amacuro (Serranía de Imataca), Bolívar (basins of Río Caroní, Río Cuyuní, and Río Orinoco, Serranía de Imataca). Guyana, Suriname, French Guiana, Brazil. Psychotria irwinii is generally recognizable as similar to P. mapourioides but with larger flowers. Psychotria irwinii may eventually be shown to represent only a vigorous form of that variable species. Psychotria jauaensis Steyerm., Mem. New York Bot. Gard. 23: 889. 1972. [Subg. Heteropsychotria]. Shrub or small tree to 4 m tall, glabrous; leaves 4.5–27 × 2.5–13 cm; petioles 5–20 mm long; stipules persistent, united around stem, sheath ca. 1 mm long, lobes 2 per side, 1.5–2 mm long; inflorescences terminal, congested-cymose, peduncles 0.5–3 cm long, branched portion 1–2 × 1–2 cm; floral bracts none or reduced, pedicels 0.5–1 mm long; calyx limb ca. 1 mm long, lobed to ca. 1/2 its length; corolla white to cream, pink, or purple, tube 8–12 mm long, lobes 2.5–4 mm long, with abaxial appendage 0.5–1 mm long; fruit didymous, ca. 5 × 7 mm, blue-purple to dark blue; pyrenes 2, ventral face not seen, dorsally smooth. Tepui scrub, 2100– 2200 m; Bolívar (Cerro Sarisariñama), Amazonas (Cerro Marahuaka). Endemic. ◆ Fig. 572. Psychotria jauaensis is similar to P. ceratantha; see comments about their separation there.

Psychotria kappleri (Miq.) Müll. Arg. ex Benoist, Bull. Soc. Bot. France 58: 140. 1921. —Carapichea kappleri Miq., Stirp. Surinam. Select. 181. 1850 [1851]. —Uragoga kappleri (Miq.) Pulle, Enum. Vasc. Pl. Surin. 446. 1906. —Cephaelis kappleri (Miq.) Standl., Publ. Field Columbian Mus., Bot. Ser. 4: 335. 1929. [Subg. Heteropsychotria]. Shrub to 3 m tall, glabrous; leaves 5.5–13 × 2.5–5 cm; petioles 3–11 mm long; stipules fragmenting below apical nodes, interpetiolar and shortly intrapetiolar, 1–2 mm long, broadly rounded, with 1 caducous arista to 1 mm long; inflorescences terminal, capitate, sessile, capitula 5–8 × 5–10 mm, involucral bracts 4–6 × 4–5 mm, green; flowers sessile; calyx limb 1–1.2 m long, undulate to shallowly dentate; corolla cream, tube 1–1.5 mm long, lobes ca. 1 mm long; fruit ovoid to ellipsoid, 6–8 × 4–5 mm, red to red-purple; pyrenes 2, ventrally plane, dorsally shallowly ridged. Not collected in the flora area, but expected here in wet forests between 50 and 500 m elevation. Barinas; Central America, Colombia, Guyana, French Guiana, northeastern Brazil. Psychotria kappleri will belong to the segregate genus Margaritopsis Griseb. when it is eventually separated from Psychotria. It is similar to Psychotria oblonga, and to Psychotria muscosa (Jacq.) Steyerm. of Guyana. Psychotria leiantha Steyerm., Mem. New York Bot. Gard. 23: 634, fig. 85. 1972. [Subg. Heteropsychotria]. Psychotria steinii Steyerm., Ann. Missouri Bot. Gard. 75: 345, fig. 346. 1988. Subshrub or shrub to 0.7 m tall, pilosulous or hirtellous to glabrous; leaves 2–4.7 × 1–2 cm; petioles 2–4 mm long; stipules persistent, united around the stem, sheath 0.5–1.5 mm long, lobes 2 per side, linear, 0.5–1 mm long; inflorescences terminal, congested-cymose, peduncle 0.3–1 cm long, branched portion 1–2 × 1–2.5 cm, bracts green, foliaceous, 8–13 mm long; flowers sessile; calyx limb 0.8–1.5 mm long, lobed; corolla white to lavender, tube 10–15 mm long, lobes 4–5 mm long; fruit subglobose, ca. 5.5 × 4 mm, blue; pyrenes 2, ventrally concave and smooth or perhaps with a short central slit in the basal half, dorsally ridged. Tepui scrub forests and meadows, 1900–2100 m; Amazonas (Sierra

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R UBIACEAE

de la Neblina). Brazil (Amazonas: Serra da Neblina). The foliaceous, relatively large bracts on the nodding inflorescences, the relatively large showy corollas, and the relatively small leaves are distinctive for Psychotria leiantha. The internally glabrous white corollas with straight bases are no doubt what persuaded Steyermark (1972) to place this species in Psychotria rather than Palicourea, but the flowers are larger than is usual in Psychotria. Steyermark in his description of Psychotria steinii compared it with P. duidana and P. oblita, both of which have well-developed inflorescence bracts and relatively long corollas, but did not mention P. leiantha, which is much more similar. The only difference between P. steinii and P. leiantha appears to be the smaller corolla that Steyermark described for P. steinii, 11–11.5 mm long versus 15–20 mm long in P. leiantha. However, the flowers of both the type and the paratype specimen of P. steinii appear to be rather old and wilted, so their dry size is reduced compared to the well-preserved corollas on the type specimen of P. leiantha. No other features separate these, and they are here treated as synonyms. Psychotria longicuspis Müll. Arg., Flora 59: 549. 1872. [Subg. Heteropsychotria]. Psychotria cincta Standl., Publ. Field Columbian Mus., Bot. Ser. 7: 90. 1930. Psychotria loretensis Standl., Publ. Field Columbian Mus., Bot. Ser. 7: 198. 1930. Shrub or small tree to 3.5 m tall, glabrous; leaves 12–21 × 5–9 cm; petioles 10–15 mm long; stipules persistent, united around stem, sheath 1–3 mm long, lobes 2 per side, linear, 3.5–17 mm long; inflorescences terminal, paniculate, peduncle 3.5–9 cm long, branched portion 7.5–13 × 1.5–6 cm, floral bracts 0.5–1 mm long; flowers sessile; calyx limb ca. 0.5 mm long, dentate; corolla white to yellow, tube ca. 3 mm long, lobes ca. 2 mm long; fruit ellipsoid to subglobose, 3.5–4 × 4–5 mm, becoming orange then purple-black; pyrenes 2, ventrally smoothly concave, dorsally shallowly ridged. Riparian forests, 100– 300 m; Amazonas (Río Yureba). Mérida, Zulia; Central America, Colombia, Ecuador, Peru, Brazil, Bolivia. Psychotria longicuspis can be recognized by its secondary leaf veins extending un-

branched to unite with the rather thickened margins; it is otherwise quite similar to Psychotria deflexa. Psychotria pakaraimensis Steyerm. of Guyana is similar and will key out here, but differs from P. longicuspis in its thicker-textured leaves, truncate stipules, and inflorescence axes branched only to a few orders with the flowers sessile and well separated along the axes. The inflorescences and infructescences are pendulous, and vary markedly in size. Steyermark (1974) described the fruits as yellowish white when dry, but collectors describe them in the field as orange then purple-black. This species has long been treated as Psychotria cincta, but the type collection (Spruce 2871, BR!) as well as the description of P. longicuspis show that these names are synonymous. Psychotria longicuspis is similar to P. limitanea Standl. of western Amazonia, to P. plocamipes Wernham of the Guianas, and to P. subundulata of the flora area. Psychotria lourteigiana Steyerm., Mem. New York Bot. Gard. 23: 586, fig. 81. 1972. [Subg. Heteropsychotria]. Subshrub or shrub to 1 m tall, glabrous; leaves 7–11 × 1–2.5 cm; petioles 3–10 mm long; stipules persistent, united around the stem, sheath 1–3 mm long, lobes 2 per side, narrowly triangular, 1–3 mm long, inflorescences terminal, capitate to subcapitate or congested-cymose, peduncle 2–3.5 cm long, capitulum or branched portion 0.5–1 × 1–2 cm, bracts subtending capitula 8–9 mm long, remaining bracts 2–4 mm long; flowers sessile; calyx limb 1–1.5 mm long, lobed; corolla white, tube 3–4.5 mm long, lobes 1.5–2 mm long; fruit subglobose, ca. 4 × 3 mm, white; pyrenes 2, ventrally with a central longitudinal furrow, dorsally ridged. Riparian and lowland to upland forests, 100–900 m; Bolívar (Río Orinoco and basin of Río Icabarú, near Urbana), Amazonas (Río Autana, Río Cunucunuma, Río Mawarinuma, Sierra de la Neblina). Endemic. Psychotria lourteigiana is similar to P. egensis and P. phaneroloma. The leaves are usually relatively narrow, and are distinctive because of the secondary veins that extend unbranched to unite with the thickened margin. Psychotria lupulina Benth., J. Bot.

Psychotria 745

(Hooker) 3: 230. 1841. [Subg. Heteropsychotria]. Shrub or small tree to 4 m tall, puberulous to glabrous; leaves 6–24 × 2–10 cm; petioles 0–6 mm long; stipules persistent, united around stem, sheath 1.5–3 mm long, lobes 2 per side, narrowly triangular, 1–5 mm long, inserted below top of sheath; inflorescence terminal, congested-cymose to corymbiform, peduncle 1–5 cm long, branched portion 1– 2.5 × 1.5–5.5 cm, bracts 7–25 mm long, green to white; flowers sessile; calyx limb 0.5–1.2 mm long; corolla white, tube 6–8.5 mm long, lobes 2.5–4.5 mm long; fruit subglobose to ovoid, 4–5 × 4–5 mm, purple; pyrenes 2, ventrally with a central longitudinal fissure, dorsally smooth. Riparian forests, 50–400 m. Venezuela, Colombia, Guyana, Suriname, Ecuador, Peru, Brazil, Bolivia; 2 subspecies, both in the flora area. The stipules of Psychotria lupulina are distinctive, and similar to those of only a few other species. When young they resemble those of other species of subgenus Heteropsychotria, with the lobes arising from the top of the sheath. However, as the stipules age the sheath continues to elongate beyond the bases of the lobes, typically for ca. 1 mm, so the lobes are left inserted well below the top of the sheath. Similar stipules are found in P. amplectans, P. lindenii, and P. venulosa; those of P. blakei have not been seen in good condition but may also be similar. Psychotria lupulina is quite variable morphologically, especially in bract size and shape, leaf base size and shape, and petiole length. However, even Steyermark was unable to partition this variation clearly into separate taxa. He recognized two subspecies, both reported from the flora area, two varieties, both reported from the flora area, and two forms, both from the flora area. Steyermark (1974, 633) noted that the variation in bract size is continuous between the subspecies on at least one individual collection and thus presumably within populations, so this character is not informative. His two subspecies appear distinct, though, in spite of the subtleness of their distinction, and are recognized here pending further study. Likewise, the basally obtuse to rounded, sessile to subsessile leaves that Steyermark used to distinguish var. maypurensis also vary continuously between this form and the petiolate, cuneate-based leaves

he described for var. rhodoleuca; consequently, these two varieties are not separable and are not recognized here. Key to the Subspecies of P. lupulina 1. External or basalmost inflorescence bracts oblong-obovate, elliptic-ovate, oblong-lanceolate, or lanceolate, abruptly cuspidate or acute at apex, usually broadest at the middle, mainly (1.8–) 2.5–5(–10) mm wide, usually 1.5–3.66 (–4.75) times longer than broad .............. ................................... subsp. lupulina 1. External or basalmost inflorescence bracts subulate to linear, lanceolate-linear, linear-oblanceolate, or linear-spatulate, acuminate, mainly of the same width throughout or slightly broader toward the apex or above the middle, mainly 1–2 mm wide, usually (5.5–)6– 13(–15) times longer than broad ............. ............................... subsp. rhodoleuca P. lupulina subsp. lupulina. [Subg. Heteropsychotria]. Bolívar (upper Río Caroní, Río Paragua), Amazonas (basins of Río Guainía, Río Negro, Río Orinoco, and Río Ventuari). Zulia; Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 575. P. lupulina subsp. rhodoleuca (Müll. Arg.) Steyerm., Mem. New York Bot. Gard. 72: 633. 1972. —Psychotria rhodoleuca Müll. Arg., Flora 59: 541, 545. 1876. [Subg. Heteropsychotria]. —Cafecillo, Paraparillo. Psychotria lupulina subsp. rhodoleuca var. maypurensis (Humb. & Bonpl. ex Roem. & Schult.) Steyerm., Mem. New York Bot. Gard. 23: 634. 1972. —Psychotria maypurensis Humb. & Bonpl. ex Roem. & Schult., Syst. Veg. 5: 190. 1819. Patabea alba Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 3: 375. 1818 [1819]. Psychotria persimilis Müll. Arg., Flora 59: 541, 545. 1876. Psychotria flavovirens Suess., Revista Sudamer. Bot. 7(6): 168. 1943. Psychotria lupulina subsp. rhodoleuca var. maypurensis f. pubens Steyerm., Mem. New York Bot. Gard. 23: 634. 1972. Bolívar (Cerro Cotorra, Río Cuchivero, Río Orinoco, Río Paragua basin), Amazonas (Río

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Guainía, Río Orinoco basin southeast of Puerto Ayacucho, Río Ventuari). Brazil. Psychotria maguireorum Steyerm., Mem. New York Bot. Gard. 23: 625, fig. 84. 1972. —Psychotria bracteata var. latifolia Müll. Arg. in Mart., Fl. Bras. 6(5): 313, pl. 49, fig. 2. 1881, not Psychotria latifolia Humb. & Bonpl. ex Roem. & Schult. 1819. [Subg. Heteropsychotria]. Psychotria bracteata var. tenuifolia Müll. Arg. in Mart., Fl. Bras. 6(5): 313. 1881. Psychotria bracteata var. intermedia Müll. Arg. in Mart., Fl. Bras. 6(5): 312. 1881. Subshrub or shrub to 1.5 m tall, glabrous; leaves 8–15 × 2.5–7 cm; petioles 0.3–1.2 cm long; stipules persistent on distalmost nodes, interpetiolar and sometimes shortly united around stem, sheath 2–5 mm long, lobes 2 per side, narrowly triangular, 4–10 mm long; inflorescence terminal, congested-cymose to corymbiform, peduncle 2–7 cm long, branched portion 1.5–6 × 2–5 cm, bracts 4–11 mm long; flowers sessile; calyx limb ca. 1 mm long, dentate; corolla white to pink or purple, tube 7–9 mm long, lobes 3–4.5 mm long; fruit ellipsoid, 4–5 × 4–5 mm, purple-black; pyrenes 2, ventrally with a longitudinal fissure, dorsally ridged. Seasonally flooded black-water forests, riparian forests, 50–200 m; Amazonas (basins of Río Casiquiare, Río Guainía, Río Negro, and Río Pasimoni). Guyana, Amazonian Brazil. Psychotria maguireorum is a handsome species, with the inflorescence bracts and/or axes quite colorful: pink, magenta, or redpurple to orange-maroon. This species is similar to P. capitata; see comments regarding their distinction under that species. Psychotria mapourioides DC., Prodr. 4: 509. 1830. —Mapouria guianensis Aubl., Hist. Pl. Guiane 175, pl. 67. 1775, not Psychotria guianensis Raeusch. 1797. —Psychotria nitida Willd., Sp. Pl. 1: 963. 1798, nom. superfl. illeg. —Psychotria mapuria Roem. & Schult., Syst. Veg. 5: 187. 1819, nom. illeg. superfl. [Subg. Psychotria]. Psychotria chionantha DC., Prodr. 4: 526. 1830. Shrub or tree to 15 m tall, glabrous; leaves 12–28 × 3–12 cm; petioles 0.7–4 cm long; stipules caducous, interpetiolar, ligulate to obovate, 8–23 mm long, obtuse to rounded;

inflorescences terminal, paniculate, peduncles 2.8–13 cm long, branched portion 3–10 × 4–12 cm, floral bracts ca. 0.5 mm long, pedicels 0.5–2 mm long; calyx limb 0.5–1.2 mm long, subtruncate to shallowly dentate; corolla white, tube 4–5.5 mm long, lobes 2– 4.5 mm long; fruit subglobose to ellipsoid, 5– 7.5 × 4–5 mm, red, with pedicels or stipes to 5 mm long; pyrenes 2, dorsally ridged to nearly smooth. Riparian forest, semideciduous to evergreen lowland and lower montane forests, 50–700 m; Bolívar (Río Caroní, Río Paramichi). Anzoátegui, Sucre; Trinidad, Tobago, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. Psychotria mapourioides is generally recognizable by its well-developed stipules and relatively large leaves, inflorescences, flowers, and fruits. It is similar to P. irwinii and P. yavitensis, which are recognized provisionally here pending further study but may eventually be shown to be smaller forms of P. mapourioides. Steyermark (1972; 1974) considered this species morphologically variable and recognized four varieties based largely on size ranges of the leaves and flowers and degree of development of the secondary leaf veins. Apart from var. tobagoensis (Urb.) Steyerm., these varieties were considered largely sympatric in the Guianas with var. mapourioides distributed more widely and the only variety he reported from Venezuela. Some of the specimens Steyermark treated in some of his varieties of P. mapourioides are here included in P. cupularis, and in general there is so much morphological variation in any given site within P. mapourioides that these varieties do not seem informative and are not recognized here. Psychotria maturacensis Steyerm., Mem. New York Bot. Gard. 23: 578, fig. 78. 1972. [Subg. Heteropsychotria]. Shrub or tree to 4 m tall, glabrous; leaves 13–25 × 6–14 cm; petioles 20–45 mm long; stipules deciduous below apical node, shortly united around stem, ovate, 10–13 mm long; inflorescences terminal, paniculate, peduncle 5.5–10 cm long, branched portion 3.5–8 × 3– 6.5 cm, floral bracts 5–10 mm long; flowers sessile; calyx limb 2.5–3.5 mm long, truncate to undulate; corolla white, tube 15–16 mm long, lobes ca. 10 mm long; fruit ellipsoid to subglobose, 8–13 × 6–10 mm, red-purple to

Psychotria 747

red-violet; pyrenes 2, ventrally plane, dorsally angled to shallowly ridged. Seasonally flooded riparian forests, lower montane forests, 100–300 m; Amazonas (Río Emoni, Sierra de la Neblina). Brazil (Amazonas: Serra da Neblina). Psychotria maturacensis may belong to the segregate genus Carapichea when it is separated from Psychotria, but the limits of Carapichea are not yet entirely clear. Psychotria maturacensis is similar to P. vasivensis. Psychotria medusula Müll. Arg., Flora 59: 541, 545. 1876. [Subg. Heteropsychotria]. Tapogomea glabra Aubl., Hist. Pl. Guiane 165, t. 63. 1775, not Psychotria glabra (Turrill) Fosberg 1942. —Uragoga glabra (Aubl.) Kuntze, Revis. Gen. 2: 960. 1891. Cephaelis blepharophylla Standl., Publ. Field Columbian Mus., Bot. Ser. 8: 182. 1930. —Psychotria blepharophylla (Standl.) Steyerm., Mem. New York Bot. Gard. 23: 649. 1972. Subshrub or low shrub to 1 m tall, strigillose and/or hirsute often becoming glabrescent; leaves 6–14 × 1.5–7 cm; petioles 5– 12 mm long; stipules persistent, united around stem, sheath 1–2 mm long, lobes 2 per side, narrowly triangular, 3–10 mm long; inflorescence terminal, capitate or subcapitate, peduncle 2–10 mm long, capitulum 1–2 cm diameter, bracts 6–20 mm long; flowers sessile; calyx limb 0.5–2 mm long, denticulate to dentate; corolla white, tube 10–14 mm long, lobes 1.5–3 mm long; fruit subglobose to ellipsoid, 4–5 × 4–5 mm, purple to bright blue; pyrenes 2, ventrally with a slender longitudinal furrow, dorsally shallowly ridged. Evergreen lowland forests, tall caatinga forests on white sand, 100–200 m; Amazonas (Río Autana, Río Casiquiare, Río Guainía, Río Negro). Colombia, French Guiana, Peru, Brazil. ◆Fig. 581. Psychotria medusula is similar to P. iodotricha of the flora area, and to P. adpressipilis Steyerm. and P. trichocephala Poepp. & Endl., both of Amazonian Peru. Psychotria medusula is rather variable morphologically, in particular in pubescence, leaf size, and size of the inflorescence bracts. Psychotria microbotrys Ruiz ex Standl., Publ. Field Columbian Mus., Bot. Ser. 8: 204. 1930. [Subg. Heteropsychotria].

Psychotria palicoureoides Müll. Arg. in Mart., Fl. Bras. 6(5): 305. 1881, nom. illeg., not Psychotria palicoureoides Mart. 1841. Subshrub or shrub to 3 m tall, glabrescent; leaves 14–27 × 7–13 cm; petioles 10–35 mm long; stipules generally persisting on apical nodes, interpetiolar and sometimes shortly united around stem, ovate in outline, 11–20 mm long, bilobed for 1/2–2/3 its length; inflorescences terminal, paniculate, peduncle 3.5–7 cm long, floral bracts 0.8–1.5 mm long; pedicels 0–2 mm long; calyx limb ca. 0.3 mm long, undulate to dentate; corolla white, tube 2–3.5 mm long, lobes ca. 1 mm long; fruit subglobose, 3–4 mm diameter, blue or white; pyrenes 2, ventrally with a smooth rather large central cavity, dorsally ridged and with the surface alveolate. Riparian forests, tepui slope forests, 100–1100 m; Bolívar (basins of Río Cuyuní and Río Icabarú, headwaters of Río Paramichi), Amazonas (Cerro Duida, Cerro Huachamacari, Cerro Yutajé, lower Río Ventuari, Sierra de la Neblina). Central America, Colombia, Amazonian Ecuador, Peru, Brazil, and Bolivia. Psychotria microbotrys is well marked by its relatively large, bifid, sub-interpetiolar stipule. Psychotria microdon (DC.) Urb., Symb. Ant. 9: 539. 1928. —Rondeletia microdon DC., Prodr. 4: 408. 1830. —Mapouria microdon (DC.) Bremek., Recueil Trav. Bot. Néerl. 31: 286. 1934. [Subg. Heteropsychotria]. —Boquilla. Psychotria microdon var. meridionalis Steyerm., Mem. New York Bot. Gard. 23: 446. 1972. Erect or recumbent to clambering shrub to 5 m tall, puberulous to glabrous; leaves 3– 15 × 2–8 cm; petioles 5–35 mm long; stipules becoming indurated and often fragmenting below apical nodes, interpetiolar, triangular to broadly so, 1.5–2.5 mm long, obtuse, sometimes splitting and appearing bilobed with age; inflorescences terminal, paniculate, peduncle 0.5–2.5 cm long, branched portion 2–3 × 4–5 cm, floral bracts 0.5–1.5 mm long; pedicels 0–4 mm long; calyx limb 2–2.5 mm long, truncate to undulate; corolla white, tube 5–9 mm long, lobes 2–4 mm long; fruit ellipsoid, 7–8 × 5–6 mm, orange-red; pyrenes 2, ventrally plane, dorsally ridged. Deciduous forests, riparian forests, forest islands in

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savannas, 100–400 m; Bolívar (Altiplanicie de Nuria, lower Río Paragua, Río Orinoco, Río Yuruaní, vicinity of San Pedro de las Dos Bocas). Widespread elsewhere in lowland Venezuela; Mexico, West Indies, Central America, Colombia, Ecuador, Peru. Psychotria microdon will be included in the segregate genus Margaritopsis when it is separated from Psychotria. It is quite distinctive, though inexplicably it is often confused with Psychotria pubescens Sw. of the Caribbean region and P. hebeclada of the flora area. Psychotria microdon is a characteristic species of deciduous forests. Steyermark recognized two partly sympatric varieties of this species based on variation in the distribution of its pubescence; these are not recognized here because this species is continuously variable in pubescence.

1972. [Subg. Heteropsychotria]. Shrub to 1.5 m tall, glabrous; leaves 5–7.5 × 1.5–2.5 cm; petioles 4–7 mm long; stipules persistent, united around stem, sheath ca. 0.5 mm long, lobes 2 per side, narrowly triangular, ca. 1 mm long; inflorescence terminal, paniculate, peduncle 1.5–3.5 cm long, branched portion 2–4 × 1.5–6 cm, floral bracts 1–4 mm long, pale green to cream or pink; flowers sessile; calyx limb ca. 0.5 mm long, dentate; corolla white, tube ca. 2.5 mm long, lobes 1.8–2 mm long; fruit subglobose, ca. 2.5 mm diameter, color unknown; pyrenes 2, ventrally with a central longitudinal fissure, dorsally ridged. Lower montane forests, 100–500 m; Amazonas (Sierra de la Neblina). Brazil (Amazonas: Serra da Neblina). The filiform flexuous inflorescence axes of Psychotria nematostachya are distinctive.

Psychotria nana K. Krause, Verh. Bot. Vereins Prov. Brandenburg 50: 109. 1908. [Subg. Heteropsychotria]. Shrub to 2 m tall, glabrous; leaves 7–16 × 1.8–7 cm; petioles 3–6 mm long; stipules fragmenting below apical nodes, interpetiolar, 0.5–1.5 mm long, bilobed to 1/2 their length, each lobe with caducous arista 0.5–1 mm long; inflorescences terminal, subcapitate becoming congested-cymose or shortly fasciculate in fruit, peduncles 1–5 mm long, capitula ca. 5–8 mm diameter, bracts reduced; flowers sessile; calyx limb 0.5–0.8 mm long, subtruncate to denticulate; corolla white, in bud with tube to 2 mm long, lobes to 1 mm long; fruit ellipsoid to subglobose, 5–6 × 5–6 mm, red to red-purple; pyrenes 2, ventrally plane, dorsally shallowly ridged. Lower montane forests, 200–1200 m; Amazonas (base of Cerro Cuao, between Cerro Duida and Cerro Marahuaka). Apure; eastern Colombia, Peru, Amazonian Brazil, Bolivia (0– 90 m outside the flora area). Psychotria nana will probably be included in the segregate genus Margaritopsis when it is separated from Psychotria. It is similar to P. pallidinervia. Psychotria nudiceps Standl. (1931, 378), described from Amazonian central Peru, is probably a synonym of P. nana. More than one species may be included here, but the plants are rather reduced and flowers are rarely collected.

Psychotria oblita Wernham, J. Bot. 52: 314. 1914. [Subg. Heteropsychotria]. Palicourea roraimae Standl., Field Mus. Nat. Hist., Bot. Ser. 17: 280. 1937. Shrub to 2 m tall, glabrous; leaves 1.5–5 × 1–1.8 cm; petioles 3–10 mm long; stipules persistent, united around the stem, sheath 1–3 mm long, lobes 2 per side, triangular, 1– 3.5 mm long; inflorescences terminal, subcapitate to shortly cymose, peduncle 0.5– 2 cm long, capitulum or branched portion 0.5–1.5 × 1–2 cm (not including corollas), bracts green to purple, 7–15 mm long; flowers sessile; calyx limb ca. 1 mm long, lobed; corolla orange, tube 18–21 mm long, lobes 2.5–4 mm long; fruit unknown. Tepui scrub, 2300–2600 m; Bolívar (Ilú-tepui, Roraimatepui, Yuruaní-tepui). Guyana (Mount Roraima). Psychotria oblita is a handsome shrub. It is similar to P. hemicephaelis and P. concinna. Their separation is outlined in the key to the species of subgenus Heteropsychotria.

Psychotria nematostachya Steyerm., Mem. New York Bot. Gard. 23: 512.

Psychotria oblonga (DC.) Steyerm., Mem. New York Bot. Gard. 23: 673. 1972. —Cephaelis oblonga DC. Prodr. 4: 535. 1830. —Uragoga oblonga (DC.) Kuntze, Revis. Gen. 2: 961. 1891. [Subg. Heteropsychotria]. Shrub or subshrub to 2 m tall, glabrous; leaves 3.5–10 × 0.8–4 cm; petioles 2–5 mm long; stipules persistent, united around the stem, sheath 1–1.5 mm long, lobes 2 per side, 0.5–6 mm long, linear and sometimes with 1–

Psychotria 749

several linear appendages; inflorescence terminal, capitate, peduncle 0.5–3 mm long, capitulum ca. 1 × 0.3–0.6 cm, bracts 7–10 mm long, green; flowers sessile; calyx limb 1–1.5 mm long, lobed for ca. 1/2; corolla white, tube 10–11 mm long, lobes ca. 3 mm long; fruit subglobose, 4–5 × 4–5 mm, violet; pyrenes 2, ventrally with a central longitudinal fissure, dorsally angled to ridged. Montane forests, 1000–1100 m; Bolívar (Gran Sabana in headwaters of Río Maurak), Amazonas (Cerro Aratitiyope, Cerro Yutajé). Guyana, Suriname, French Guiana, northeastern Brazil. Psychotria oblonga is similar to P. apoda, P. kappleri, and P. muscosa (Jacq.) Steyerm. of northern Venezuela and the Guianas. Psychotria muscosa differs in its broadly oblong inflorescence heads (versus lanceoloid in P. oblonga) and its corolla tubes ca. 6 mm long. Psychotria occidentalis Steyerm., Mem. New York Bot. Gard. 23: 545. 1972. [Subg. Psychotria]. Subshrub, glabrous; leaves 10.5–19 × 2.5– 6 cm; petioles 1–3 mm long; stipules interpetiolar, caducous, ovate, 9–13 mm long, shortly bilobed, lobes rounded; inflorescence terminal, capitate, peduncle 4.5–9 cm long, head 1–1.2 × 1.2–2 cm, bracts green, 5–5.5 mm long; flowers sessile; calyx limb ca. 3.5 mm long; corolla in bud with color unknown, to 3 mm long; fruit unknown. Riparian forests, ca. 50 m; Delta Amacuro (lower Río Orinoco). Trinidad. Psychotria occidentalis can be recognized by its shortly petiolate leaves that are abruptly and shortly rounded to truncate at the base and its short rounded stipule lobes. Psychotria occidentalis is similar to P. peduncularis Salisb., a widespread and morphologically variable species of wet tropical Africa. Steyermark (1972, 545–546) considered carefully whether the New World collections represent a native neotropical species or a set of mislabelled collections, and concluded that this is indeed a rare native species. He placed P. occidentalis in his subgenus Heteropsychotria section Notopleura, which is here treated as Notopleura. However, P. occidentalis is excluded here from that group based on its terminal inflorescences, though its membranaceous stipules do resemble the laminar stipule appendages of some species of Notopleura. The type speci-

men of P. occidentalis shows the distinctive gray-brown dried color that characterizes most species of subgenus Psychotria, and its stipules are similar to those of P. ernestii and several other species of this group in both their form and their persistence on only the apical 1 or 2 nodes; consequently, it is provisionally here included in subgenus Psychotria. Steyermark (1972) incorrectly considered subgenus Psychotria to be restricted to the Neotropics and all the African Psychotria species, including P. peduncularis, to belong to subgenus Heteropsychotria. However, subgenus Psychotria is actually pantropical and includes P. peduncularis. Psychotria officinalis (Aubl.) Raeusch. ex Sandwith, Kew Bull. 1931: 473. 1931. —Nonatelia officinalis Aubl., Hist. Pl. Guiane 182, pl. 70, fig. 1. 1775. —Psychotria officinalis (Aubl.) Raeusch., Nom.. Bot. 55. 1797, nom. nud. —Psychotria involucrata Sw., Fl. Ind. Occid. 1: 413. 1797, nom. superfl. illeg. [Subg. Heteropsychotria]. Subshrub or shrub to 1 m tall, puberulous to glabrous; leaves 4–14 × 1.2–3.5 cm; petioles 2–6 mm long; stipules persistent, united around stem, sheath 1.5–2 mm long, lobes 2 per side, narrowly triangular, 1–2 mm long; inflorescences terminal, corymbiform, peduncle 1.3–5 cm long, branched portion 0.5– 1.5 × 1–4 cm, floral bracts 4–9 mm long, apparently green; flowers sessile; calyx limb ca. 0.4 mm long, dentate; corolla white, tube ca. 3.5 mm long, lobes ca. 1 mm long; fruit ellipsoid, ca. 2.5 × 3.5 mm, purple; pyrenes 2, ventrally with a central longitudinal fissure, dorsally ridged. Lowland to upland forests, 100– 600 m; Bolívar (basin of Río Cuyuní, upper Río Yuruaní), Amazonas (Río Negro basin). Colombia, Guyana, Suriname, French Guiana, Brazil. Psychotria officinalis is similar to P. hoffmannseggiana and P. adderleyi. See comments regarding their separation under P. adderleyi. Steyermark recognized two subspecies of Psychotria officinalis, with subsp. officinalis found throughout the range of the species and subsp. wilhelmensis Steyerm. reported only from one site in Suriname. The name “Psychotria involucrata” was a superfluous and thus illegitimate new name provided by Swartz for Nonatelia officinalis.

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It was used by a number of 19th-century authors, who also greatly confused the identity of this species by incorrectly applying the name P. involucrata to what are now considered about half a dozen species. The confusion was greatly increased in the 20th century by Standley in his various Mesoamerican and western South American floras. There he applied the name P. involucrata to several species that are not conspecific with the Guianan plant. Later Dwyer (1980) followed this incorrect application of the name. The attribution of authorship here of P. officinalis follows Steyermark (1972).

long, bilobed, lobes deltoid; inflorescences terminal, subcapitate, peduncles 1–2 mm long, capitula 5–8 × 9–10 mm, bracts reduced; flowers sessile; calyx limb 1–1.5 mm long, shallowly dentate; corolla white, tube ca. 2 mm long, lobes ca. 1.5 mm long; fruit unknown. Lower montane forests, 500–700 m; Amazonas (Sierra de la Neblina). Endemic. Psychotria pallidinervia may be included in the segregate genus Margaritopsis when it is separated from Psychotria, but the limits of that group are not yet completely clear. Psychotria pallidinervia is poorly known, but is similar to P. nana and perhaps not distinct.

Psychotria pacimonica Müll. Arg. in Mart., Fl. Bras. 6(5): 337. 1881. [Subg. Heteropsychotria]. Psychotria necopinata Standl., Field Mus. Nat. Hist., Bot. Ser. 11: 242. 1936. Shrub or small tree to 10 m tall, glabrous; leaves 12–27 × 3–6 cm; petioles 4–22 mm long; stipules generally persisting on apical nodes, united around stem, sheath 1.5–4 mm long, truncate to undulate; inflorescences terminal, capitate or subcapitate, peduncle 3–6 cm long, capitula 1–1.5 × 1.5–2 cm (not including corollas), external bracts green, 1.5–4.5 cm long; flowers sessile; calyx limb 0.5–1.5 mm long, truncate; corolla white, tube 9–10 mm long, lobes ca. 2.5 mm long; fruit ellipsoid, ca. 10 × 7 mm, color unknown; pyrenes 2, ventrally plane, dorsally ridged. Riparian forests, seasonally flooded forests, 100–200 m; Amazonas (basins of Río Negro and Río Pasimoni). Amazonian Brazil. Psychotria pacimonica may be included in the segregate genus Carapichea when it is separated from Psychotria, but its limits are not yet completely clear. The well-developed peduncles, flowering heads surrounded by 4– 8 relatively long and narrow green bracts, truncate stipules, and closely set intersecondary leaf veins are distinctive for Psychotria pacimonica. Delprete (2001) outlined its separation from Psychotria ligularis (Rudge) Steyerm. of the Guianas.

Psychotria paniculata (Aubl.) Raeusch., Nom. Bot. 56. 1797. —Nonatelia paniculata Aubl., Hist. Pl. Guiane 181, t. 70, fig. 2. 1775. —Psychotria flexuosa Willd., Sp. Pl. 1: 966. 1798, nom. superfl. illeg. [Subg. Heteropsychotria]. Shrub or small tree to 3 m tall, glabrous; leaves 18–21 × 3.5–12 cm; petioles 5–10 mm long; stipules persistent, interpetiolar and sometimes shortly united around stem, triangular, 2.5–4 mm long, emarginate or bilobed to 1/2 their length, lobes triangular; inflorescence terminal, paniculate, peduncle 3.5–7 cm long, branched portion 5–9 × 4–7 cm, floral bracts reduced, pedicels 0–0.5 mm long; calyx limb ca. 0.2 mm long, subtruncate; corolla white or pale green, tube ca. 3 mm long, lobes ca. 1 mm long; fruit subglobose, 3–3.5 × 4–4.5 mm, black; pyrenes 2, ventrally with a rounded rather deep central cavity, dorsally ridged. Upland forests, 300–1100 m; Bolívar (near El Paují, near Santa Elena de Uairén), Amazonas (uppermost Río Orinoco). Guyana, Suriname, French Guiana, Brazil. Psychotria paniculata can be recognized by the combination of its stipules that are essentially interpetiolar and emarginate to bilobed with the lobes closely set with its inflorescence axes branched to several orders.

Psychotria pallidinervia Steyerm., Mem. New York Bot. Gard. 23: 492. 1972. [Subg. Heteropsychotria]. Shrub to 2 m tall, glabrous; leaves 8–15 × 2–3 cm; petioles 5–10 mm long; stipules fragmenting after distalmost node, interpetiolar and shortly united around stem, ca. 1.5 mm

Psychotria parimensis Steyerm., Mem. New York Bot. Gard. 23: 691. 1972. [Subg. Heteropsychotria]. Subshrub, hirsute; leaves 7.5–13.5 × 2–3.5 cm, sessile; stipules apparently persistent, apparently united around stem, sheath not described, lobes 2 per side, narrowly triangular, 8–9 mm long; inflorescence and flowers unknown; infructescence terminal, capitate,

Psychotria 751

peduncle ca. 5 mm long, capitulum ca. 1.8 × 2.5 cm, external bracts ca. 18 mm long, color unknown; fruit subglobose, ca. 5 mm diameter, color unknown, with persistent calyx limb ca. 1.5 mm long, dentate; pyrenes 2, ventrally with a central longitudinal fissure, dorsally ridged. Upland forests, 500–600 m; Amazonas (Sierra Parima). Endemic. Psychotria parvibractea Steyerm., Mem. New York Bot. Gard. 23: 581, fig. 80. 1972. [Subg. Heteropsychotria]. Shrub or small tree to 4 m tall, glabrous; leaves 9–15 × 3.5–7.5 cm; petioles 11–15 mm long; stipules persistent, united around stem, sheath 2–2.5 mm long, lobes 2 per side, deltoid, 0.8–1 mm long; inflorescence terminal, paniculate, peduncle 3–5 cm long (to 6 cm long in Guyana), branched portion 3.5–7 × 4–6 cm, bracts 2–3.5 mm long, pale green (to pink in Guyana); flowers sessile; calyx limb ca. 0.8 mm long, dentate; corolla cream, tube 4–4.5 mm long, lobes ca. 2 mm long; fruit ellipsoid, ca. 4 × 3 mm, pale blue (to deep blue in Guyana); pyrenes 2, ventrally with a central longitudinal fissure, dorsally ridged. Wet montane forests, ca. 1000 m; Bolívar (Cerro Venamo, basin of Río Cuyuní). Guyana. Psychotria parvibractea is similar in general aspect to P. wurdackii, but differs from that species in its smaller bracts and capitula. Psychotria paupertina Standl. & Steyerm., Fieldiana, Bot. 28: 603. 1953. [Subg. Heteropsychotria]. Shrub to 4 m tall, glabrous; leaves 10.5– 19.5 × 4–9 cm; petioles 1–5 mm long; stipules interpetiolar and sometimes very shortly united around stem, ligulate to ovate, 4.5–6 mm long, rounded to obtuse, with 1 or 2 caducous aristas 0.5–2 mm long; inflorescences terminal, subcapitate to congested-cymose, peduncle 5–12 mm long, branched portion 1– 1.2 × 1.5–2.5 cm, bracts reduced; flowers sessile; calyx limb 2–3 mm long, shallowly dentate; corolla cream, tube ca. 3.5 mm long, lobes 2–2.5 mm long; fruit ellipsoid to subglobose, 17–25 × 15–17 mm, dark purplered; pyrenes 2, ventrally plane, dorsally ridged. Evergreen lowland and upland forests, 100–600 m; Bolívar (Río Cuyuní basin, Río Paramichi), Amazonas (base of Cerro Duida, Cerro Sipapo, Río Orinoco below San Fernando de Atabapo, Río Sipapo). Endemic.

Psychotria paupertina may be included in the segregate genus Margaritopsis when it is separated from Psychotria, but the limits of that group are not yet completely clear. Psychotria paupertina is similar to P. podocephala. Psychotria pectinata Steyerm., Ann. Missouri Bot. Gard. 75: 344, fig. 11. 1988. [Subg. Psychotria]. Shrub to 2 m tall, glabrous; leaves 6–15 × 1.5–3 cm; petioles 3–15 mm long; stipules interpetiolar, usually persistent with the leaves, sheath 1.5–3 mm long, lobes linear, 1.5–6 mm long, densely hirtellous; inflorescences terminal, subcapitate to congested-cymose, peduncle 0.1–1.3 cm long, flower-bearing portion ca. 0.5 × 1 cm, floral bracts ca. 1.5 mm long, pedicels 0.5–1 mm long; calyx limb 1–2 mm long, lobed for 1/2 or more its length; corolla white, tube ca. 2 mm long, lobes ca. 1.8 mm long; fruit ellipsoid to ovoid, 8–9 × 5 mm, bright red; pyrenes 2, dorsally ridged. Evergreen lowland and riparian forests, 100–200 m; Amazonas (base of Sierra de la Neblina). Endemic. The stipules of Psychotria pectinata are distinctive, though they are similar to those of P. graciliflora Benth. of Central America. One specimen from French Guiana (Feuillet 10006, MO!) may represent a range extension of this species, though the calyx limbs of its immature flowers are apparently shorter so its identification as P. pectinata is not completely confirmed. Psychotria phaneroloma Standl. & Steyerm., Fieldiana, Bot. 28: 604. 1953. [Subg. Heteropsychotria]. —Wairá-yuyek (Pemón). Psychotria phaneroloma var. angustior Steyerm., Mem. New York Bot. Gard. 23: 591. 1972. Subshrub or shrub to 1.7 m tall, glabrous; leaves 3–9.5 × 0.5–4 cm; petioles 4–9 mm long; stipules persistent, united around the stem, sheath 1–2 mm long, lobes 2 per side, narrowly triangular, 1–2 mm long; inflorescences terminal, spiciform to subcapitate or paniculate, peduncles 1–6 cm long, capitula or branched portion 1–3 × 1–2.5 cm, bracts 3–6 mm long; flowers sessile; calyx limb ca. 0.8 mm long, dentate; corolla white, tube 2.5–3 mm long, lobes ca. 1 mm long; fruit subglobose, ca. 3 mm diameter, blue to

752

R UBIACEAE

purple; pyrenes 2, ventrally with a central invagination, dorsally ridged. Gallery forests, riparian forests, rocky igneous sites near rapids, tepui slope forests, savannas, 400–1300 m; Bolívar (Río Aparamán, Río Ichún, Río Pauo in basin of Río Caura, Río Tírica), Amazonas (Cerro Aratitiyope, Cerro Parú). Guyana, Brazil. ◆Fig. 579. The secondary leaf veins that extend unbranched to unite with the thickend margins and the usually subcapitate to spiciform or congested-cymose inflorescences with the flowers borne in capitula are distinctive for Psychotria phaneroloma. It is similar to P. egensis, P. lourteigiana, P. polycephala, and P. moyobamba Standl. of Peru. Steyermark recognized two varieties of Psychotria phaneroloma, var. angustior and var. phaneroloma, but these are not recognized as distinct here due to continuous variation. Steyermark (1972; 1974 indirectly) cited Schomburgk s.n. from Guyana as the type of Psychotria phaneroloma, but Standley and Steyermark’s protologue actually cites a different type, “Steyermark 58117, F.” This latter specimen has not been seen; however, Steyermark 59117 (MO) is annotated in Steyermark’s hand as “Psychotria phaneroloma sp. nov.,” its label data correspond exactly with the data cited in the protologue, and this second number is not cited in the protologue. Therefore, the protologue appears to contain a typographical error with respect to the type collection number. Psychotria phelpsiana Steyerm., Mem. New York Bot. Gard. 23: 889, fig. 19. 1972. [Subg. Heteropsychotria]. Shrub or small tree to 3 m tall, puberulous or strigillose to glabrous; leaves 2.5–5 × 0.5–3 cm, sessile or subsessile, the petioles to 4 mm long; stipules persistent, united around stem, sheath 1.5–2 mm long, lobes 1 per side, narrowly triangular, 1–2 mm long; inflorescences pseudoaxillary and/or axillary, subcapitate to congested-cymose, peduncle 1–3.5 cm long, capitula or cymes ca. 0.5 × 0.5 cm, bracts 1–2 mm long; flowers sessile; calyx limb 1–1.2 mm long, lobed for ca. 1/2 its length; corolla white, tube ca. 4 mm long, lobes ca. 1 mm long; immature fruit ellipsoid, to 4.5 × 3 mm, reportedly 8–10-winged; pyrenes unknown. Tepui summit scrub, ca.

2100 m; Bolívar (Cerro Sarisariñama). Endemic. ◆Fig. 580. Psychotria phelpsiana is similar to P. carrenoi; see additional comments there. Steyermark described the young fruits of this species as winged, but this unusual feature is not evident on the hypanthia of the specimens seen, and the type, apparently the only collection with fruits, has not been seen. Psychotria pilosa Ruiz & Pav., Fl. Peruv. 2: 60, pl. 208. 1799. [Subg. Heteropsychotria]. Shrub to 3.5 m tall, hirsute or pilose; leaves 12–20 × 4.5–11 cm; petioles 0.8–4 cm long; stipules persistent on apical nodes, interpetiolar and sometimes shortly intrapetiolar, ovate in outline, 10–24 mm long, lobed for 1/4–1/2 their length; inflorescences terminal, paniculate, peduncles 1.5–4.5 cm long, branched portion 2–6 × 2.5–6 cm, bracts 4–12 mm long, floral bracts 4–7 mm long; flowers sessile; calyx limb 1–1.2 mm long, deeply lobed; corolla white, tube 2.5–3 mm long, lobes ca. 1 mm long; fruit ellipsoid, 4–5 × 4–5 mm, blue; pyrenes 2, ventrally with a longitudinal furrow, dorsally ridged. Tepui slope forests dominated by Iriartea palms, 1200–1300 m; Amazonas (Sierra de la Neblina). Central America, Colombia, Ecuador, Peru, western Brazil, Bolivia. Psychotria pilosa is distinctive due to its pilose or hirsute pubescence and well-developed, almost interpetiolar stipules. Psychotria platypoda DC., Prodr. 4: 510. 1830. [Subg. Heteropsychotria]. Cephaelis dichotoma Rudge, Pl. Guian. 1: 29, pl. 44. 1805, not Cephaelis dichotoma Willd. ex Roem. & Schult. 1819. —Psychotria dichotoma (Rudge) Bremek., Recueil Trav. Bot. Néerl. 31: 301. 1934, nom. illeg., not Psychotria dichotoma Humb. & Bonpl. ex Roem. & Schult. 1819. Psychotria martiana Müll. Arg. in Mart., Fl. Bras. 6(5): 339, pl. 51. 1881. Subshrub or shrub to 3 m tall, glabrous; leaves 8–20 × 3–8.5 cm, petioles 2–12 mm long; stipules persistent, united around the stem, sheath 0.5–2 mm long, lobes 2 per side, narrowly triangular, 1.5–4 mm long; inflorescences terminal, capitate, peduncles 0.5–1 cm long, capitula 1.2–2 × 0.8–1 cm, external

Psychotria 753

bracts green, 7–10 mm long; calyx limb 0.8–1 mm long, undulate to dentate; corolla white sometimes tinged pink, tube 5–6.5 mm long, lobes 1.2–2 mm long; fruit ellipsoid, 4–5 × 3.5–5 mm, purple; pyrenes 2, ventrally with a central longitudinal fissure, dorsally smooth to ridged. Semideciduous upland forests, 600–800 m; Bolívar (Río Icabarú basin), Amazonas (Río Matacuni, Sierra Parima). Central America, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. The inflorescences of Psychotria platypoda are capitate and enclosed by involucral bracts, but their axes frequently begin to elongate as the fruits develop. The plants with branched infructescences are usually still recognizable as P. platypoda by their persistent elliptic, ovate, or suborbicular bracts that persist at the bases of the axes. The inflorescences are distinctively nodding. This species is similar to P. casiquiaria (which may not be distinct) and P. gracilenta. The largest pyrenes are smooth dorsally but smaller ones are often shallowly to notably ridged, which may be because the pyrenes expand and become smooth as they mature. Psychotria podocephala (Müll. Arg.) Standl., Publ. Field Columbian Mus., Bot. Ser. 7: 109. 1930. —Mapouria podocephala Müll. Arg., Flora 59: 460, 466. 1876. [Subg. Heteropsychotria]. —Caruto, Medi-wadi (Yekwana). Shrub or small tree to 5 m tall, glabrous; leaves 12–27 × 4.5–11 cm; petioles 3–5 mm long; stipules fragmenting after nodes, interpetiolar and sometimes shortly united around stem, triangular to ovate, 3–10 mm long, obtuse to acute and sometimes shortly 1- or 2apiculate; inflorescences terminal, capitate, peduncle 0–9 mm long, capitula 12–18 mm diameter, bracts reduced; flowers sessile; calyx limb 2.5–5 mm long, shallowly lobed; corolla white, tube ca. 5 mm long, lobes ca. 2.5 mm long; fruit ellipsoid, 10–15 × 8–11 mm, red; pyrenes 2, ventrally plane, dorsally ridged. Riparian forests, lowland to upland forests, 100– 600 m; Bolívar (upper Río Paragua basin), Amazonas (basins of Río Guainía and Río Orinoco). Amazonian Brazil. ◆Fig. 585. Psychotria podocephala may be included in the segregate genus Margaritopsis when it is separated from Psychotria, but the limits

of this other group are not yet completely clear. The crude bark of this species is used as a remedy for the bite of the 24-hour ant by the Yekwana Amerindians. This species is similar to P. paupertina. Psychotria poeppigiana Müll. Arg. in Mart., Fl. Bras. 6(5): 370. July 1881. [Subg. Heteropsychotria]. Tapogomea tomentosa Aubl., Hist. Pl. Guiane 160, pl. 61. 1775, not Psychotria tomentosa Hemsl., April 1881. —Callicocca tomentosa (Aubl.) J.F. Gmel., Syst. Nat. 1: 371. 1791. —Cephaelis tomentosa (Aubl.) Vahl, Eclog. Amer. 1: 19. 1796. —Uragoga tomentosa (Aubl.) K. Schum. in Engl. & Prantl, Nat. Pflanzenfam. 4(4): 120. 1891. Cephaelis hirsuta M. Martens & Galeotti, Bull. Acad. Roy. Sci. Bruxelles 11(1): 135. 1844, not Psychotria hirsuta Sw. 1797. Subshrub or shrub to 3.5 m tall, strigose or hirsute; leaves 7–30 × 2.5–11.5 cm; petioles 6–50 mm long; stipules persistent, united around stem, sheath 2–5 mm long, lobes 2 per side, narrowly triangular, 4–11 mm long; inflorescences terminal, capitate, peduncle 0.5–12(22) cm long, capitulum 1.5– 3 × 1–2 cm (not including involucral bracts), involucral bracts 2, orange to red, 3–7 cm long; flowers sessile; calyx limb 0.5–2 mm long, deeply lobed; corolla yellow, tube 13– 14.5 mm long, lobes ca. 2.5 mm long; fruit ellipsoid, 5.5–7 × 3 mm, bright blue; pyrenes 2, ventrally with a central longitudinal fissure or invagination, dorsally ridged to angled. Evergreen lowland to montane forests, 50– 1300 m. Mexico, Central America, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Bolivia, Argentina; 2 subspecies, both in the flora area. Psychotria poeppigiana is very showy with its brightly colored bracts, yellow flowers, and blue fruits. It is one of the most frequently collected species of Psychotria in the Neotropics. Steyermark (1972) described the pyrenes as smooth ventrally, but all of those examined have a longitudinal invagination or fissure, though this is smoothly concave rather than sharply edged. Steyermark’s two subspecies are apparently often sympatric; these are here maintained provisionally pending a comprehensive, careful study of this species.

754

R UBIACEAE

Key to the Subspecies of P. poeppigiana 1. Pubescence of midrib and lateral veins of the lower leaf surface appressed, strigose ......................... subsp. barcellana 1. Pubescence of midrib and lateral veins of the lower leaf surface spreading, hirsute or villous .............. subsp. poeppigiana P. poeppigiana subsp. barcellana (Müll. Arg.) Steyerm., Mem. New York Bot. Gard. 23: 680. 1972. —Psychotria barcellana Müll. Arg. in Mart., Fl. Bras. 6(5): 369. 1881. —Cephaelis barcellana (Müll. Arg.) Standl., Publ. Field Columbian Mus., Bot. Ser. 8: 184. 1930. [Subg. Heteropsychotria]. —Kowa-wakaanamahu (Yanomami), Oreja de picure, Oreja de rabo pelado, Tulipán, Tulipán rojo montañero. 50–1300 m; Bolívar (widespread), Amazonas (widespread). Monagas; Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil. P.

poeppigiana subsp. poeppigiana. [Subg. Heteropsychotria]. 90–1300 m; Delta Amacuro (Serranía de Imataca), Bolívar (Gran Sabana, Ilu-tepui, Río Ichún), Amazonas (Isla Ratón, Puerto Ayacucho, region of San Carlos de Río Negro). Widespread elsewhere in lowland Venezuela; Mexico, Central America, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Bolivia, Argentina. ◆Fig. 584. Subspecies poeppigiana is the most frequently collected subspecies throughout the range of the species and in most of Venezuela, but in the flora area it is found in widely scattered sites and is less common than subsp. barcellana. Psychotria polycephala Benth., J. Bot. (Hooker) 3: 231. 1841. [Subg. Heteropsychotria]. Psychotria racemifera Standl., Publ. Field Columbian Mus., Bot. Ser. 8: 71. 1841. Subshrub or shrub to 1 m tall, glabrous; leaves 4.5–11 × 1.5–4 cm; petioles 3–10 mm long; stipules persistent, united around stem, sheath 0.5–2.5 mm long, lobes 2 per side, narrowly triangular, 1.5–4 mm long; inflorescence terminal, paniculate, peduncle 1– 3 cm long, branched portion 3–7.5 × 1–1.5

cm, bracts subtending capitula 4–5 mm long, floral bracts 1.5–2 mm long; flowers sessile; calyx limb 0.5–0.8 mm long, dentate; corolla white, tube 3–3.5 mm long, lobes 1–1.2 mm long; fruit subglobose, ca. 3 mm diameter, purple to blue; pyrenes 2, ventrally with a central longitudinal invagination or furrow, dorsally ridged and pustulate. Evergreen lowland to montane forests, 50–1200 m; Delta Amacuro (San Victor), Bolívar (common in basins of Río Caroní, Río Caura, Río Cuyuní, and Río Paragua), Amazonas (Mavaca, upper Río Orinoco). Zulia; Guyana, Peru, Brazil, possibly Colombia. ◆Fig. 590. Psychotria polycephala is distinctive due to the termination of each of the secondary axes in a relatively small (ca. 3 × 3 mm) capitulum and the union of most or all of the unbranched secondary leaf veins with the cartilaginous margin. This species is similar to Psychotria phaneroloma. Psychotria prunifolia (Kunth) Steyerm., Mem. New York Bot. Gard. 23: 655. 1972. —Cephaelis prunifolia Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 3: 377. 1818 [1819]. [Subg. Heteropsychotria]. Cephaelis microcephala Willd. ex Roem. & Schult., Syst. Veg. 5: 214. 1819. —Psychotria microcephala (Willd. ex Roem. & Schult.) Müll. Arg. in Mart., Fl. Bras. 6(5): 351. 1881, nom. illeg., not Psychotria microcephala Miq. 1850 [1851]. Psychotria microcephala var. tripotamica Müll. Arg. in Mart., Fl. Bras. 6(5): 352. 1881. Psychotria xanthocephala Müll. Arg. in Mart., Fl. Bras. 6(5): 351, pl. 53, fig. 1. 1881. Subshrub or shrub to 1.5 m tall, hirtellous or strigillose to glabrescent; leaves 4–13 × 2– 7 cm; petioles 3–7 mm long; stipules persistent, united around stem, sheath ca. 0.5 mm long, lobes 2 per side, narrowly triangular, 2– 4 mm long; inflorescences terminal, capitate, peduncle 0–3 mm long, capitulum 1–2 × 0.5 cm, external bracts 10–20 mm long, pandurate, yellow at base to green above; flowers sessile; calyx limb 2.5–3 mm long, dentate; corolla white, tube 10–11 mm long, lobes ca. 3 mm long; fruit ellipsoid to subglobose, 4–5 × 3–4 mm, blue to black; pyrenes 2, ventrally with a central longitudinal fissure, dorsally angled. Riparian forests, forests over igneous substrate, ca. 100 m; Amazonas (base of

Psychotria 755

Cerro Arauicaua, Río Casiquiare, Río Yatúa). Peru, Brazil, Bolivia. ◆Fig. 583. Psychotria prunifolia is similar to Psychotria turbinella. The external inflorescence bracts are unusual in their pandurate shape, with the broadened apical portion foliaceous.

This species was long treated under the name Psychotria racemosa (Aubl.) Raeusch., but Kirkbride (1997) showed that although this name applies to the same species, it is based on a different type than Richard’s name and thus is a later homonym.

Psychotria racemosa Rich., Actes Soc. Hist. Nat. Paris 1: 107. 1792. [Subg. Heteropsychotria]. —Bonita de noche. Nonatelia racemosa Aubl., Hist. Pl. Guiane 187, pl. 72. 1775. —Oribasia racemosa (Aubl.) J.F. Gmel., Syst. Nat. 367. 1791. —Psychotria racemosa (Aubl.) Raeusch., Nom. Bot. 56. 1797, hom. illeg., not Psychotria racemosa Rich. 1792. Psychotria longistipula Benth., J. Bot. (Hooker) 3: 227. 1841. Subshrub or shrub to 3 m tall, hirtellous to glabrescent; leaves 8.5–18 × 2.5–8.5 cm; petioles 3–15 mm long; stipules persistent, united around stem, sheath 1.5–3 mm long, lobes 2 per side, narrowly triangular, 5–12 mm long; inflorescence terminal, paniculate, peduncle 0.5–3.5 cm long, branched portion 1.5–3.5 × 3–4.5 cm, floral bracts 0.5–1.5 mm long, green, pedicels 0–2 mm long; calyx limb 0.5–1 mm long, lobed; corolla white, tube 1.5–3 mm long, lobes 1.2–1.5 mm long, with abaxial projections 0.5–1 mm long; fruit subglobose to oblate, 3.5–4 × 5–6 mm, becoming orange then red then finally black; pyrenes 5, ventrally with a central longitudinal fissure, dorsally ridged. Lowland forests, riparian forests, Río Negro caatinga forests on white sand, 50–600 m; Delta Amacuro (Serranía de Imataca), Bolívar (Altiplanicie de Nuria, basins of Río Caroní, Río Caura, Río Cuyuní, Río Paragua, and Río Orinoco), Amazonas (Río Guainía, basin of Río Orinoco, Río Siapa, and Río Ventuari). Distrito Federal; Mexico, Central America, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. The five pyrenes, relatively long narrow stipule lobes, and short corollas with well-developed abaxial projections borne on backs of the lobes are distinctive for Psychotria racemosa. The fruits are frequently reported to be orange or red but they do finally turn black or purple-black when mature (personal observation), and then are quickly removed by frugivores. Vegetatively only P. anartiothrix is similar. Their distinctions are discussed under the latter species.

Psychotria remota Benth., J. Bot. (Hooker) 3: 225. 1841. —Mapouria remota (Benth.) Müll. Arg. in Mart., Fl. Bras. 6(5): 407. 1881. [Subg. Psychotria]. —Hoja de guacuraya, Pata de picure. Psychotria alboviridula K. Krause, Notizbl. Königl. Bot. Gart. Berlin 6: 208. 1914. Shrub or small tree to 8 m tall, glabrous; leaves 7–24 × 2–10 cm; petioles 2–15 mm long; stipules caducous, interpetiolar, triangular to ligulate, 3–5 mm long, acute; inflorescences terminal, paniculate, peduncles 2– 10.5 cm long, branched portion 5–15 × 4.5–8 cm, floral bracts ca. 0.5 mm long, pedicels 0– 0.5 mm long; calyx limb 0.5–1 mm long, subtruncate to shortly dentate; corolla white, tube ca. 2 mm long, lobes ca. 1 mm long; fruit ellipsoid, 6–9 × 4–6 mm, red; pyrenes 2, dorsally ridged. Riparian forests, evergreen lowland forests, seasonally flooded forests, 50– 200 m; Amazonas (Río Casiquiare, Río Guainía, Río Negro, upper Río Orinoco basin, Río Ventuari). Central America, Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Amazonian Brazil, Bolivia. The subspiciform inflorescence axes with the higher-order axes, flowers, and fruit typically spreading at nearly 90° from each other are distinctive for this species. Psychotria remota is frequently confused with P. viridis Ruiz & Pav. of Central America and western South America, which has not yet been found in the flora area, but may be expected. Psychotria viridis can be distinguished by its fully spiciform inflorescence axes with the flowers and fruits sessile in dense sessile glomerules and its shortly bifid stipules. Psychotria rosea (Benth.) Müll. Arg. in Mart., Fl. Bras. 6(5): 360. 1881. —Cephaelis rosea Benth., J. Bot. (Hooker) 3: 224. 1841. [Subg. Heteropsychotria]. —Palo de sortija. Psychotria rosea var. rosea f. calvescens Steyerm., Mem. New York Bot. Gard. 23: 688. 1972. Subshrub or shrub to 2 m tall, puberulous, pilosulous, or strigillose to glabrescent;

756

R UBIACEAE

leaves 8.5–27 × 2.5–11 cm; petioles 4–25 mm long; stipules persistent, united around stem, sheath 1–3 mm long, lobes 2 per side, narrowly triangular, 4–15 mm long; inflorescence terminal, capitate, peduncle 0.5–6 cm long, capitulum 1–2 × 2–7 cm, external bracts purple to white, 15–25 mm long; flowers sessile; calyx limb 1.5–2 mm long, lobed; corolla white to purple, tube 10–11 mm long, lobes ca. 2 mm long; fruit ellipsoid, 4–5 × 3– 4.5 mm, purple; pyrenes 2, ventrally with a narrow central longitudinal fissure, dorsally ridged. Riparian forests, periodically flooded forests, lowland forests, 50–200 m; Amazonas (basins of Río Emoni, Río Guainía, Río Negro, Río Orinoco, Río Pasimoni, and Río Ventuari, Río Yatúa). Táchira; Colombia, Guyana, Brazil. See comments under Psychotria colorata, which is similar (and may not be distinct). Steyermark recognized two varieties and two forms of P. rosea based on details of pubescence, but the pubescence seems to vary widely and continuously and consequently those are not recognized here. Psychotria schomburgkii Benth., J. Bot. (Hooker) 3. 231. 1841. [Subg. Heteropsychotria]. Psychotria chondroloma Standl., Publ. Field Columbian Mus., Bot. Ser. 7: 440. 1931. Shrub or small tree to 2(–10) m tall, puberulous to glabrous; leaves 7.5–23 × 2–7 cm; petioles 4–17 mm long; stipules persistent, united around stem, sheath 1–1.5 mm long, lobes 2 per side, triangular to narrowly so, 1–1.5 mm long; inflorescences terminal, subcapitate to corymbiform, peduncle 0.5–6 cm long, capitulum or branched portion 1–3 × 1.5–5 cm, bracts green, 2–5 mm long; flowers sessile; calyx limb 0.8–1 mm long, deeply lobed; corolla white, tube 2.5–3 mm long, lobes ca. 1.8 mm long; fruit ellipsoid, 4–4.5 × 3 mm, orange; pyrenes 2, ventrally with a rather broad central longitudinal fissure, dorsally ridged. Montane and lower montane forests, 400–1500 m; Amazonas (Cerro Duida, Cerro Huachamacari, Cerro Marahuaka, Cerro Yutajé, Sierra Parima). Guyana, Brazil. Psychotria schomburgkii is distinctive due to its lanceolate, rather thick-textured leaves with the secondary veins extending unbranched to unite with the thickened mar-

gins, its corymbiform inflorescences with the secondary axes each terminating in a capitulum, and its orange fruits. Psychotria hyptoides Benth. is probably a synonym of P. schomburgkii, but is poorly known. Psychotria sipapoensis Steyerm., Mem. New York Bot. Gard. 23: 708, fig. 90. 1972. [Subg. Heteropsychotria]. Apparently a shrub (habit and height not noted), glabrous; leaves 10–16 × 5.5–12 cm, sessile; stipules persistent, united around stem, sheath ca. 2 mm long, truncate to shallowly emarginate; inflorescence terminal, paniculate, peduncle 7–10 cm long, branched portion 9–12 × 5–7 cm, floral bracts 0.5–1 mm long; flowers sessile; calyx limb ca. 0.5 mm long, dentate; corolla in bud 3–4 mm long, color not noted; fruit unknown. Montane forests, ca. 600 m; Amazonas (Cerro Sipapo). Endemic. Psychotria sipapoensis is a poorly known species similar to Psychotria cardiomorpha and Palicourea foldatsii. It may eventually be demonstrated to be a well-marked form of P. cardiomorpha, which probably has more variation in leaf and inflorescence size than Steyermark knew in 1972. Psychotria spadicea (Pittier) Standl. & Steyerm., Fieldiana, Bot. 28: 1105. 1957. —Pagamea spadicea Pittier, Bol. Soc. Venez. Ci. Nat. 9: 123. 1944. [Subg. Heteropsychotria]. —Carisillo, Paripari. Shrub to 1.5 m tall, puberulous to glabrescent; leaves 4.5–10.5 × 1–3.5 cm; petioles 1–5 mm long; stipules persistent, united around stem, sheath ca. 0.5 mm long, lobes 2 per side, narrowly triangular, 1–2.5 mm long; inflorescences terminal or sometimes displaced to pseudoaxillary, capitate, sessile or with peduncle to 0.3 cm long, capitulum 0.5–0.8 cm diameter, bracts green, 3–4 mm long; flowers sessile; calyx limb ca. 1 mm long, dentate; corolla white, tube 2.5–3 mm long, lobes 1–1.5 mm long; fruit subglobose, ca. 3 × 3 mm, purple, blue, or black; pyrenes 2, ventrally with a longitudinal central fissure that is widened in the distal 1/2, dorsally ridged. Riparian forests, lowland to upland forests, white-sand savannas, 100–800 m; Bolívar (Cerro Guaiquinima, Río Paragua basin, Río Tonoro), Amazonas (basins of Río Autana, Río Guainía, Río Negro, and Río Orinoco). Colombia. ◆Fig. 564.

Psychotria 757

The leaves of Psychotria spadicea are used by some of the local inhabitants to relieve itching caused by other plants. It is similar to P. casiquiaria and P. vichadensis. Psychotria speluncae Standl. & Steyerm., Fieldiana, Bot. 28: 606. 1953; emend. Steyerm., Mem. New York Bot. Gard. 17(1): 436. 1967. [Subg. Heteropsychotria]. Psychotria speluncae subsp. brevicalyx Steyerm., Mem. New York Bot. Gard. 17(1): 436. 1967. Psychotria speluncae subsp. exserta Steyerm., Mem. New York Bot. Gard. 17(1): 436. 1967. Subshrub or shrub to 2.5 m tall, glabrous; leaves 3–6.4 × 1–2 cm; petioles 2–7 mm long; stipules persistent, united around stem, sheath 1–3 mm long, lobes 2 per side, linear, 1.5–4.5 mm long; inflorescence terminal, congested-cymose to corymbiform, peduncle 1– 3.5 cm long, branched portion 1–1.5 × 1–2.5 cm, floral bracts 3.5–5 mm long, apparently green; flowers sessile; calyx limb 3–4 mm long, shallowly to deeply lobed; corolla white, tube 6.5–7.5 mm long, lobes 3–4.5 mm long, with abaxial projections to 0.5 mm long; fruit ellipsoid, 6–8 × 6.5–7 mm, blue or purple; pyrenes 2, ventrally with a central longitudinal fissure, dorsally ridged. Tepui slope and summit forests, 1200–2300 m; Bolívar (Auyán-tepui, Cerro Guaiquinima, Cerro Venamo, Ilú-tepui, Macizo del Chimantá, Ptari-tepui, Roraima-tepui), Amazonas (Sierra de la Neblina). Guyana. ◆Fig. 586. Steyermark’s emendation of Psychotria speluncae consisted of noting that the flowers are not always 4-merous, as in the original description, but sometimes may be 5-merous. Psychotria speluncae is similar to P. vellosiana and P. heteroneura. Steyermark recognized three subspecies of Psychotria speluncae, all from the flora area. One of these, subsp. exserta Steyerm., was distinguished primarily by its stamens exserted rather than included; however, the presence of both of these conditions in the same species is normal in distylous species, which P. speluncae apparently is, rather than a taxonomically informative character. Steyermark also based the separation of subsp. exserta on its having the “lateral peduncles” arranged as “solitary” instead of “usually 2–3-flowered;” the meaning of this is

not clear but this character appears to be continuously variable. Steyermark separated his subsp. brevicalyx Steyerm. by its calyx lobes 0.7–1 mm long and stipule lobes 1.5–2 mm long, versus 1.2–2 and 1.5–4 mm long, respectively, in subsp. speluncae. Each of these subspecies was based on a single collection from a different tepui, and with more collections now available the range of variation in all of these features is now clearly similar on each tepui; consequently, these subspecies are not recognized here. Psychotria sphaerocephala Müll. Arg., Flora 59: 550, 553. 1876. —Cephaelis sphaerocephala (Müll. Arg.) B.L. Rob., Proc. Amer. Acad. 45: 408. 1910. [Subg. Heteropsychotria]. Subshrub or low shrub to 2 m tall, strigillose to hirtellous; leaves 6.5–12 × 2.5–5 cm; petioles 3–5 mm long; stipules persistent, united around stem or often subinterpetiolar, 3–5 mm long, deeply lacerate; inflorescences terminal, capitate, peduncle 2.5–4 cm long, capitulum 0.5–1 cm diameter, bracts 2.5–5 mm long; flowers sessile; calyx limb 1.5–2 mm long, deeply lobed; corolla white, tube ca. 4 mm long, lobes ca. 1.5 mm long; fruit subglobose, 3–4 × 1.5–3 mm, white; pyrenes 2, ventrally with a central furrow, dorsally smooth. Riparian and evergreen lowland forests, ca. 100 m; Amazonas (Isla Ratón, downstream from Santa Barbara, Trapichote). Amazonian Brazil, Bolivia. Psychotria sphaerocephala is similar to P. iodotricha of the flora area and to P. bremekampiana Steyerm. of French Guiana, Suriname, Brazil, and Bolivia. Psychotria bremekampiana can be recognized by its deeply bilobed stipules with the lobes often 2or 3-fid and/or erose and peduncles 3–20 mm long. The stipules of P. sphaerocephala are deeply laciniate similarly to those of some species of Rudgea, but appear to have been derived independently from those. These stipules are usually irregularly lacerate but occasionally have two groups of segments that suggested that they have two much-divided lobes. The stipules of P. bremekampiana are intermediate in form between those of P. sphaerocephala and those of other related species such as P. medusula, with two clearly defined lobes. The reportedly white fruits of P. sphaerocephala are unusual in Psychotria, but typical for Rudgea.

758

R UBIACEAE

Psychotria spiciflora Standl., Publ. Field Columbian Mus., Bot. Ser. 7: 115. 1930. —Psychotria spicata Müll. Arg., Flora 59: 550. 1876, hom. illeg., not Psychotria spicata Benth. 1841. [Subg. Heteropsychotria]. Psychotria vaupesana Standl. ex Steyerm., Acta Biol. Venez. 4: 101, fig. 50. 1964. Shrub to 4 m tall, glabrescent; leaves 10– 34 × 3.5–13 cm; petioles 5–20 mm long; stipules persistent, united around stem, sheath 0.5–1.5 mm long, lobes 2 per side, deltoid, 1.5–2.5 mm long; inflorescences terminal, spiciform, peduncle 0.8–3 cm long, branched portion 1.5–4.5 × 1–2 cm; floral bracts reduced; flowers sessile; calyx limb 0.3–0.5 mm long, dentate; corolla white, tube 4–5 mm long, lobes 1.5–2 mm long; fruit didymous, ca. 4 × 8 mm, white; pyrenes 2, ventrally with a slender longitudinal furrow, dorsally smooth or angled. Evergreen lowland forests, 50–200 m; Amazonas (basins of Río Casiquiare and Río Negro). Amazonian Colombia, Peru, and presumably adjacent Brazil. Psychotria spiciflora is distinctive due to the spiciform inflorescence. It is similar to P. huampamiensis C.M. Taylor of Amazonian northern Colombia, Ecuador, and Peru. Psychotria huampamiensis differs in its leaves usually very shiny adaxially (versus matte in P. spiciflora), its pedicels 0–3 mm long, and its slender inflorescence axes (versus quite stout in P. spiciflora). Although Steyermark classified P. spiciflora in his section Chytropsia together with the species here referred to Margaritopsis, P. spiciflora appears to be more closely related to the species he placed in his section Didymocarpos (e.g., P. cornigera, P. acuminata). Psychotria stipulosa Müll. Arg. in Mart., Fl. Bras 6(5): 334. 1881. —Cephaelis stipulosa (Müll. Arg.) Standl., Publ. Field Columbian Mus., Bot. Ser. 7: 430. 1931. [Subg. Heteropsychotria]. Psychotria homoplastica Moore, Trans. Linn. Soc. London 4: 378, pl. 26, figs. 1– 5. 1896. Subshrub, shrub, or small tree to 2(–5) m tall, puberulous to glabrescent; leaves 13–28 × 4–9 cm; petioles 5–40 mm long; stipules generally persisting on distalmost nodes, interpetiolar or shortly united around stem, ovate in outline, 10–20 mm long, acute to usually bilobed to 1/4 their length; inflores-

cence terminal, congested-cymose to paniculate, peduncle 1–6 cm long, branched portion 2.5–4.5 × 2.5–6 cm, bracts 8–25 mm long; flowers sessile; calyx limb ca. 0.8 mm long, undulate to dentate; corolla white, tube 8–11 mm long, lobes 3.5–4 mm long; fruit ellipsoid, ca. 4.5 × 4 mm, purple-black; pyrenes 2, ventrally with a longitudinal fissure, dorsally ridged. Evergreen lowland forests, granitic outcrops, 50–200 m; Amazonas (basins of Río Casiquiare, Río Guainía, and Río Orinoco). Eastern Colombia, Amazonian Brazil. ◆Fig. 588. Psychotria stipulosa is similar to P. capitata; for comments see the discussion under the latter species. Psychotria subundulata Benth., J. Bot. (Hooker) 3: 227. 1841. [Subg. Heteropsychotria]. Psychotria atabapoensis Standl., Publ. Field Columbian Mus., Bot. Ser. 8: 190. 1930. Shrub or small tree to 5 m tall, glabrous; leaves 12–20 × 3.5–11 cm; petioles 7–20 mm long; stipules persistent, united around stem, sheath ca. 1 mm long, lobes 2 per side, triangular, 0.5–1 mm long; inflorescences terminal, paniculate, peduncle 3.5–6 cm long, branched portion 4.5–6 × 4–5 cm, floral bracts ca. 0.5 mm long, green; flowers sessile; calyx limb 0.2–0.3 mm long, dentate; corolla white, tube 2–3 mm long, lobes ca. 1.8 mm long; fruit not seen, reportedly subglobose, ca. 2 × 3 mm, white; pyrenes unknown. Evergreen lowland to montane forests, 100–1500 m; Bolívar (Río Torono), Amazonas (basin of Río Casiquiare, Río Negro, and Río Orinoco). Táchira; Guyana, Suriname, French Guiana, Brazil. Psychotria subundulata can be recognized by the combination of its thin-textured leaves with the secondary veins extending delicately but clearly to unite with the margins, either directly or after reticulating once; its relatively short stipule lobes; and its inflorescences with the secondary axes dichotomously branched once into two usually straight, secund axes. The source of Steyermark’s (1972) report of this species from Bolivia is unknown. Psychotria tapirapecoana Steyerm., Mem. New York Bot. Gard. 23: 628. 1972. [Subg. Heteropsychotria].

Psychotria 759

Tree ca. 10 m tall, glabrous; leaves 10–18 × 2–5.5 cm; petioles 10–20 mm long; stipules apparently persistent, apparently united around stem, sheath ca. 3 mm long, lobes 2 per side, narrowly triangular, ca. 2.5 mm long; inflorescence terminal, paniculate, peduncle 5–6 cm long, branched portion ca. 4.5 × 3 cm, bracts 7–11 mm long, perhaps pale green or blue; flowers sessile; calyx limb 1– 1.2 mm long, lobed; corolla pale blue, tube ca. 7 mm long, lobes ca. 2.5 mm long; fruit unknown. Evergreen lowland forests, riparian forests, 100–200 m; Venezuelan-Brazilian border in Brazil (Río Castanho on Sierra Tapirapecó). Endemic. Psychotria tapirapecoana is known only from the type; based on the protologue it appears to generally resemble P. capitata, except with the stipules apparently persistent. Psychotria tatei Standl., Publ. Field Columbian Mus., Bot. Ser. 7: 460. 1931. —Psychotria everardii subsp. tatei (Standl.) Steyerm., Mem. New York Bot. Gard. 23: 540. 1972. [Subg. Heteropsychotria]. Psychotria breweri Steyerm., Acta Bot. Venez. 14(3): 25, fig. 14. 1984. Shrub to 7 m tall, glabrous; leaves 1.7–6.5 × 1.5–4.5 cm; petioles 2–7 mm long; stipules persistent, united around stem, sheath 0.8–1 mm long, lobes 2 per side, ligulate, 0.5–1 mm long; inflorescence terminal, paniculate, sessile, branched portion 3–3.5 × 3–4 cm, floral bracts reduced, pedicels 0–3 mm long; calyx limb 0.5–0.8 mm long, lobed; corolla white or yellow flushed with pink, tube 8–10 mm long, lobes ca. 4 mm long; fruit subglobose, 4–5 × 3–4 mm, dark blue; pyrenes apparently 2, ventrally with a central longitudinal fissure, dorsally ridged. Tepui scrub and meadows, 1200–2600 m; Amazonas (Cerro Duida, Cerro Marahuaka). Endemic. The small, stiff-textured leaves, small stipules, and sessile pyramidal inflorescences of Psychotria tatei are distinctive. It is similar to P. everardii (see comments there), P. phelpsiana, and P. carrenoi. Psychotria tepuiensis (Steyerm.) Steyerm., Mem. New York Bot. Gard. 23: 661. 1972. —Cephaelis tepuiensis Steyerm., Mem. New York Bot. Gard. 17(1): 430. 1967. [Subg. Heteropsychotria].

Shrub or tree to 6 m tall, glabrous; leaves 7–16.5 × 2.5–6.5 cm; petioles 7–15 mm long; stipules persistent, united around stem, sheath 5–10 mm long, lobes triangular to narrowly so, 6–13 mm long; inflorescences terminal, paniculate, peduncle 4–7.5 cm long, branched portion 5–7 × 4–8 cm, primary and secondary axes terminating in capitula, bracts 6–13 mm long; flowers sessile; calyx limb 6–7.5 mm long, denticulate; corolla white, tube 12–17 mm long, lobes 3.5–6 mm long; fruit subglobose to ellipsoid, 8–15 × 10–13 mm, becoming orange then red then purple; pyrenes 2, ventrally with a longitudinal furrow, dorsally shallowly ridged. Montane forests, 900–1800 m; Bolívar (Macizo del Chimantá, basins of Río Caroní, Río Cuyuní, and Río Yuruaní, Sierra de Lema), Amazonas (Cerro Duida, Sierra de la Neblina, Sierra Parima). Guyana, Peru, northern Brazil. ◆Fig. 582. Psychotria tepuiensis is similar to P. transiens. Psychotria transiens Wernham, J. Bot. 52: 314. Dec. 1914. —Cephaelis ernestii K. Krause, Notizbl. Bot. Gart. Berlin-Dahlem 6: 210. 1914, not Psychotria ernestii K. Krause 1909. [Subg. Heteropsychotria]. Shrub or small tree to 4 m tall, glabrous; leaves 4.5–16 × 1.5–5 cm; petioles 3–20 mm long; stipules persistent, united around stem, sheath 2–3 mm long, lobes narrowly triangular, 1–2.5 mm long; inflorescences terminal, corymbiform, peduncle 2.5–5 cm long, branched portion 1.5–5 × 2–6 cm, primary and secondary axes terminating in capitula, bracts 3.5–6 mm long, apparently green; flowers sessile; calyx limb 2.5–3 mm long, shallowly lobed; corolla white, tube 7–8 mm long, lobes 4–4.5 mm long; fruit subglobose to ellipsoid, ca. 15 × 10 mm, becoming red then black; pyrenes 2, ventrally with a longitudinal furrow, dorsally shallowly ridged. Tepui slope and summit forests, 1400–2100 m; Bolívar (Auyán-tepui, Macizo del Chimantá, Ptari-tepui, Roraima-tepui, Sororopán-tepui), Amazonas (Cerro Aratitiyope, Cerro Yutajé, Sierra de la Neblina, Sierra Parima). Guyana. Psychotria transiens is similar to P. tepuiensis. Their separation is outlined in the key to the species of subgenus Heteropsychotria. Psychotria mazaruniensis Standl. of

760

R UBIACEAE

Guyana and French Guiana is somewhat intermediate between these species, with the pyramidal inflorescences of P. tepuiensis, but stipules with sheaths 4–6 mm long and lobes 2–4 mm long, i.e., intermediate in size between these other two species. One specimen of P. transiens, Nee 31141 (MO!), from Sierra de la Neblina, is atypical in having two rather than one pair of secondary inflorescence axes. Psychotria trichotoma M. Martens & Galeotti, Bull. Acad. Roy. Sci. Bruxelles 11: 227. 1844. [Subg. Psychotria]. Shrub or small tree to 5 m tall, densely puberulous or hirtellous to glabrous; leaves 10–30 × 6–12.5 cm; petioles 1.5–4.5 cm long; stipules caducous, calyptrate (i.e., united into a conical cap), 6–25 mm long, the cap sharply acute; inflorescences terminal, paniculate, depending on interpretation sessile or with 2 or 3 fasciculate peduncles, peduncles or principal axes 2–4.5 cm long, bearing branched portions 2–8 × 3–10 cm, floral bracts reduced, pedicels 1–2.5 mm long; calyx limb 0.2–0.5 mm long, subtruncate; corolla white, tube ca. 4 mm long, lobes 2–2.5 mm long; fruit ellipsoid to subglobose, 6–10 × 4–7 mm, red, on pedicels to 6 mm long; pyrenes 2, dorsally ridged. Montane forests, ca. 1200 m; Amazonas (uppermost Río Orinoco near Brazilian border). Apure, Barinas, Mérida, Táchira, Yaracuy, Zulia; Mexico, Central America, Colombia, Ecuador, Peru, Bolivia. Psychotria trichotoma can be recognized by its calyptrate stipules, its relatively large leaves, its sessile or fasciculate inflorescences, and its relatively large fruits borne on relatively long pedicels. The fruits are notably variable in shape throughout the range of this species. Steyermark distinguished two varieties of this species in Venezuela based on variations in the distribution of pubescence; var. trichophylla Steyerm. was known from only one specimen from Barinas while var. trichotoma was considered common. However, pubescence is variable throughout the Mexican and Central American plants of Psychotria trichotoma and is not considered taxonomically informative there; consequently, Steyermark’s Venezuelan varieties are not recognized here.

Psychotria turbinella Müll. Arg. in Mart., Fl. Bras. 6(5): 374. 1881. [Subg. Heteropsychotria]. Venezuela, Brazil, Bolivia, 2 varieties, 1 in Venezuela. The small, pedunculate, capitate inflorescences with four rather small involucral bracts are distinctive for this species. See the discussion under Psychotria podocephala for comments on identification confusions in P. turbinella. Steyermark recognized two varieties of this species. He distinguished var. turbinella by its glabrous stems, petioles, peduncles, and leaf blades, versus at least some of these structures hirtellous in var. sororiella. Little material of this species is available, and the status of these varieties cannot be evaluated in the scope of this flora. P. turbinella var. sororiella (Müll. Arg.) Steyerm., Mem. New York Bot. Gard. 23: 677. 1972. —Psychotria sororiella Müll. Arg. in Mart., Fl. Bras. 6(5): 375. 1881. —Cephaelis sororiella (Müll. Arg.) Standl., Publ. Field Columbian Mus., Bot. Ser. 7: 429. 1931. [Subg. Heteropsychotria]. Subshrub or shrub to 2.5 m tall, hirtellous or pilosulous, sometimes becoming glabrescent; leaves 7–16.5 × 3–7.5 cm; petioles 3–7 mm long; stipules persistent, united around stem, sheath ca. 1 mm long, lobes 2 per side, narrowly triangular, 3–6 mm long; inflorescences terminal, capitate, peduncle 0.9–1.6 cm long, capitulum 0.8–1 × 2–3 m, involucral bracts usually 4, green or white to purple, 6– 15 mm long; flowers sessile; calyx limb 0.8–1 mm long, dentate; corolla white, tube ca. 5 mm long, lobes ca. 2 m long; fruit ellipsoid, ca. 5 × 3.5 mm, purple; pyrenes 2, ventrally with a central longitudinal ridge, dorsally ridged. Evergreen lowland and upland forests, 100–600 m; Amazonas (Cerro Aracamuni, basin of Río Siapa, near San Carlos de Río Negro). Western Brazil, Bolivia. Psychotria ulviformis Steyerm., Mem. New York Bot. Gard. 23: 638. 1972, not Psychotria alba Ruiz & Pav. 1799. —Tapogomea alba Aubl., Hist. Pl. Guiane 164, pl. 62, fig. 4. 1775. —Callicocca alba (Aubl.) J.F. Gmel., Syst. Nat. 5: 371. 1791. —Cephaelis alba (Aubl.) Willd.,

Psychotria 761

Sp. Pl. 1: 978. 1798. —Uragoga alba (Aubl.) Kuntze, Revis. Gen. Pl. 2: 959. 1891. —Gamotopea alba (Aubl.) Bremek., Recueil Trav. Bot. Néerl. 31: 294. 1934. [Subg. Heteropsychotria]. Geophila picta Rolfe, Kew Bull. 1896: 18. 1896, not Psychotria picta Wall. 1830. Cephaelis paraensis Standl., Publ. Field Columbian Mus., Bot. Ser. 8: 180. 1930, not Psychotria paraensis Müll. Arg. 1881. —Geophila paraensis Huber ex Standl., Publ. Field Columbian Mus., Bot. Ser. 8: 180. 1930, nom. nud., pro syn. Creeping herb, rooting at nodes, villous; leaves 3.5–10 × 2–8.5 cm; petioles 4–32 mm long; stipules persistent, united around stem or sometimes sub-interpetiolar, sheath 2–3 mm long, lobes 2 per side, narrowly triangular, 2–6 mm long; inflorescences terminal or occasionally pseudoaxillary, capitate or subcapitate, peduncles 1.5–4 cm long, capitulum 1–2.5 cm diameter, bracts 3.5–12 mm long; flowers sessile; calyx limb 1.5–2 mm long, deeply lobed; corolla cream, tube 5.5–6 mm long, lobes ca. 1.5 mm long; fruit ellipsoid, 4– 9 × 4–7 mm, blue; pyrenes 2, ventrally with a low longitudinal ridge, dorsally smooth. Riparian forests, lowland to upland forests, 100–800 m; Bolívar (basins of Río Aparurén, Río Carrao, Río Icabarú, and Río Supamo), Amazonas (basin of upper Río Orinoco to south of Puerto Ayacucho). Colombia, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil. ◆Fig. 562. Psychotria ulviformis is similar to P. variegata and to P. callithrix (Miq.) Steyerm. of Trinidad, Guyana, Suriname, French Guiana, Brazil, and Bolivia. These last two species can be separated from P. ulviformis by their leaves that are acute at the apex, pilosulous on the lower surface. Both P. ulviformis and P. variegata frequently have showy colorful foliage. The leaves of P. ulviformis have alternate zones of bronzy brown and lavender brown trichomes on their upper surfaces (Steyermark 1974, and in manuscript) and bright purple lower surfaces (Berry 4760, MO!). The inflorescences of Psychotria ulviformis have been described as “axillary” (Steyermark 1972; 1974; Boom and Delprete 2002), but in fact most of them are clearly terminal though usually overtopped by two

long stems, one produced in each axil of the leaves that subtend the peduncle (e.g., KnabVispo 221, MO!). A few specimens show a similar arrangement, but with only one axillary stem (e.g., Berry 4760, MO!); this condition is here interpreted as pseudoaxillary rather than truly axillary. Psychotria vareschii Steyerm., Mem. New York Bot. Gard. 23: 706, fig. 89. 1972. [Subg. Heteropsychotria]. Subshrub or shrub to 1 m tall, puberulous to glabrous; leaves 3–4 × 0.5–1.5 cm; petioles ca. 1 mm long; stipules persistent, united around stem, sheath 1–1.5 mm long, lobes 2 per side, triangular, ca. 0.5 mm long; inflorescences terminal, subcapitate to congested-cymose, peduncle 2–3.5 cm long, capitulum or branched portion 0.5–0.8 × 0.5–1 cm, bracts 1–2 mm long; flowers sessile; calyx limb ca. 0.8 mm long; corolla white, tube ca. 2.5 mm long, lobes ca. 2 mm long; immature fruit subglobose, to 3 mm diameter; pyrenes unknown. White-sand savannas, 50–200 m; Amazonas (basins of Río Atabapo, Río Casiquiare, and Río Guainía). Endemic. The relatively small elliptic-oblong leaves with their margins revolute and their venation prominulous on the upper surfaces are distinctive for Psychotria vareschii. Psychotria variegata Steyerm., Mem. New York Bot. Gard. 23: 638. 1972, not Psychotria purpurea Merrill & Perry 1946. —Tapogomea purpurea Aubl., Hist. Pl. Guiane 162, pl. 62, fig. 3, 1775. —Callicocca purpurea (Aubl.) J.F. Gmel., Syst. Nat. 1: 371. 1791. —Cephaelis purpurea (Aubl.) Willd., Sp. Pl. 1: 978. 1798. —Uragoga purpurea (Aubl.) Kuntze, Revis. Gen. 2: 962. 1891. —Gamotopea purpurea (Aubl.) Bremek., Recueil Trav. Bot. Néerl. 31: 294. 1934. [Subg. Heteropsychotria]. Cephaelis surinamensis Standl., Publ. Field Columbian Mus., Bot. Ser. 4: 335. 1929, not Psychotria surinamensis Bremek. 1934. —Gamotopea surinamensis (Standl.) Bremek., Recueil Trav. Bot. Néerl. 31. 294. 1934. Creeping herbs, densely villous to hirtellous; leaves 2.5–6 × 1–3 cm; petioles 3–10 mm long; stipules persistent, united around stem or sometimes sub-interpetiolar, sheath

762

R UBIACEAE

2–3 mm long, lobes 2 per side, narrowly triangular, 3–5 mm long; inflorescences terminal, capitate, peduncle 4–15 mm long, capitulum 0.8–1 × 1–2 cm, bracts 6–9 mm long; flowers sessile; calyx limb 1.2–1.5 mm long, deeply lobed; corollas white to pale yellow, tube 5–7 mm long, lobes 2–2.5 mm long; fruit subglobose, 4–7 × 2–7 mm, bright blue; pyrenes 2, ventrally with a low longitudinal ridge, dorsally smooth. Lowland to upland forests, also on soils derived from igneous rocks, 100–1100 m; Bolívar (basins of Río Acanán, Río Caroní, Río Icabarú, and Río Paragua), Amazonas (basins of Río Orinoco and Río Ventuari). Guyana, Suriname, French Guiana, Brazil, Amazonian Bolivia. Steyermark (1974, and in manuscript) and collectors noted that the leaves of Psychotria variegata often have the upper surface dark green except for a silvery or gray longitudinal stripe down the middle and the lower surfaces bright purple. However, the leaves of this species are sometimes plain green; the significance and genetics of this variation are unknown. Psychotria vasivensis (Müll. Arg.) Standl., Publ. Field Columbian Mus., Bot. Ser. 7: 461. 1931. —Mapouria vasivensis Müll. Arg., Flora 59: 460, 466. 1876. [Subg. Heteropsychotria]. —Carriso de picure. Shrub or small tree to 3 m tall, glabrous; leaves 10–31 × 3.5–14 cm; petioles 2–7.5 cm long; stipules persistent with leaves, interpetiolar, ovate to triangular, 3–11 mm long, obtuse to rounded or truncate; inflorescences terminal, paniculate, peduncle 6–11.5 cm long, branched portion 3.5–6.5 × 5–8 cm, floral bracts 3–9 mm long, pedicels 0–1 mm long; calyx limb 3–3.5 mm long, truncate to irregularly dentate; corolla white, tube 5–6 mm long, lobes 4–5 mm long; fruit narrowly ellipsoid to lanceoloid, 7–10 × 4–5 mm, lavender to purple; pyrenes 2, dorsally plane, dorsally ridged. Riparian forests, 100–200 m; Amazonas (Caño Emoni in Río Siapa basin, upper Río Temi, Tamatama). Endemic. ◆Fig. 587. Psychotria vasivensis may be included in the segregate genus Carapichea when it is separated from Psychotria, but the limits of this group are not yet entirely clear. Psychotria vasivensis is similar to P. maturacensis, P. wurdackii, and P. franquevilleana.

Psychotria vellosiana Benth., Linnaea 23: 464. 1850, not Psychotria sessilis Vell. 1825 [1829]. —Coffea sessilis Vell., Fl. Flum. 64, t. 20. 1825 [1829]. —Psychotria sessilis (Vell.) Müll. Arg. in Mart., Fl. Bras. 6(5): 358. 1881, nom. illeg., non Psychotria sessilis Vell. 1825 [Subg. Heteropsychotria]. Psychotria hancorniifolia Benth., Linnaea 23: 463. 1850, “hancorniaefolia.” Psychotria hancorniifolia var. angustifolia Müll. Arg. in Mart., Fl. Bras. 6(5): 356. 1881. —Psychotria sessilis var. angustifolia (Müll. Arg.) Steyerm., Mem. New York Bot. Gard. 23: 716. 1972, nom. illeg., not Psychotria sessilis Vell. 1825 [1829]. Psychotria hancorniifolia var. longifolia Müll. Arg. in Mart., Fl. Bras. 6(5): 356. 1881. Psychotria sororopanensis Standl. & Steyerm., Fieldiana, Bot. 28: 605. 1953. Shrub or small tree to 5 m tall, densely hirtellous to glabrous; leaves 3.5–12 × 0.5–3 cm; petioles 1–6 mm long; stipules persistent, united around stem, sheath 2–3 mm long, lobes 2 per side, linear, 1–2.5 mm long; inflorescences axillary, capitate to subcapitate, peduncle 0.4–6 cm long, capitulum 0.3–0.7 × 0.6–0.8 cm, external bracts 1.5–3 × 1 mm; flowers sessile; calyx limb 0.5–1 mm long, dentate; corolla white, tube 3.5–5 mm long, lobes 3.5–4 mm long; fruit ellipsoid to subglobose, 3.5–4 × 3–4 mm, blue to black; pyrenes 2, dorsally ridged, ventrally with a central longitudinal furrow. Lowland to montane forests, 100–2300 m; Bolívar (Auyántepui, Perai-tepui, Sororopán-tepui), Amazonas (Río Castanho tributary of Río Padauiri on the Brazilian border). Guyana, Peru, Brazil. ◆Fig. 589. Psychotria vellosiana can be recognized by its axillary inflorescences and well marked, closely set tertiary leaf venation. Vegetatively it is difficult to distinguish from Psychotria speluncae. Psychotria vellosiana is infrequently collected in the flora area but quite common in southeastern Brazil. The names P. hancorniifolia and P. vellosiana were published simultaneously; the name P. vellosiana is chosen here based on the extensive previous usage of this name by Kirkbride (in herbaria). Steyermark (1972) recognized four varieties of this species, three in Brazil and var. angustifolia found from the flora area to

Psychotria 763

southeastern Brazil. These were separated by details of the pubescence and leaf size and shape. However, this species is quite variable in these features, as documented carefully by Steyermark (1972, 716), so the taxonomic recognition of this variation does not seem useful and is not done here. Psychotria venezuelensis Steyerm., Mem. New York Bot. Gard. 23: 493, fig. 69. 1972. [Subg. Heteropsychotria]. Shrub to 2 m tall, glabrous; leaves 7–12.5 × 2–4.5 cm; petioles 2–8 mm long; stipules persistent, united around stem, sheath 1–2 mm long, truncate, on each interpetiolar side with 2 groups of glands inserted near sheath base; inflorescences terminal, paniculate, peduncle 1.5–3.5 cm long, branched portion 4–6 × 2.5–3 cm, bracts ca. 1 mm long, apparently green; flowers sessile; calyx limb ca. 0.7 mm long, dentate; corolla white, tube 6.5–7 mm long, lobes ca. 3 mm long; fruit ellipsoid, ca. 6 × 4 mm, blue-black; pyrenes 2, ventrally with a shallow central furrow, dorsally smooth. Montane forests, sandstone escarpment slopes and border of forested canyons, 1600– 2000 m; Amazonas (Sierra de la Neblina). Endemic. The combination of the rather small, stifftextured leaves, truncate stipules with glands inserted near the sheath base, 4merous flowers in dense glomerules, and dorsally smooth pyrenes is distinctive in Psychotria venezuelensis. This poorly known species probably belongs to Rudgea, as shown by its stipules with groups of caducous glands inserted near their bases. It is also keyed in Rudgea but provisionally treated in Psychotria pending its further study. Steyermark (1972) grouped it with Psychotria ventuariana, which is clearly a species of Rudgea, and with several species that are probably referable to Margaritopsis, though the fruits of P. venezuelensis are blue-black rather than red like these others. Steyermark examined material of this species with the calyx and corolla 4-lobed. There is often variation in this feature in Psychotria subgenus Heteropsychotria species, but all the flowers seen of P. venezuelensis are indeed 4-merous. Psychotria ventuariana Standl. & Steyerm., Fieldiana, Bot. 28: 609. 1953. [Subg. Heteropsychotria].

Tree to 10 m tall, glabrous; leaves 11–16.5 × 3–7 cm; petioles 6–11 mm long; stipules persistent, shortly united around stem, 1–3 mm long, apically with a dense group of caducous glands ca. 1 mm long; inflorescences terminal, paniculate, peduncle 2.5–3 cm long, branched portion 4–5.5 × 3–3.5 cm, with secondary axes 3–4 at basal node, floral bracts ca. 0.5 mm long; flowers sessile; calyx limb in bud ca. 0.5 mm long, undulate; corolla in bud with color unknown, to 3 mm long; fruit unknown. Riparian forests, lowland forests, 50–200 m; Amazonas (Río Ventuari at Raudal Trapichote). Barinas. Psychotria ventuariana is a poorly known species that appears to belong to Rudgea, as shown by its stipules that terminate in a group of caducous glands ca. 1 mm long. It is also keyed in Rudgea but treated here in Psychotria pending further study. Psychotria venulosa Müll. Arg. in Mart., Fl. Bras. 6(5): 294. 1881. —Psychotria deflexa subsp. venulosa (Müll. Arg.) Steyerm., Mem. New York Bot. Gard. 23: 503. 1972. [Subg. Heteropsychotria]. —Bujiyuju (Bale), Cadajo-como-mamujei-du (Yanomamo), Cafecillo, Chuta, Viejita. Subshrub or shrub to 1 m tall, glabrous; leaves 5–16 × 1–6 cm; petiole 2–8 mm long; stipules persistent, united around stem, sheath 1–2.5 mm long, lobes 2 per side, triangular, 1.5–5 mm long; inflorescence terminal, paniculate, peduncle 1–3 cm long, branched portion 3–8 × 3–5 cm, floral bracts reduced, pedicels 0–1 mm long; calyx limb ca. 0.2 mm long, undulate; corolla white, tube 2.5–3 mm long, lobes ca. 1 mm long; fruit subglobose, ca. 3 mm diameter, white; pyrenes 2, ventrally smooth or with a longitudinal cavity that sometimes extends to a complete furrow, dorsally ridged and with the surface alveolate. Evergreen lowland to lower montane forests, 100–500 m; Bolívar (base of Amaruay-tepui, base of Guaiquinima-tepui, Río Caura basin), Amazonas (basins of Río Caura, Río Negro, and Río Orinoco, base of Sierra de la Neblina). Colombia, Guyana, Peru, Brazil. The stipules with the sheath elongating behind the lobes on older nodes and the tertiary leaf venation generally subparallel and raised on the lower surface are distinctive for Psychotria venulosa. It is similar to

764

R UBIACEAE

Psychotria deflexa. See comments under that species regarding their separation and historical treatment. Steyermark (in manuscript) reported that the fruit of P. venulosa is the source of a blue dye in the flora area. Psychotria vichadensis Standl., Field Mus. Nat. Hist., Bot. Ser. 22: 207. 1940. [Subg. Heteropsychotria]. Subshrub or shrub to 1 m tall, puberulous to glabrous; leaves 1–5.5 × 0.3–2 cm; petioles 1–4 mm long; stipules persistent, united around stem, sheath 0.5–1 mm long, lobes 2 per side, narrowly triangular, 0.5–1 mm long; inflorescence terminal, capitate, peduncle 0.1–0.3 cm long, capitulum 0.3–0.5 × 0.5 cm, bracts green, 0.5–5 mm long; flowers sessile; calyx limb 0.5–0.8 mm long, dentate; corolla white to cream, tube ca. 3.5 mm long, lobes ca. 1 mm long; fruit ellipsoid, 2.5–3 × 2.5–3 mm, black; pyrenes 2, ventrally with a central longitudinal fissure, dorsally ridged. Semideciduous to evergreen lowland forests, 100–300 m; Bolívar (Río Paragua), Amazonas (vicinity of Isla Ratón, Río Coro Coro west of Cerro Yutajé, Río Orinoco, Río Parú). Colombia. ◆Fig. 576. The relatively low slender stature and small leaves, stipules, and inflorescences are distinctive for Psychotria vichadensis. It is frequently reported to form dense understory colonies. It is similar to P. hoffmannseggiana, and because of some of the details of Steyermark’s key (1972; 1974) has sometimes also been confused with P. spadicea. Psychotria viridis Ruiz & Pav., Fl. Peruv. 2: 61, t. 210, fig. B. 1799. [Subg. Psychotria]. Shrub or tree to 4 m tall, glabrous; leaves 6.5–15 × 2.5–5 cm; petioles 3–10 mm long; stipules caducous, interpetiolar, ovate, 10–20 mm long, acuminate to bilobed; inflorescences terminal, spiciform, peduncle 3–8 cm long, axis or branched portion 4–10 × 3–8 cm, floral bracts 1–2 mm long; calyx limb 0.3–0.5 mm long, subtruncate; corolla white, tube ca. 1.5 mm long, lobes ca. 1 mm long; fruit ellipsoid, ca. 5 × 4 mm, red; pyrenes 2, dorsally ridged. Upland forests, ca. 700 m; Amazonas (Sierra Parima). Central America, Colombia, Ecuador, Peru, Brazil, Bolivia. Psychotria viridis contains a hallucinogenic alkaloid, and is sometimes used by indigenous peoples as an admixture in the tra-

ditional hallucinogenic mixture ayahuasca, based on Banisteriopsis caapi in the Malpighiaceae. Psychotria wurdackii Steyerm., Mem. New York Bot. Gard. 23: 576, fig. 77. 1972. [Subg. Heteropsychotria]. Subshrub or shrub to 2.5 m tall, pilosulous to glabrous; leaves 10–26 × 4–10 cm; petioles 15–20 mm long; stipules persistent, united around stem, sheath 2–4 mm long, truncate to broadly emarginate or lobed to 1/2 their length, lobes ligulate; inflorescences terminal, paniculate, peduncle 3–6 cm long, branched portion 3–6 × 3–5 cm, bracts 4.5–9 mm long, pale green; flowers sessile; calyx limb 0.8–1.2 mm long, dentate; corolla white, tube 6.5–7 mm long, lobes ca. 3 mm long; fruit ellipsoid, ca. 4 × 3 mm, color unknown; pyrenes 2, ventrally with a central longitudinal furrow, dorsally ridged. Montane forests usually overlying sandstone of tepuis and in evergreen seasonally dry mountains overlying igneous and quartzitic substrate, 1000– 1700 m; Amazonas (Cerro Camani, Cerro Duida, saddle between Cerro Duida and Cerro Marahuaka, Sierra Parima, Cerro Parú, Sierra de la Neblina). Ecuador, Peru, Brazil. ◆Fig. 565. Psychotria wurdackii is similar in general aspect to P. vasivensis, P. franquevilleana, P. maturacensis, and P. cerronis. At least the first three of these may be included in the segregate genus Carapichea when it is fully delimited and separated from Psychotria, but P. cerronis and P. wurdackii will more likely remain in Psychotria. Psychotria wurdackii is also similar to P. silvae Steyerm. of Pará, Brazil, which may not actually be distinct; Steyermark separated this by its “outer” inflorescence bracts glabrous adaxially (versus pilosulous in P. wurdackii), its tertiary leaf venation “prominent both sides” (sic; versus “faint, not prominently elevated” in P. wurdackii), and its inflorescence axes 8–13 (versus 5–18 in P. wurdackii). Psychotria yapacanensis Steyerm., Pittieria 9: 10. 1981. [Subg. Heteropsychotria]. Shrub to 2 m tall, puberulous to pilosulous; leaves 5.5–9 × 2.3 cm; petioles 1–3 mm long; stipules apparently persistent, united around stem, sheath length unknown, lobes

Psychotria 765

2 per side, linear, 4.5–5 mm long; inflorescence terminal, apparently cymose, peduncle ca. 1.5 cm long, branched portion ca. 1 × 1 cm, floral bracts ca. 0.7 mm long, perhaps green; flowers sessile; calyx limb 0.6–0.7 mm long, dentate; corolla yellow-green, tube ca. 4 mm long, lobes ca. 1.5 mm long, with abaxial appendages of unknown length; fruit subglobose, 5–7 mm diameter, color unknown; pyrenes 2, ventral face unknown, dorsally ridged. White-sand savannas, shrub formation, 100–200 m; Amazonas (base of Cerro Yapacana). Endemic. ◆Fig. 578. Psychotria yapacanensis is said in its protologue to be distinguished by its secondary leaf veins 12–17 pairs and its puberulous to pilosulous pubescence. It is apparently generally similar to P. anartiothrix. Psychotria yavitensis Steyerm., Mem. New York Bot. Gard. 23: 456. 1972. [Subg. Psychotria]. Tree to 12 m tall, glabrous; leaves 15–19 × 6–9.5 cm; petioles 2.5–4 cm long; stipules caducous, interpetiolar, obovate to ligulate, ca. 12 mm long, rounded; inflorescences terminal, paniculate, peduncle 2.2–4.5 cm long,

branched portion 6–8.5 × 4–7 cm, floral bracts perhaps ca. 0.5 mm long, pedicels 0.5– 1 mm long; calyx limb ca. 1 mm long, subtruncate; corolla white, tube ca. 2.5 mm long, lobes ca. 3 mm long; fruit unknown. Evergreen lowland forests, 100–200 m; Amazonas (Río Guainía basin). Endemic. Psychotria yavitensis is similar to P. mapourioides, but Steyermark separated it from that species by its secondary leaf veins 5 or 6 pairs and “more widely separate,” its shorter peduncles, its “more compactly flowered and shorter” inflorescences, and the corolla throat “more conspicuously barbate.” In Steyermark’s circumscription, P. mapourioides has 6–15 pairs of secondary leaf veins that are more closely set, peduncles 2.8–13 cm long, and inflorescences with the branched portion 3–10 cm long, so these two species in fact overlap in these features. Psychotria yavitensis probably represents only a combined extreme form of the variation in several morphological characters, but because no material of this species has yet been seen by Taylor it is maintained provisionally here pending further study.

Fig. 559. Psychotria capitata subsp. inundata

766

R UBIACEAE

Fig. 560. Psychotria hemicephaelis

Fig. 561. Psychotria amplectans

Fig. 562. Psychotria ulviformis

Psychotria 767

Fig. 563. Psychotria coussareoides

Fig. 564. Psychotria spadicea

Fig. 565. Psychotria wurdackii

768

R UBIACEAE

Fig. 566. Psychotria durifolia

Fig. 567. Psychotria duricoria var. longiloba

Fig. 568. Psychotria everardii Fig. 569. Psychotria campylopoda

Psychotria 769

Fig. 570. Psychotria casiquiaria

Fig. 571. Psychotria crocochlamys

770

R UBIACEAE

Fig. 572. Psychotria jauaensis

Fig. 573. Psychotria humboldtiana

Fig. 574. Psychotria imthurniana

Psychotria 771

Fig. 575. Psychotria lupulina subsp. lupulina

Fig. 576. Psychotria vichadensis

Fig. 578. Psychotria yapacanensis Fig. 577. Psychotria iodotricha

772

R UBIACEAE

Fig. 579. Psychotria phaneroloma

Fig. 580. Psychotria phelpsiana

Fig. 581. Psychotria medusula

Psychotria 773

Fig. 582. Psychotria tepuiensis

Fig. 583. Psychotria prunifolia

Fig. 584. Psychotria poeppigiana subsp. poeppigiana

774

R UBIACEAE

Fig. 585. Psychotria podocephala

Fig. 587. Psychotria vasivensis

Fig. 586. Psychotria speluncae

Psychotria 775

Fig. 588. Psychotria stipulosa

Fig. 589. Psychotria vellosiana Fig. 590. Psychotria polycephala

776

R UBIACEAE

68. RANDIA L., Sp. Pl. 1192. 1753. Basanacantha Hook. f. in Benth. & Hook. f., Gen. Pl. 2: 82. 1873. by Julian A. Steyermark Trees or shrubs, terrestrial, unarmed or frequently armed with axillary or supra-axillary spines, frequently with the flowers and leaves borne at the ends of short lateral stems. Leaves opposite though sometimes closely grouped and appearing whorled or fasciculate, sometimes slightly to markedly anisophyllous, sessile or petiolate, venation not lineolate; stipules interpetiolar and sometimes shortly united around the stem, persistent, triangular, generally imbricate to valvate in bud. Inflorescences terminal, axillary, or cauliflorous, ebracteate, the staminate flowers solitary to several and cymose to fasciculate, the pistillate flowers solitary. Flowers small to large, often showy, often nocturnal and fragrant, unisexual (the plants dioecious), sessile or pedicellate. Staminate flowers: hypanthium reduced; calyx limb truncate, denticulate, or shortly to deeply lobed, teeth or lobes 5 or 6, without calycophylls; corolla salverform to funnelform, white becoming yellow with age, internally usually pubescent at least in throat, lobes 5 or 6, convolute in bud; stamens 5 or 6, inserted in upper part of corolla tube; anthers narrowly oblong, dorsifixed or basifixed, subsessile, included or partially exserted; pistillode present, similar to style and stigma, included or exserted. Pistillate flowers: hypanthium turbinate to ellipsoid or obovoid; calyx limb and corolla similar to the staminate or usually a little larger; staminodes present, similar to though smaller than stamens; ovary 1-locular; ovules numerous, on parietal placentas. Fruit baccate, subglobose to ellipsoid, smooth to longitudinally ridged or winged, yellow, orange, or brown, endocarp coriaceous to woody. Seeds subcircular to ellipsoid or angled, usually compressed, small to medium-sized, smooth, embedded in usually orange to brown pulp. New World tropics and subtropics: southeastern U.S.A., Mexico, the Antilles, Central America, South America except Chile and Uruguay; ca. 70 species, 12 in Venezuela, 7 of these in the flora area. Since Steyermark (1972, 325–343; 1974) reviewed this genus, the circumscription of Randia has been modified. Steyermark considered Randia a pantropical genus of 200–300 species, but more recent authors have restricted it to a smaller group of neotropical species. Randia in its old, broadly pantropical circumscription comprised a diverse group of species that are now treated in nearly a dozen smaller genera, including the neotropical genus Rosenbergiodendron and the paleotropical genera Aidia Lour. and Catunaregam Tirv. Randia sensu stricto differs from similar neotropical genera in its unisexual flowers (plants dioecious), parietal placentation, and pollen in tetrads. The systematics of this genus (e.g., Gustafsson 2000) as well as its placement in a tribe (e.g., Persson 2000b) are as yet poorly known, and several species circumscriptions and names of Randia in the flora area are likely to change when this group is studied in detail. Key to the Species of Randia 1. 1. 2(1). 2. 3(2). 3.

Plants unarmed; leaves 20–29 cm long .............................. R. amazonasensis Plants unarmed or usually armed; leaves 0.4–15 cm long ...................... 2 Calyx lobes elliptic, 5–25 mm long ................................................. R. armata Calyx lobes triangular to narrowly elliptic, 0.4–10 mm long .................. 3 Calyx lobes densely pubescent on both surfaces .............................. R. dioica Calyx lobes glabrous to puberulous internally or on both surfaces......... 4

Randia 777

4(3). 4. 5(4). 5. 6(5). 6.

Fruits, hypanthium, and corolla exterior of both staminate and pistillate flowers pilosulous to hirtellous .............................................. R. hebecarpa Fruits, hypanthium, and corolla exterior of both staminate and pistillate flowers glabrous to puberulous or strigillose ....................................... 5 Fruits 22–27 mm long; calyx lobes 3.5–7 mm long ...................... R. brevipes Fruits 7–20 mm long; calyx lobes 0.4–3.5 mm long ................................. 6 Leaves closely grouped and often apparently fasciculate; calyx lobes 0.4– 2 mm long; corolla tube 2.5–7 mm long .................................... R. aculeata Leaves loosely grouped or distributed along stems, usually clearly paired; calyx lobes 2–3.5 mm long; corolla tube 7–10 mm long ...... R. venezuelensis

Randia aculeata L., Sp. Pl. 1192. 1753. U.S.A. (Florida), Mexico, Central America, Antilles, Colombia, Venezuela, Trinidad; 3 varieties, 2 in Venezuela, 1 of these in the flora area. Steyermark distinguished the variety found in the flora area by its pubescent lower leaf surfaces, calyx tube, and hypanthium; the varieties of Randia aculeata recognized by him have not been collected in the Guayana region. There is wide morphological variation in this species throughout its range, though, making the separation of infraspecific taxa controversial. R. aculeata var. dasyphylla Steyerm., Mem. New York Bot. Gard. 23: 342. 1972. Shrub to 3 m tall, with 2 or more spines along each of the short lateral stems; leaves 0.4–11.5 × 0.2–5.2 cm; staminate flowers 1–3; calyx limb with tube 0.5–2 mm long, lobes 5 or 6, 0.4–2 mm long; corolla with tube 2.5–7 mm long, lobes 5 or 6, 3–7 mm long; fruit 7– 17 × 5–15 mm. Savannas, scrub vegetation, 300–500 m; Bolívar (extreme northeastern portion). Carabobo, Miranda, Yaracuy, Nueva Esparta; Colombia, Trinidad. Randia amazonasensis Steyerm., Mem. New York Bot. Gard. 23: 336, fig. 60. 1972. Basanacantha spinosa f. macrophylla K. Schum. in Mart., Fl. Bras. 6(6): 377. 1998, not Randia macrophylla Benth. & Hook. 1873, not Randia macrophylla C. Moore 1895. Tree to 8 m tall, unarmed; leaves 20–29 × 6–7.5 cm; staminate flowers 1 or 2; calyx limb with tube ca. 0.5 mm long, lobes 5, 4–5 mm long; corolla with tube ca. 17 mm long, lobes 5, ca. 9 mm long; fruit 30–35 × 25 mm. Ripar-

ian forests, evergreen lowland forests, 50– 200 m; Amazonas (basins of Río Casiquiare, Río Orinoco, and Río Ventuari). Endemic. Randia armata (Sw.) DC., Prodr. 4: 387. 1830. —Mussaenda spinosa Jacq., Select. Stirp. Amer. Hist. 70, t. 49. 1763. —Gardenia armata Sw., Prodr. 51. 1788, not Gardenia spinosa Thunb. 1780, not Gardenia spinosa L. f. 1782. —Randia spinosa (Jacq.) H. Karst., Fl. Columb. 2: 128. 1869, hom. illeg., not Randia spinosa (Thunb.) Poir. 1812. —Basanacantha spinosa (Jacq.) K. Schum. in Mart., Fl. Bras. 6(6): 376. 1889. —Jazmín de monte, Zajarito montañero. Tree or shrub to 8 m tall, with spines 3 or 4 at apices of stems; leaves 3.5–11 × 2–6 cm; staminate flowers 1–3; calyx limb with tube 0.8–1.5 mm long, lobes 5, 5–25 mm long; corolla with tube 20–30 mm long, lobes 5, 10– 16 mm long; fruit 20–25 × 15–30 mm. Deciduous and semideciduous forests, riparian forests, 50–300 m; Delta Amacuro (lower Río Orinoco, Serranía de Imataca), Bolívar (Altiplanicie de Nuria, Cerro Bolívar, lower Río Caroní, lower Río Caura, lower Río Paragua, Serranía de Imataca), Amazonas (north of Puerto Ayacucho). Miranda, Sucre, Yaracuy, Zulia; widespread in Mexico, Central America, the West Indies, and perhaps through South America. ◆Fig. 591. The name Randia spinosa has periodically been incorrectly resurrected for this species, but this combination is precluded by an earlier publication of the same name for an African species (Taylor and Lorence 1993). The name R. armata has been applied widely to plants of Central America and South America and the Antilles, but the plants treated under this name are heterogeneous and probably not all conspecific.

778

R UBIACEAE

Fig. 591. Randia armata

Randia brevipes Steyerm., Acta Bot. Venez. 6: 126. 1971. —Jazmín de monte, Punteral. Shrub or small tree to 8 m tall, with spines borne in pairs below tips of older branches; leaves 5–6.5 × 2.5–4 cm; staminate flowers 1 or 2; calyx limb with tube ca. 2 mm long, lobes 5, 3.5–7 mm long; corolla with tube 18–20 mm long, lobes 5, 8–10 mm long; fruit 22–27 × 15–18 mm. Along streams in deciduous forests, thickets in deciduous forests, 100–500 m; Bolívar (Altiplanicie de Nuria, lower Río Caroní). Zulia; Trinidad.

Carabobo, Distrito Federal, Falcón, Guárico, Lara, Miranda, Zulia; adjacent Colombia. Steyermark (1972; 1974) cited the name Randia armata var. pubescens (Kunth) Knuth as a synonym of R. dioica. This name is based on a type from Guayaquil on the coast of Ecuador, and was applied by Schumann in the Flora Brasiliensis to the Venezuelan plants as well. However, R. dioica has not yet been confirmed from Ecuador, so it seems unlikely at present that these names could apply to the same species and these are here considered separate names.

Randia dioica H. Karst., Linnaea 30: 153. 1850; Fl. Columb. 2: 127, pl. 167, fig. 1. 1869. —Basanacantha dioica (H. Karst.) Hook.f. in Benth. & Hook.f., Gen. Pl. 2: 83. 1873. —Cruceto, Punteral negro. Randia caracasana Standl., Publ. Field Columbian Mus., Bot. Ser. 8: 57. 1930. Shrub or small tree to 10 m tall, with 2–4 spines borne at or below the ends of the stems; leaves 3.5–15 × 2–8.5 cm; staminate flowers 1–5; calyx limb with tube ca. 1.5 mm long, lobes 5, 3–10 mm long; corolla with tube 7–14 mm long, lobes 5, 3.5–7 mm long; fruit 15–25 × 13–18 mm. Deciduous forests, ca. 300 m; Bolívar (east of El Callao). Aragua,

Randia hebecarpa Benth., J. Bot. (Hooker) 3: 213. 1841. —Basanacantha hebecarpa (Benth.) Hook. f. in Benth. & Hook. f., Gen. Pl. 2: 82. 1873. —Cafecito, Cruceta real, Pajarito. Shrub or small tree to 2 m tall, usually with paired slender spines borne at ends of short lateral stems; leaves 1.5–5 × 0.8–1.8 cm; staminate flower solitary; calyx limb with tube ca. 0.5 mm long, lobes 5, ca. 3 mm long; corolla with tube 7–8 mm long, lobes 5, 7–7.5 mm long; fruit 12–14 × 10–12 mm. Borders of savannas, gallery forests, 50–100 m; Bolívar (lower Río Caura and lower and middle Río Orinoco toward Maripa and Cai-

Remijia 779

cara). Anzoátegui, Barinas, Cojedes, Guárico, Monagas, Yaracuy; Colombia, Guyana, Brazil. Randia venezuelensis Steyerm., Bol. Soc. Venez. Ci. Nat. 29(119–120): 23. 1971. —Barba de tigre, Cachito, Cruceta, Punteral. Shrub to 2.5 m tall, with 1–4 stout spreading spines borne at and slightly below the tips of stems; leaves 3–9 × 1.5–4.5 cm; flowers 1–5; calyx limb 0.5–1.5 mm long, lobes 5,

2–3.5 mm long; corolla with tube 7–10 mm long, lobes 5, 4–6 mm long; fruit 15–20 × 16– 17 mm. Shrub islands in savannas, periodically flooded savannas, low gallery forests, around igneous outcrops, 50–200 m; Bolívar (northwestern portion in basin of middle Río Orinoco and south of Caicara), Amazonas (basin of Río Manapiare, upper Río Orinoco at El Motin, Río Orinoco south of Puerto Ayacucho). Apure, Barinas, Guárico, Lara, Portuguesa, Yaracuy; Colombia.

69. REMIJIA DC., Biblioth. Universelle Sci., Sci. Arts 41: 155. 1829. Cephalodendron Steyerm., Mem. New York Bot. Gard. 23: 228. 1972. by Charlotte M. Taylor and Julian A. Steyermark Trees or shrubs, terrestrial, unarmed, sometimes resinous, sometimes monocaulous, often symbiotic with ants that are housed in the stems or infrequently the leaf bases. Leaves opposite or 3- or 4-verticillate, sometimes very large, sessile to petiolate, venation not lineolate; stipules interpetiolar, caducous, triangular to lanceolate, in bud generally erect and flatly appressed, often very large. Inflorescences axillary, capitate to cymose, subsessile to usually pedunculate, bracteate or bracts reduced, with flowers solitary or usually multiflowered, paniculate, racemiform, or capitate. Flowers medium to large, sessile to pedicellate, fragrant, often distylous. Hypanthium turbinate to ellipsoid. Calyx limb truncate to spathaceous or lobed, lobes 4–6, without calycophylls; corolla salverform, white to pink or red, internally glabrous or variously pubescent, lobes 4–6, valvate in bud. Stamens 4–6, inserted at or below the middle of the corolla tube; anthers narrowly oblong, dorsifixed near base, included or partially exserted. Ovary 2-locular; ovules numerous in each locule, on axile placentas. Fruit capsular, cylindrical to fusiform, ovoid, or oblong, septicidal from apex or base and sometimes the valves later splitting, chartaceous to woody, smooth. Seeds flattened, papery, small, fusiform to elliptic, marginally winged. Colombia, Venezuela, Guyana, Ecuador, Peru, Brazil, Bolivia (most species in Venezuela and Brazil): ca. 45 species, 23 species in Venezuela, all in the flora area. Steyermark distinguished Cephalodendron from Remijia only by its capitate inflorescences. No doubt his idea of Cephalodendron’s distinctness was reinforced by its corollas that are pubescent internally. Steyermark considered this character taxonomically informative in Remijia, though it varies in a number of Rubiaceae genera. However Andersson (1995) in his morphological study of the tribe Cinchoneae, concluded that Cephalodendron is a synonym of Remijia, though without making any nomenclatural combinations for its species. Key to the Species of Remijia 1. 1. 2(1). 2. 3(2). 3.

Petiole conspicuously inflated; inflorescence sessile ...................... R. sessilis Petiole not inflated; inflorescence pedunculate ........................................ 2 Leaves 3- or 4-verticillate .......................................................................... 3 Leaves opposite .......................................................................................... 6 Leaves 1–2.5 cm wide, > 5 times as long as wide ........................ R. delascioi Leaves 3–35 cm wide, < 3 times as long as wide ...................................... 4

780

R UBIACEAE

4(3). 4. 5(4). 5. 6(2). 6. 7(6). 7. 8(6). 8. 9(8). 9. 10(9). 10. 11(10). 11. 12(9).

12. 13(12). 13. 14(13).

14.

15(8). 15.

Leaves 3–10 cm wide, the upper surface plane; capsules 15–25 mm long ................................................................................................ R. amazonica Leaves 10–35 cm wide, the upper surface plane to bullate; capsules 12– 48 mm long ............................................................................................. 5 Young growth and lower surface of leaves densely hirtellous to pilosulous; the upper surface of leaves bullate ...................................... R. pacimonica Young growth and lower surface of leaves strigose to glabrescent; the upper surface of leaves plane ................................................................R. ulei Inflorescence 1–3-flowered, unbranched or fasciculate; peduncle or stipe 0.1–0.8 cm long ...................................................................................... 7 Inflorescence several- to many-flowered, capitate to branched; peduncle 2.5–100 cm long ..................................................................................... 8 Stems, stipules, leaves, peduncules, and fruits hirsute; capsules 13–32 × 8–13 mm ...................................................................................... R. reducta Stems, stipules, leaves, peduncles, and fruits hirtellous to glabrescent; capsules 40–45 × 10 mm ........................................................... R. uniflora Leaves obtuse, rounded or truncate at apex ............................................. 9 Leaves acute to acuminate at apex, sometimes shortly or abruptly so .............................................................................................................. 15 Young growth, lower surface of leaves, and capsules glabrous or sparsely pubescent ............................................................................................. 10 Young growth, lower surface of leaves, and capsules moderately to densely pubescent ................................................................................ 12 Leaves 4.5–10 × 2.5–8 cm, the upper surface with secondary and tertiary venation sulcate, margins revolute ................................... R. pilosinervula Leaves 7.5–34 × 3.5–15 cm, the upper surface smooth, margins plane .............................................................................................................. 11 Leaves oblong to elliptic-oblong, 7.5–15 × 3.5–6 cm; corolla lobes ca. 8 mm long ..............................................................................................R. morilloi Leaves obovate-elliptic, obovate, or oblanceolate, 15–34 × 5.5–16 cm; corolla lobes 12–16 mm long .................................................... R. sipapoensis Leaves 5.5–17 × 2.5–7.5 cm, narrowly elliptic-oblong to oblanceolate or lanceolate-oblong; petioles 5–16 mm long; peduncles 4–12 cm long ................................................................................................. R. wurdackii Leaves 23–36 × 7.5–15 cm, elliptic to elliptic-oblong; petioles 30–45 mm long; peduncules 12–29 cm long .......................................................... 13 Inflorescences branched .............................................................. R. longifolia Inflorescences capitate, involucrate ........................................................ 14 Calyx limb ca. 15 mm long; corolla white, 35–37 mm long; stems, petioles, peduncles, and lower surface of leaves villosulous to villous or hirsute with spreading or lax trichomes 2–3 mm long ............ R. aracamuniensis Calyx limb 5.5–6 mm long; corolla pink, ca. 25 mm long; stems, petioles, peduncles, and lower surface of leaves closely tomentose, the trichomes not lax nor spreading .................................................... R. globosa Stipules, petioles, peduncles, and leaves hirsute; calyx limb spathaceous ..................................................................................................... R. hispida Stipules, petioles, peduncles, and leaves strigose, strigillose, pilosulous, or hirtellous to glabrescent or glabrous; calyx limb spathaceous to tubular, truncate to lobed ....................................................................... 16

Remijia 781

16(15). Calyx limb truncate, sinulate, or lobed to 1/4 its length, the lobes obtuse to rounded ................................................................................................ 17 16. Calyx limb spathaceous or lobed for 1/3 or more, the lobes obtuse to acute or acuminate ........................................................................................ 20 17(16). Leaf blades rounded to subcordate at base; petioles 1.5–4 mm long ............................................................................................. R. steyermarkii 17. Leaf blades acute to cuneate or obtuse at base; petioles 5–35 mm long .............................................................................................................. 18 18(17). Calyx limb 3–4 mm long; corolla white; leaves 4.5–10 cm wide, the lower surface densely hirtellous to glabrescent ................................ R. roraimae 18. Calyx limb 1–2 mm long; corolla pink; leaves 2–6 cm wide, the lower surface strigillose to glabrescent .............................................................. 19 19(18). Corolla with tube 21–24 mm long, lobes 10–13 mm long ........... R. maguirei 19. Corolla with tube 10–15 mm long, lobes 5–6 mm long ..... R. marahuacensis 20(16). Lower surface of leaves densely but minutely tomentulose in the areoles, with the veins glabrous and forming a distinctive tesselated pattern ................................................................................................... R. argentea 20. Lower surface of leaves glabrous to strigillose, strigose, or hirtellous, with the veins pubescent similarly to the lamina .............................. 21 21(20). Calyx limb 1.3–2 mm long; plants puberulous to glabrescent ................... ............................................................................................. R. pedunculata 21. Calyx limb 2–10 mm long; plants strigose or hirtellous to glabrescent .............................................................................................................. 22 22(21). Leaves chartaceous to subcoriaceous, 3–5.5 cm wide, with the secondary veins spreading; capsules 10–15 × 3–5 mm; corollas 7.5–12.5 mm long ................................................................................................. R. densiflora 22. Leaves subcoriaceous to coriaceous, 3.5–12 cm wide, with the secondary veins ascending; capsules 6–30 × 6–7 mm; corollas 16–20 mm long ..................................................................................................... R. firmula Remijia amazonica K. Schum. in Mart., Fl. Bras. 6(6): 153. 1889. Venezuela, Brazil; 2 varieties, 1 in Venezuela. R. amazonica var. amazonica Shrub to 3(–6) m tall, strigose or hirtellous to glabrescent; leaves 3- or 4-verticillate, 18–35 × 3–10 cm, adaxially plane or infrequently bullate; petioles 0–8 mm long; peduncles 6–30 cm long; calyx limb 1–2 mm long; corolla white, tube ca. 10 mm long, lobes ca. 5 mm long; capsules 15–25 × 6–9 mm. Evergreen lowland forests, 100–200 m; Amazonas (Río Ocamo and Río Padamo north of Sierra Unturán). Amazonian Brazil. Remijia aracamuniensis (Steyerm.) C.M. Taylor, comb. nov. —Cephalodendron aracamuniensis Steyerm., Ann. Missouri Bot. Gard. 76: 967. 1989. Tree to 4 m tall, villosulous to villous or

hirsute; leaves opposite, 32–35 × 16–26 cm, adaxially rugulose; petioles 25–40 mm long; peduncles 9–29 cm long; calyx limb ca. 15 mm long; corolla white, tube ca. 27 mm long, lobes ca. 8 mm long; capsules unknown. Wet forested ravines on tepui summits, 1500– 1600 m; Amazonas (Cerro Aracamuni). Endemic. Remijia argentea Steyerm., Mem. New York Bot. Gard. 23: 267, fig. 55. 1972. Shrub or tree to 6 m tall, glabrescent to glabrous; leaves opposite, 15–30 × 12–15 cm, adaxially with secondary veins sulcate; petioles 30–40 mm long; peduncles 9–14 cm long; calyx limb 5–7 mm long; corolla white, tube 17–18 mm long, lobes 14–16 mm long; capsules 20–35 × 7–9 mm. White-sand savannas, gallery forests, lowland to upland forests, 100–800 m; Amazonas (Cerro Moriche, Cerro Yapacana, basins of Río Guainía, Río Orinoco, and Río Ventuari). Endemic. ◆Fig. 593.

782

R UBIACEAE

Remijia delascioi Steyerm., Ann. Missouri Bot. Gard. 71: 332. 1984. Shrub to 1.5 m tall, generally unbranched, hirtellous to strigose or glabrescent; leaves 3verticillate, 11–21 × 1–2.5 cm, adaxially smooth; petioles 6–15 mm long; peduncles 6.5–17 cm long; calyx limb ca. 4 mm long; mature corolla and fruits unknown. Gallery forests bordering small streams overlying sandstone substrate, ca. 400 m; Amazonas (Cerro Vinilla). Endemic. Remijia densiflora Benth., J. Bot. (Hooker) 3: 215. 1841. Shrub or small tree to 8 m tall, puberulous to glabrescent; leaves opposite, 9–26 × 3–5.5 cm, adaxially smooth; petioles 5–15 mm long; peduncles 10–18 cm long; calyx limb 2–10 mm long; corolla white, tube 4.5–11 mm long, lobes 2.5–4 mm long; capsules 10–15 × 3–5 mm. Lowland to montane forests and disturbed areas, 100–1700 m. Venezuela, Guyana, adjacent Brazil; 2 subspecies, both in the flora area. This species varies notably in length of the calyx lobes, pubescence of all parts including the inner surface of the calyx tube, and size of the corollas. Key to the Subspecies of R. densiflora 1. Calyx lobes narrowly to broadly triangular or lanceolate, usually unequal, 0.5–5 mm long in anthesis, some of them elongating in fruit to as much as 8 mm long; bracts subtending flowers 1–6 mm long ................................ subsp. densiflora 1. Calyx lobes linear to narrowly triangular, equal, 2.5–4.5 mm long in anthesis, elongating in fruit to ca. 9 mm; bracts subtending flowers 4–19 mm long ................ .............................. subsp. stenopetala R. densiflora subsp. densiflora Venezuela, Guyana, adjacent Brazil; 2 varieties, both in the flora area. Key to the Varieties of R. densiflora subsp. densiflora 1. Corolla 12–12.5 mm long, with tube 8– 10 mm long, lobes 2.8–3.5 mm long ..................................... var. densiflora 1. Corolla 7.5–8 mm long, with tube 4.7– 5.2 mm long, lobes 2.5–3 mm long ........................................ var. minima

R. densiflora subsp. densiflora var. densiflora Remijia densiflora subsp. densiflora var. densiflora f. glabricalyx Steyerm., Mem. New York Bot. Gard. 23: 265. 1972. Bolívar (Cerro Guaiquinima, Río Caura), Amazonas (Cerro Cariche, Cerro Duida, Cerro Marahuaka, Sierra Parima). Guyana, adjacent Brazil. R. densiflora subsp. densiflora var. minima Steyerm., Mem. New York Bot. Gard. 23: 265. 1972. Bolívar (Cerro Ichún, Serranía Marutaní). Endemic. R. densiflora subsp. stenopetala (Standl. & Steyerm.) Steyerm., Mem. New York Bot. Gard. 23: 265. 1972. Bolívar (basins of Río Caroní, Río Caura, Río Cuyuní, and Río Paragua), Amazonas (Cerro Duida, Cerro Huachamacari). Endemic. ◆Fig. 594. Remijia firmula (Mart.) Wedd., Hist. Quinquin. 93. 1849. —Cinchona firmula Mart., Reise Bras. 3: 1286. 1831. —Caruto. Remijia spruceana Benth. ex K. Schum. in Mart., Fl. Bras. 6(6): 156. 1889. Tree to 7 m tall, densely hirtellous or pilosulous to glabrous; leaves opposite, 17–30 × 3.5–12 cm, adaxially smooth; petioles 15– 45 mm long; peduncles 7–30 cm long; calyx limb 2.5–5 mm long; corolla white, tube 6–11 mm long, lobes ca. 10 mm long; capsules 6–30 × 6–7 mm long. Savannas, seasonally flooded forests, evergreen lowland to montane forests, 100–1000 m; Amazonas (saddle between Cerro Duida and Cerro Marahuaka, Río Casiquiare, Río Negro basin, Río Orinoco). Amazonian Colombia, Peru, Brazil, Bolivia. Remijia firmula as circumscribed by Steyermark (1972) shows notable morphological variation, especially in pubescence and length of the calyx limb. Similar variation is found in some other species of Remijia, for example R. densiflora. Specimens from the southern part of the range of this species show less morphological variation, with leaves relatively small and consistently pubescent. Remijia globosa (Steyerm.) C.M. Taylor, comb. nov. —Cephalodendron globosum Steyerm., Mem. New York Bot. Gard. 23: 228, fig. 49. 1972.

Remijia 783

Tree to 8 m tall, tomentose; leaves opposite, 33–40 × 12–18 cm, adaxially smooth to rugulose; petioles 15–35 mm long; peduncles 21–29 cm long; calyx limb 5.5–6 mm long; corolla pink, tube ca. 18 mm long, lobes 4–6.5 mm long; capsules 9–17 × 3–7 mm. Wet forests along rocky streams in canyons, 1100– 1400 m; Amazonas (Sierra de la Neblina). Endemic. ◆Fig. 595. Remijia hispida Spruce ex K. Schum. in Mart., Fl. Bras. 6(6): 150. 1889. Shrub or tree to 8 m tall, hirsute; leaves opposite, 5.5–17 × 2.5–5.5 cm, adaxially smooth; petioles 5–20 mm long; peduncles (1–)5–18 cm long; calyx limb 7–10 mm long; corolla white, tube 15–27 mm long, lobes 10– 20 mm long; capsules 13–18 × 7–8 mm. White-sand savannas, sandstone ridges, along streams by igneous outcrops, open places overlying igneous substrate, 100–600 m; Amazonas (Río Autana, Río Pamoni, basins of Río Pasimoni, Río Orinoco, and Río Siapa). Adjacent Colombia. ◆Fig. 597. Remijia longifolia Benth. ex Standl., Publ. Field Columbian Mus., Bot. Ser. 8: 335. 1931. Small tree or shrub to 6 m tall, hirtellous; leaves opposite, 23–36 × 7.5–15 cm, adaxially smooth to rugulose; petioles 30–45 mm long; peduncles 12–15 cm long; calyx limb ca. 2.5 mm long; corolla color unknown, tube 10–14 mm long, lobes 5.5–9 mm long; capsules 15– 45 × 10–12 mm. White-sand savannas, gallery forests, lower tepui slope forests, 50–400 m; Amazonas (Río Atabapo, Río Sipapo, base of Sierra de la Neblina). Amazonian Brazil. Remijia maguirei Steyerm., Mem. New York Bot. Gard. 23: 256, fig. 53. 1972. Shrub or small tree to 4 m tall, strigose or sericeous to glabrescent; leaves opposite, 6.5–16 × 2–6 cm, adaxially smooth; petioles 5–15 mm long; peduncles 6–9.5 cm long; calyx limb 1–2 mm long; corolla pink, tube 21– 24 mm long, lobes 10–13 mm long; capsules 22–27 × 5 mm. Bonnetia scrub forests along streams on tepui slopes and summits, 1100– 2000 m; Amazonas (Sierra de la Neblina). Brazil (Amazonas: Serra da Neblina). Remijia marahuacensis Steyerm., Ann. Missouri Bot. Gard. 74: 111, fig. 11. 1987. Shrub to 2 m tall, strigillose or strigose to glabrescent; leaves opposite, 11–18 × 2.5–5

cm, adaxially the secondary veins sulcate; petioles 10–15 mm long; peduncles 6–15 cm long; calyx limb 1–1.3 mm long; corolla pink, tube 10–15 mm long, lobes 5–6 mm long; capsules 15–23 × 5–6 mm. Tepui slope forests, 1100–1200 m; Amazonas (Cerro Marahuaka). Colombia, Guyana. Remijia morilloi Steyerm., Fl. Venezuela 9(1): 115. 1974. —Bada (Bare), Palo de agujón. Shrub or small tree to 4 m tall, strigillose to glabrescent; leaves opposite, 7.5–15 × 3.5– 6 cm, adaxially smooth; petioles 7–25 mm long; peduncles 4–25 cm long; calyx limb 2.5– 3.5 mm long; corolla cream, tube ca. 9 mm long, lobes ca. 8 mm long; capsules 15–24 × 7–8 mm. White-sand savannas (caatinga formation), shrub and dwarf tree formation on white sand, 100–200 m; Amazonas (Río Negro basin, Río Sipapo). Amazonian Brazil. ◆Fig. 596. Remijia morilloi has been confused with R. involucrata K. Schum. of Colombia, which can be distinguished by its acute leaf apices, ferruginous-tomentose involucral bracts, and deeply divided calyx limb that is much longer than the hypanthium, versus rounded to truncate apices, glabrescent free bracts, and calyx limb that is lobed for only ca. 1/2 and about equal to the hypanthium in R. morilloi. Remijia pacimonica Standl., Publ. Field Columbian Mus., Bot. Ser. 7: 360. 1931. —Hoja de baba, Palo hinchazón. Shrub or tree to 15 m tall, generally unbranched, densely hirtellous or pilosulous at least on young growth and lower surface of leaves; leaves 3- or 4-verticillate, 48–93 × 18– 35 cm, adaxially bullate; petioles 0–10 mm long; peduncles 50–100 cm long; calyx limb 3–4 mm long; corolla white, tube ca. 23 mm long, lobes ca. 12 mm long; capsules 12–48 × 5–20 mm. Evergreen lowland forests, Río Negro caatinga, 400 m; Amazonas (basins of Río Guainía, Río Orinoco, and Río Pasimoni north to Río Sipapo). Amazonian Colombia, Ecuador, Peru, Brazil. ◆Fig. 598. The sap of Remijia pacimonica is said to be used to reduce inflammation of the skin. This species is more common in the western Amazon basin than in the flora area. Remijia chelamophylla G. Sullivan of Peru is similar to R. pacimonica and can be distinguished by its sparser, appressed rather than spreading

784

R UBIACEAE

pubescence on the lower surface of the leaves. Remijia pedunculata (H. Karst.) Flueck., Chinarind. 17, t. 6. 1883. —Cinchona pedunculata H. Karst., Fl. Colomb. 1: 53, t. 36. 1859. Remijia berryi Steyerm., Ann. Missouri Bot. Gard. 71: 332. 1984. Tree to 15 m tall, puberulous to glabrescent; leaves opposite, 8–25 × 2.5–13 cm, adaxially smooth; petioles 7–23 mm long; peduncles 8–16 cm long; calyx limb 1.3–2 mm long; corolla white, tube 5.5–12 mm long, lobes 3.5–8.5 mm long; capsules 7–19 × 4–7 mm. Evergreen lowland forests, 100–200 m; Amazonas (Santa Bárbara). Colombia, Ecuador, Peru. ◆Fig. 599. Remijia pilosinervula Steyerm., Mem. New York Bot. Gard. 17(1): 232, fig. 30. 1967. Small tree or shrub to 4 m tall, strigose to glabrescent; leaves opposite, 4.5–10 × 2.5–8 cm, adaxially rugulose; petioles 3–10 mm long; peduncles 3–10 cm long; calyx limb 2–4 mm long, lobed; corolla pink, tube 5–12 mm long, lobes 4–6 mm long; capsules 8–10 × 5–6 mm. Tepui scrub forests, 1900–2200 m; Bolívar (Macizo del Chimantá). Guyana. ◆Fig. 603. Remijia reducta Steyerm., Ann. Missouri Bot. Gard. 76: 968, fig 16. 1989. Shrub to 1 m tall, hirsute; leaves apparently opposite, 4–10 × 2–3 cm, apparently smooth adaxially; petioles 2–9 mm long; peduncles 1–7 mm long; calyx limb 1–2 mm long; corolla white, tube ca. 11 mm long, lobes 10–12 mm long; capsules 13–32 × 8–13 mm. Tepui meadows, 1400–1600 m; Amazonas (Cerro Aracamuni). Endemic. Remijia roraimae (Benth.) K. Schum. in Mart., Fl. Bras. 6(6): 154. 1889. —Cinchona roraimae Benth., J. Bot. (Hooker) 3: 214. 1841. —Ladenbergia roraimae (Benth.) Klotzsch in Hayne, Gestr. Darstell. Gew. 14: sub. pl. 15. 1846. —Cascarilla roraimae (Benth.) Wedd., Hist. Quinquin. 89. 1849. Shrub or small tree to 5 m tall, hirtellous or pilosulous to glabrescent; leaves opposite, 9.5–25 × 4.5–10 cm, adaxially smooth; petioles 5–25 mm long; peduncles 9–18 cm long;

calyx limb 3–4 mm long; corolla white, tube ca. 21 mm long, lobes 9–14 mm long; capsules 14–17 × 10 mm. Savannas, tepui slope and summit forests, 900–1900 m. Guyana, Brazil (Roraima); 4 varieties, all in the flora area. Remijia roraimae is quite variable in the degree and type of pubescence of the leaves and the styles, and in pedicel length. Steyermark has recognized the following infraspecific taxa, which may deserve reconsideration. Key to the Varieties of R. roraimae 1. Lower surface of leaf appressed-puberulous to glabrescent; leaves 17–28 cm long ................................ var. adpressa 1. Lower surface of leaf tomentellous or hirtellous to glabrescent; leaves 9.5–25 cm long .................................................... 2 2. Style appressed-pubescent throughout or at least in upper or basal half ................. ...................................... var. roraimae 2. Style glabrous throughout .................. 3 3. Leaves with secondary veins 13 or 14 pairs; flowers with pedicels 2–4 mm long ........................... var. auyantepuiensis 3. Leaves with secondary veins 10–25 pairs; flowers sessile or subsessile .................... .................................... var. guianensis R. roraimae var. adpressa Steyerm., Mem. New York Bot. Gard. 23: 262. 1972. Bolívar (Río Ichún, Río Paragua, Sierra Ichún), Amazonas (Cerro Parú). Endemic. R.

roraimae var. auyantepuiensis Steyerm., Mem. New York Bot. Gard. 23: 263. 1972. Bolívar (Auyán-tepui). Endemic.

R. roraimae var. guianensis Steyerm., Mem. New York Bot. Gard. 23: 263. 1972. Bolívar (Gran Sabana east of San Ignacio). Guyana. ◆Fig. 602. R. roraimae var. roraimae. —Dorona (Arekuna). Bolívar (Gran Sabana, Macizo del Chimantá, Perai-tepui, Roraima-tepui), Amazonas (Sierra Parima). Guyana, adjacent Brazil. Pemón Amerindians of the Gran Sabana use the bark, after boiling it in water, for colds and headaches.

Remijia 785

Fig. 592. Remijia steyermarkii

Remijia sessilis Steyerm., Ann. Missouri Bot. Gard. 75: 1084. 1988. Shrub to 2 m tall, hirsute; leaves apparently opposite, ca. 36 × 16 cm, adaxially apparently smooth; petioles 2.5–3 cm long, inflated; stipules unknown; inflorescences sessile, glomerate, 2 × 1.5–2 cm; calyx limb 4.5–6 mm long; corolla 30–38 mm long, color unknown; capsules 32–38 × 5–11 mm. Lower tepui slope forests, 400–500 m; Amazonas (Sierra de la Neblina). Endemic. Remijia sipapoensis Steyerm., Mem. New York Bot. Gard. 23: 255, fig. 52. 1972. Tree to 10 m tall, strigose to glabrescent; leaves opposite, 15–34 × 5.5–16 cm, adaxially smooth; petioles 20–30 mm long; peduncles 7.5–18 cm long; calyx limb 3.5–5 mm long; corolla color unknown, tube 9–10 mm long, lobes 12–16 mm long; capsules unknown. Tepui forests, ca. 1500 m; Amazonas (Cerro Sipapo). Endemic. ◆Fig. 600. Remijia steyermarkii Standl., Fieldiana, Bot. 28: 612, fig. 135. 1953. Tree to 8 m tall, strigose, sericeous, or hirtellous to glabrescent; leaves opposite, 6.5– 18.5 × 3–9.5 cm, adaxially plane or the secondary veins sulcate; petioles 1.5–4 mm long; peduncles 6–17 cm long; calyx limb 1.5–2 mm long; corolla pale grayish lavender, tube ca. 18 mm long, lobes ca. 6 mm long; capsules 20 × 6–8 mm. Tepui forests, 1100–1500 m; Amazonas (Cerro Duida, Serranía Parú). Endemic. ◆Fig. 592. Remijia ulei K. Krause, Notizbl. 6: 201. 1914. —Carutillo. Shrub or tree to 12 m tall, strigillose to glabrescent; leaves 3- or 4-verticillate, 38–76 × 10–23 cm, adaxially plane to a little bul-

late; petioles 5–20 mm long; peduncles 10–35 cm long; calyx limb 2.5–4 mm long; corolla white, tube 18–19 mm long, lobes 12–14 mm long; capsules 32–43 × 10–12 mm. Evergreen lowland forests, 100–200 m; Amazonas (middle Río Baría, Yavita). Amazonian Brazil. Remijia uniflora C.M. Taylor, Novon 6: 218, fig. 3. 1996. Shrub to 3 m tall, hirtellous to glabrescent; leaves opposite, 13.5–17.5 × 4.7–5.5 cm, rugulose adaxially; petioles 10–13 mm long; peduncles 0.5–0.7 cm long; calyx and corolla unknown; capsules 40–45 × 10 mm. Shrub forests on granitic substrates, 600–700 m; Amazonas (Cuao-Sipapo massif). Endemic.

786

R UBIACEAE

Fig. 593. Remijia argentea Fig. 594. Remijia densiflora subsp. stenopetala

Remijia 787

Fig. 595. Remijia globosa

788

R UBIACEAE

Fig. 596. Remijia morilloi

Fig. 597. Remijia hispida

Remijia 789

Fig. 598. Remijia pacimonica

790

R UBIACEAE

Fig. 599. Remijia pedunculata

Fig. 600. Remijia sipapoensis

Remijia 791

Fig. 601. Remijia wurdackii

Fig. 602. Remijia roraimae var. guianensis

792

R UBIACEAE

Fig. 603. Remijia pilosinervula

Remijia wurdackii Steyerm., Mem. New York Bot. Gard. 23: 259, fig. 54. 1972. Slender tree or shrub to 5 m tall, strigose to glabrescent; leaves opposite, 5.5–17 × 2.5– 7.5 cm, adaxially smooth; petioles 5–16 mm long; peduncles 4–12 cm long; calyx limb 4.5–

8 mm long; corolla white, tube 15–16 mm long, lobes ca. 14 mm long; capsules 15–60 × 5–8 mm. Gallery forests, Río Negro caatinga, 100–200 m; Amazonas (Río Atabapo, Río Autana, Río Guayapo, Río Sipapo). Amazonian Brazil. ◆Fig. 601.

70. RETINIPHYLLUM Bonpl. in Humb. & Bonpl., Pl. Aequin. 1: 86, pl. 25. 1806 [1808]. Commianthus Benth., J. Bot. (Hooker) 3: 223. 1841. —Retiniphyllum sect. Commianthus (Benth.) in Mart., Fl. Bras. 6(5): 6. 1881. Ammianthus Spruce ex Hook. f., Gen. Pl. 2: 98. 1873, nom. illeg., stat. confusus. —Retiniphyllum sect. Ammianthus Spruce ex Müll. Arg. in Mart., Fl. Bras. 6(5): 6. 1881. by Rocio Cortés and Julian A. Steyermark Shrubs or small trees; buds, young branches, and inflorescences commonly resinous. Leaves opposite, petiolate; stipules persistent or caducous, generally valvate to imbricate in bud, fused around the stem and truncate (i.e., cup-shaped) or triangular, less often interpetiolar and triangular, or sometimes splitting along one side with margin entire or laciniate, dimorphic in two species (i.e., two different types of stipules in the same species). Inflorescences terminal or rarely axillary, multiflowered, racemose, spicate, or umbellate, sessile or pedunculate, with or without bracts subtending the most basal pair of flowers, with a bracteole and an involucel (i.e., calyculus; a pair of floral bracts) subtending each flower, the bracteoles sometimes reduced or foliose (i.e., with an expanded lamina), the

Retiniphyllum 793

involcels cupular (i.e., calyx-like) or discoid. Flowers sessile or pedicellate, homostylous, protandrous. Hypanthium turbinate, ellipsoid, or globose. Calyx limb cupular or tubular, truncate, dentate, or lobed, the lobes or teeth 5, without calycophylls; corolla salverform and sometimes zygomorphic with either tube or lobes bent in bud, dark pink, red, white, or greenish white tinged with light pink, externally sericeous or rarely glabrous, internally the lobes sericeous and the tube with a ring of trichomes near the base or at the mouth, lobes 5, convolute to the left in bud, reflexed at anthesis. Stamens 5, inserted near top of corolla tube; filaments pubescent or rarely glabrous; anthers dorsifixed near the base, narrowly oblong, exserted, with basal and apical sterile appendages. Style exserted, terete; style branches 5, recurved at maturity. Ovary (4)5(6)-locular; ovules 2 per locule, collateral, descending on axile placentas, inserted at top or middle of the septum. Fruits drupaceous, globose to ellipsoid, fleshy, red at maturity turning dark purple later; pyrenes (4)5(6), 1-locular, reniform, woody, with or without wings. Seeds 1 per locule due to abortion or rarely 2, reniform or cylindrical. South America; 22 species, 11 in Venezuela, all in the flora area. This genus was treated by Steyermark (1965) and more recently has been revised by Cortés (2003). Most Retiniphyllum species are endemic to the Guayana Shield, where they are always associated with white-sand soils. More than half of the species grow in Río Negro caatinga forests, a formation widespread along the Río Negro basin and dominant in the states of Guainía and Vaupés in Colombia, Amazonas in Venezuela, and northern Amazonas in Brazil. According to recent phylogenetic studies (Cortés 2003), the tribe Retiniphylleae and its single genus are monophyletic. The tribe Retiniphylleae is included in the subfamily Ixoroideae, and is sister to two clades: one including members of the tribes Ixoreae and Vanguerieae, and another that includes members of the tribes Coffeeae, Gardenieae, Pavetteae, and Octotropideae. The resinous branches and inflorescences, the bracteole and involucel subtending each flower, and the anthers with basal and apical appendages are distinctive. Key to the Species of Retiniphyllum 1. 1. 2(1). 2. 3(2). 3. 4(3).

4.

5(1). 5. 6(5).

Inflorescences spicate; flowers sessile ....................................................... 2 Inflorescences racemose or umbellate; flowers pedicellate ...................... 5 Corolla red or dark pink, the tube > 10 mm long .......................... R. discolor Corolla white or greenish white and tinged with light pink, the tube < 10 mm long ............................................................................................ 3 Leaves, stipules, and rachis glabrous ....................................... R. truncatum Leaves, stipules, and rachis pubescent ..................................................... 4 Stipules oblong or ovate, 15–30 mm long, papyraceous; bracteoles foliose, > 4 mm long; involucel lobulate, papyraceous; pyrenes with 2 lateral wings, dorsally smooth .............................................................. R. pilosum Stipules truncate, to 3 mm long, coriaceous; bracteoles reduced, < 2 mm; involucel truncate, coriaceous; pyrenes without wings, dorsally verrucose .................................................................................... R. schomburgkii Inflorescences umbellate; stipules dimorphic, both cup-shaped and triangular splitting at one side on the same individual plant ..................... 6 Inflorescences racemose; plants with only one type of stipule ................ 7 Involucel, pedicel, and calyx hirsute; pyrenes with 2 lateral wings; seeds cylindrical, inserted at top of the septum ................................ R. scabrum

794

R UBIACEAE

6.

Involucel, pedicel, and calyx glabrous; pyrenes without wings; seeds reniform, inserted at the middle of the septum ....................... R. pauciflorum 7(5). Corolla red or dark pink, actinomorphic ................................................... 8 7. Corolla white or greenish white and tinged with light pink, zygomorphic with either the tube or the lobes bent in bud ....................................... 9 8(7). Inflorescences terminal, pendulous; bracteoles reduced, < 2 mm long ................................................................................................ R. laxiflorum 8. Inflorescences axillary, erect; bracteoles foliose, > 2 mm long ................... .......................................................................................... R. secundiflorum 9(7). Calyx tubular, > 2 mm long; corolla tube < 10 mm long; pyrene surface verrucose .................................................................................. R. tepuiense 9. Calyx cupular, < 2 mm long; corolla tube > 10 mm long; pyrene surface smooth .................................................................................................. 10 10(9). Involucel lobulate; pyrenes with 2 lateral wings ................. R. chloranthum 10. Involucel truncate; pyrenes without wings .................................. R. concolor Retiniphyllum chloranthum Ducke, Trop. Woods 76: 31. 1943. —Palo de cogollo de piedra, Palo de gallineta, Peramán de vieja. Shrub or small tree to 10 m tall; leaves 13–30 × 6–14.5 cm; inflorescences racemose, 6–19 cm long, with 2 bracts subtending the basal pair of flowers; bracteoles reduced; involucel discoid, lobulate; calyx cupular, 0.8–1 × 1.8–2.9 mm; corolla 16–27 mm long, the tube white with pale pink lines, (8.5–)10– 19 mm long, the lobes 5.5–8.5 × 1.5–2 mm, light pink on the left half and white on the right half; fruit globose, 10-ribbed, 5–9.2 × 4.2–7 mm; pyrenes 5.2–5.5 × 1.8–3 mm, each with 2 wings. Río Negro caatinga forests and shrublands, gallery forests, basimontane and submontane forests, 100–1100(–2000) m; Bolívar (Amaruay-tepui, Cerro Apacará, Cerro Guaiquinima, Gran Sabana, Macizo del Chimantá, Río Auyán, Río Caroní, Río Caura, Río Churún, Río Tírica), Amazonas (Cerro Duida, La Esmeralda, Río Casiquiare, Río Guainía, Río Negro, Río Orinoco, Río Yatúa, San Juan de Ucata, Sierra de la Neblina, Yavita). Colombia, Guyana, Brazil. ◆Fig. 606. Retiniphyllum chloranthum is sometimes employed by local inhabitants as a vermifuge. Retiniphyllum concolor (Spruce ex Benth.) Müll. Arg. in Mart., Fl. Bras. 6(5): 8. 1881, emend. Cortés, herein. —Commianthus concolor Spruce ex Benth., Hooker’s J. Bot. Kew Gard. Misc. 5: 235. 1853. —Ají de Morrocoy, Palo ojo de picure.

Retiniphyllum martianum Müll. Arg. in Mart., Fl. Bras. 6(5): 9. 1881. Retiniphyllum angustiflorum K. Krause, Bot. Jahrb. Syst. 40: 326. 1908. Shrub or small tree to 6 m tall; leaves 8.5– 37.5 × 3–12.5 cm; inflorescences racemose, (2.5–)4–10(–18) cm long, with 2 bracts subtending the basal pair of flowers; bracteoles reduced; involucel discoid, truncate; calyx cupular, 0.8–2 × 2–2.6 mm; corolla 15.8–39 mm long, the tube 10–24 mm long, white with pale pink longitudinal lines, the lobes 4–10.5 × 1.6–2.5 mm, pale pink on the left half and white on the right half; fruit globose, 5-ribbed, 7–9 × 4.5–5 mm; pyrenes 4.8–6.2 × 1.8–2 mm, without wings. Río Negro caatinga forests and shrublands, flooded forests on white sands, submontane forests, 100–1100 m; Bolívar (Río Paragua), Amazonas (Río Atabapo, Río Autana, Río Baría, Río Casiquiare, Río Guainía, Río Negro, Río Pasimoni, Río Puruname, Río Sipapo, Yavita). Colombia, Peru, Brazil. ◆Fig. 607. When Mueller Argoviensis transferred Commianthus concolor to Retiniphyllum, he emended Bentham’s description unfortunately using a mixed specimen. That collection included R. concolor, characterized by the truncate involucel and 5-ribbed fruits, along with an undescribed species with a lobulate involucel and 10-ribbed fruits. Mueller Argoviensis apparently did not notice this problem, and described the fruits of R. concolor as 5-ribbed in his full description but as 10-ribbed in his diagnostic description. He described the new species R.

Retiniphyllum 795

martianum in the same publication, pointing out its similarities with R. concolor. He differentiated them by the 10-ribbed fruits and minutely denticulate involucel of R. concolor, versus the 5-ribbed fruits and truncate involucel of R. martianum. As a consequence, the name R. concolor has been erroneously used for the species with 10-ribbed fruits. However Bentham’s original name applies to the species with the 5-ribbed fruits, so Retiniphyllum martianum is a synonym of R. concolor. Thus, the circumscription of R. concolor is here emended to exclude the plants with 10-ribbed fruits. This other species with 10-ribbed fruits was later described as R. chloranthum Ducke. Retiniphyllum discolor (Spruce ex Benth.) Müll. Arg. in Mart., Fl. Bras. 6(5): 11. 1881. —Commianthus discolor Spruce ex Benth., Hooker’s J. Bot. Kew Gard. Misc. 5: 234. 1853. Shrub or small tree to 3 m tall; leaves 4.6– 9.6 × 2.2–4.1 cm; inflorescences spicate, 6.1– 14.8 cm long, without bracts subtending the basal pair of flowers; bracteoles reduced; involucel discoid; calyx tubular, 8.1–8.5 × 3.2–3.8 mm; corolla red, 25.5–27.8 mm long, the tube 14.5–15.8 mm long, the lobes 10.3– 12 × 1.4–2.2 mm; fruit ellipsoid, 5.5–7 × 4.2– 7.1 mm; pyrenes 5.9–6 × 2.5 mm, without wings. Río Negro caatinga forests and shrublands, savannas, 50–200 m; Amazonas (northeast of Caño Marueta, Río Ventuari). Brazil. Retiniphyllum laxiflorum (Benth.) N.E. Br., Trans. Linn. Soc. London, Bot. 6: 36. 1901. —Patima laxiflora Benth., J. Bot. (Hooker) 3: 220. 1841. Savannas, open rocky places, shrublands, submontane and montane forests, gallery forests, 100–1600 m. Guyana, Suriname, Brazil; 2 varieties, 1 in Venezuela. The second variety, var. brasiliense Steyerm., is endemic to the Atlantic forest in the state of Bahia, Brazil. Its flowers and fruits are much smaller than in the typical variety. R. laxiflorum var. laxiflorum. —Won-tayek (Pemón). Synisoon schomburgkianum Baill., Bull. Mens. Soc. Linn. Paris 208. 1879.

Retiniphyllum laxiflorum var. longilobum Steyerm., Mem. New York Bot. Gard. 12: 233. 1965. Shrub or small tree to 4 m tall; leaves 4.1– 11.3 × 1.5–5.5 cm; inflorescences racemose, (0.85–)1.5–7(–8.7) cm long, without bracts subtending the basal pair of flowers; bracteoles reduced; involucel discoid; calyx tubular, (3.2–)3.9–5.5 × 2.1–3.7 mm; corolla red, (23–)27–44 mm long, the tube 13–22 mm long, the lobes 8–20 × 1.8–3.7 mm; fruit globose, (6–)7–9.7 × 5.5–8.5 mm; pyrenes (3.8–) 5–6.2 × 2.5–2.8 mm, with 2 perpendicular wings. Savannas, open rocky places, shrublands, submontane and montane forests, gallery forests, 100–1600 m; Bolívar (Auyán-tepui, El Paují, Gran Sabana, Ilútepui, Karaurín-tepui, Macizo del Chimantá, Ptari-tepui, Túriba, Uaipán-tepui), Amazonas (Cerro Aratitiyope, Cerro Duida, Cerro Guanay, Cerro Huachamacari, Cerro Parú, Cerro Sipapo, Cerro Yapacana, Cerro Yaví, Cerro Yutajé, Río Casiquiare, Río Cuao, Río Siapa, Río Ventuari, Sierra de la Neblina). Guyana, Suriname, Brazil. ◆Fig. 611. Retiniphyllum pauciflorum Kunth ex K. Krause, Bot. Jahrb. Syst. 40: 326. 1908. Shrub 1–4 m tall; leaves 4.3–8.7 × 1.5–3.4 cm; inflorescences umbellate, 0.8–2.5 cm long, with or without 2 bracts subtending the basal pair of flowers; bracteoles reduced; involucel discoid; calyx tubular, 4.5–7.5 × 3– 3.8 mm; corolla red, 23–36 mm long, the tube 15–18 mm long, the lobes 7–12 × 1.8–3.5 mm; fruit ellipsoid, 6.5 × 4.8–5; pyrenes 5–5.8 × 1.6–2.5, without wings. Periodically flooded savannas, Río Negro caatinga forests, 50– 200 m; Amazonas (Río Atabapo, Río Casiquiare, Río Guainía, Río Guasacavi, Río Sipapo, Río Temi, San Fernando de Atabapo). Endemic. Retiniphyllum pilosum (Spruce ex Benth.) Müll. Arg. in Mart., Fl. Bras. 6(5): 7. 1881. —Commianthus pilosus Spruce ex Benth., Hooker’s J. Bot. Kew Gard. Misc. 5: 233. 1853. —Borrajón, Palo de peluza. Shrub or small tree 0.5–4 m tall; leaves 7.6–18 × 3.5–7 cm; inflorescences spicate, 5.5–12.5 cm long, with or without 2 bracts subtending the basal pair of flowers; bracteoles foliose; involucel cupular; calyx tubular, 2.8–4.2 × 2.1–3.5 mm; corolla 13.3–

796

R UBIACEAE

15.1 mm long, the tube 6.8–8.3 mm long, white or cream white with pale pink lines, the lobes 6–9 × 1.3–2.7 mm, white or cream white; fruit oblate or rarely globose, 5–6 × 4.5–7.8 mm; pyrenes 3.9–5.2 × 2.1–2.8 mm, with 2 parallel wings. Río Negro caatinga forests and shrublands, savannas, 50–200 m; Amazonas (Maroa, Río Negro, Río Pasimoni, Río Temi). Colombia, Brazil. ◆Fig. 608. Retiniphyllum scabrum Benth., J. Bot. (Hooker) 3: 222. 1841. Retiniphyllum erythranthum Standl., Publ. Field Columbian Mus., Bot. Ser. 7: 399. 1931. —Retiniphyllum scabrum var. erythranthum (Standl.) Steyerm., Mem. New York Bot. Gard. 12: 236. 1965. Retiniphyllum cearense Standl., Field Mus. Nat. Hist., Bot. Ser. 17: 220. 1937. Retiniphyllum scabrum var. ayangannense Steyerm., Mem. New York Bot. Gard. 12: 236. 1965. Shrub to 3 m tall; leaves dimorphic, blades in fertile shoots 1.7–7.4 × 1.2–4.2 cm, blades in sterile shoots 5.3–12.4 × 1.9–3.8 cm; inflorescences umbellate, 4-flowered, without bracts subtending the basal pair of flowers; bracteoles discoid; involucel discoid; calyx tubular, 3–6.5(–8) × 2.8–4.2 mm; corolla red, 30–38 mm long, the tube 22–27 mm long, the lobes 6.8–10.5 × 1.7–4.9 mm; fruit ellipsoid or globose, 6–11 × 6–9 mm, glabrous; pyrenes 4.5–8 × 2.5–3.5 mm, with 2 perpendicular wings. Savannas, open rocky places, shrublands, submontane and montane forests, gallery forests, 500–2300 m; Bolívar (Aprada-tepui, Auyán-tepui, Cerro Jaua, Cerro Sarisariñama, Cerro Venamo, Ilú-tepui, Kamarkawarai-tepui, Kukenántepui, Macizo del Chimantá [Abacapá-tepui, Acopán-tepui, Toronó-tepui], Murisipán-tepui, Ptari-tepui, Río Aponguao, Río Cuyuní, Serranía Marutaní, Sororopán-tepui, Terekéyurén-tepui, Uaipán-tepui), Amazonas (Cerro Asisa, Cerro Duida, Cerro Marahuaka, Cerro Parú, Río Ventuari, Río Yudi, Serranía Uasadi, Sierra de Maigualida, Sierra Parima). Guyana (Pakaraima Mountains), Brazil (Amazonas: Serra Aracá). ◆Fig. 612. Retiniphyllum schomburgkii (Benth.) Müll. Arg. in Mart., Fl. Bras. 6(5): 12. 1881. —Commianthus schomburgkii Benth., J. Bot. (Hooker) 3: 223. 1841. —Jovillo, Kitak-Yek.

Retiniphyllum schomburgkii var. angustiflorum Huber, Bull. Soc. Bot. Genève Ser. 2, 6: 209. 1914. Retiniphyllum schomburgkii subsp. occidentale Steyerm., Mem. New York Bot. Gard. 12: 240. 1965. Retiniphyllum schomburgkii subsp. occidentale var. hirticalyx Steyerm., Mem. New York Bot. Gard. 12: 241. 1965. Shrub or small tree to 5 m tall; leaves 3.3– 12.1 × (1.6–)2.2–5.3 cm; inflorescences spicate, 6–18.5 cm long, with or without 2 bracts subtending the basal pair of flowers; bracteoles reduced; involucel discoid; calyx tubular, 1.8–5.2 × 2–3.1 mm; corolla 12.1–23 mm long, the tube 3.8–10 mm long, white or greenish white with 5 longitudinal light pink lines, the lobes 6.5–14 × 1.2–2.8 mm, white; fruit globose, ellipsoid, or oblate, 3.5–5.5 × 4– 7 mm; pyrenes 3.1–5 × 1.5–2.5 mm, without wings. Savannas, open rocky places, shrublands, submontane and montane forests, gallery forests, 300–2300 m; Bolívar (Auyántepui, Cerro Guaiquinima, Cerro Pitón, Gran Sabana, Isla Ratón, Río Asa, Río Caroní, Río Chicanán, Río Cuyuní, Río Ichún, Río Tonoro, Sierra de Lema), Amazonas (widespread). Colombia, Guyana, Suriname, Brazil. ◆Fig. 605. Retiniphyllum secundiflorum Bonpl. in Humb. & Bonpl., Pl. Aequin. 1: 86, pl. 25. 1806 [1808]. —Nonatelia secundiflora (Bonpl.) Spreng., Syst. Veg. 1: 751. 1825. —Peramancito. Shrub or small tree 1–5 m tall; leaves 3.6– 11.5 × 2–5.6 cm; inflorescences racemose, 5– 16 cm long, with 2 bracts subtending the basal pair of flowers; bracteoles foliose; involucel cupular; calyx tubular, 3–7.5 × 2.5– 3.3 mm; corolla red, 17–34 mm long, the tube 11–17.5 mm long, the lobes 5.5–10 × 1–1.5 mm; fruit oblate, 3–6.7 × 3.8–6.7 mm; pyrenes 4 × 2.1–2.7 mm, with 2 very reduced ribs. Río Negro caatinga forests and shrublands, savannas, periodically flooded savannas, 50–200 m; Amazonas (Caño Cumare, Caño Pimichín, Caño San Miguel, La Ceiba, La Esmeralda, Río Casiquiare, Río Guainía, Río Guasacavi, Río Pasiba, Río Pasimoni, Río Siapa, Río Temi, San Fernando de Atabapo, Santa Cruz). Colombia (Guainía-Venezuelan border), Brazil (Amazonas: Rio Uaupés). ◆Fig. 610.

Retiniphyllum 797

Fig. 604. Retiniphyllum tepuiense

Fig. 605. Retiniphyllum schomburgkii

Fig. 606. Retiniphyllum chloranthum

798

R UBIACEAE

Fig. 607. Retiniphyllum concolor

Fig. 608. Retiniphyllum pilosum

Retiniphyllum 799

Fig. 609. Retiniphyllum truncatum

Fig. 611. Retiniphyllum laxiflorum var. laxiflorum

Fig. 612. Retiniphyllum scabrum

Fig. 610. Retiniphyllum secundiflorum

800

R UBIACEAE

Retiniphyllum tepuiense Steyerm., Mem. New York Bot. Gard. 12: 237. 1965. Shrub to 4 m or small tree to 8 m tall; leaves 3–10.5 × 1.4–4 cm; inflorescences racemose, 4.5–11.6 cm long, with or without 2 bracts subtending the basal pair of flowers; bracteoles reduced; involucel discoid; corolla cream-white tinged with pink, 12.9–17.7 mm long, the tube 4.3–7.2 mm long, the lobes 8– 10.5 × 1.3–2.8 mm; fruit globose, 6–7 × 6–7.5 mm; pyrenes 4.7–5.5 × 2.3–3 mm, with 2 parallel wings. Montane forests, shrublands, savannas, open rocky places, 1300–1900 m; Amazonas (Caño Piedra, Cerro Cuao, Cerro Sipapo, Cerro Yutajé, Sierra de la Neblina). Endemic. ◆Fig. 604. Retiniphyllum truncatum Müll. Arg. in Mart., Fl. Bras. 6(5): 11. 1881. —Cupi banero, Cupi hoja fina. Retiniphyllum pallidum Müll. Arg. in

Mart., Fl. Bras. 6(5): 12. 1881. Retiniphyllum truncatum var. angustifolium Steyerm., Mem. New York Bot. Gard. 12: 242. 1965. Shrub or small tree to 4 m tall; leaves 4.5– 12.7 × 2.3–6.7 cm; inflorescences spicate, 7.2–19.3 cm long, with or without 2 bracts subtending the basal pair of flowers; bracteoles reduced; involucel discoid; corolla 13–17.5 mm long, the tube 5.5–7.5 mm long, white or greenish white with 5 longitudinal light pink lines, the lobes 6–11 × 1.2–2.8 mm, white or white tinged with light pink; fruit oblate, 2.6–4.5 × 3.5–6.5 mm; pyrenes 3.2– 3.5 × 1.5–1.9 mm, without wings. Río Negro caatinga forests and shrublands, montane forests, savannas, open rocky places, 50– 1600 m; Amazonas (Cerro Aracamuni, Cerro Duida, Río Casiquiare, Río Atabapo, Río Guainía, Río Negro, Río Orinoco, Río Temi). Colombia, Brazil. ◆Fig. 609.

71. RICHARDIA L., Sp. Pl. 330. 1753. by Charlotte M. Taylor and Julian A. Steyermark Annual or perennial herbs, terrestrial, unarmed, erect to prostrate, usually conspicuously hispid or hirsute. Leaves opposite, generally subsessile, venation not lineolate; stipules interpetiolar and united to leaf bases, persistent, in bud generally erect and appressed or valvate, the sheath truncate to rounded, setose. Inflorescence capitate, terminal, pedunculate, few- to multiflowered, enclosed by 2 or 4 involucral ovate leaves or foliaceous bracts. Flowers numerous, small, homostylous, protandrous. Hypanthium turbinate to subglobose. Calyx limb deeply lobed, lobes 4–8, without calycophylls; corolla white to lavender or pink, funnelform, internally glabrous or pubescent in the throat and/or on lobes, lobes 4–6, valvate in bud. Stamens (3)4 or 6, inserted in corolla throat; anthers narrowly oblong to narrowly elliptic, exserted, dorsifixed near middle. Ovary (2)3- or 4(6)-locular; ovules 1 in each locule, axile. Fruit dry, schizocarpous; mericarps 3 or 4, indehiscent, dorsally usually muricate or papillose. Seeds elliptic-oblong, small. Southern U.S.A., Mexico, Central America, Antilles, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina, Uruguay, adventive in Africa, Asia, and some Pacific islands; 15 species, 1 in Venezuela. Richardia is similar to other members of its tribe and often confused with them, especially Borreria and Mitracarpus. These genera are all similar in their herbaceous habit and setose stipules. However the six calyx and corolla lobes in Richardia and its fruits with three mericarps distinguish it. Richardia scabra L., Sp. Pl. 330. 1753. —Richardsonia scabra (L.) A. St.-Hil., Pl. Usual. Bras. 1: 8, t. 8. 1824. —Conejilla. Annual herb; leaves (and involucral

leaves of inflorescences) 1–7 × 0.5–2 cm; stipule sheaths 1–4 mm long, setae 3–15, 1–5 mm long; calyx lobes 6, 1.5–3 mm long; corolla tube 3–8 mm long, lobes 6, 1–3 mm long; fruits 2–3.5 mm long, mericarps 3. Sa-

Ronabea 801

vannas, exposed grassy slopes, borders of Mauritia palm swamps, 100–400 m; Delta Amacuro (between Los Castillos de Guayana and Piacoa), Bolívar (Altiplanicie de Nuria, Ciudad Piar, margin of Guri Dam, Hato La Vergareña). Scattered at low altitudes thoughout Venezuela; widespread and weedy throughout the Americas and adventive in the Paleotropics. ◆Fig. 613. Richardia scabra is sometimes not recognized as Rubiaceae because of its six calyx and corolla lobes and its 3-merous fruits. It is the most commonly encountered species in the genus. Another weedy species, R. brasiliensis Gomes, is apparently spreading in the Neotropics and may be expected in the flora area. Richardia brasiliensis can be distinguished reliably from R. scabra only by its mature fruits: each mericarp has 1 narrow sulcus on the inner (i.e., adaxial) face in R. scabra, versus 2 broad parallel depressions in R. brasiliensis. Richardia brasiliensis is also adventive in the Paleotropics.

Fig. 613. Richardia scabra

72. RONABEA Aubl., Hist. Pl. Guiane 154, pl. 59. 1775. —Psychotria sect. Oppositiflorae Benth. & Hook., Gen. Pl. 2: 124. 1873. by Charlotte M. Taylor Shrubs or subshrubs, unarmed, terrestrial. Leaves opposite, petiolate to subsessile, venation not lineolate. Stipules persistent, interpetiolar or very shortly united around stem, triangular to subulate, in bud generally valvate to imbricate. Inflorescence axillary, subcapitate to congested-cymose, few- to several-flowered, subsessile to shortly pedunculate, bracteate. Flowers sessile, small, protandrous, at least sometimes distylous. Hypanthium turbinate. Calyx limb reduced, teeth or lobes 5, without calycophylls; corolla salverform, white, internally pubescent near middle of tube and sometimes on upper part of tube and lobes, lobes 5, valvate in bud. Stamens 5, inserted near or above middle of corolla tube; anthers narrowly oblong, included, dorsifixed near middle. Ovary 2-locular; ovules solitary in each locule, basal. Fruit ellipsoid, drupaceous, carnose, blue to black; pyrenes 2, bony, 1locular, hemispherical. Seeds ellipsoid, small. Belize to south-central Brazil; 3 species, 1 in Venezuela. Ronabea has long been included in Psychotria (e.g., Steyermark 1972; 1974) or considered closely related to it (e.g., Bremekamp 1934). but recent research supports its separation from Psychotria. For more information see the article by C.M. Taylor (2004).

802

R UBIACEAE

Fig. 614. Ronabea latifolia

Ronabea latifolia Aubl., Hist. Pl. Guiane 1: 154, t. 59. 1775, not Psychotria latifolia Humb. & Bonpl. ex Roem. & Schult. 1819. —Psychotria axillaris Willd., Sp. Pl. 1: 962. 1798, not Psychotria axillaris Vell. 1827, not Cephaelis axillaris Sw. —Ameu (Piaroa). Ronabea erecta Aubl., Hist. Pl. Guiane 156. 1775. —Psychotria erecta (Aubl.) Standl. & Steyerm., Publ. Field Mus. Nat. Hist., Bot. Ser. 23: 24. 1943. Coffea subsessilis Benth., Hook. J. Bot. 3: 232. 1841. —Mapouria subsessilis (Benth.) Müll. Arg., Flora 59: 460. 1876. —Psychotria fluctuans Standl., Publ. Field Columbian Mus., Bot. Ser. 7: 446. 1931, not Psychotria subsessilis Benth. 1852. —Psychotria erecta f. fluctuans (Standl.) Steyerm., Mem. New York Bot. Gard. 23: 713. 1972. Mapouria subsessilis var. [gamma]

angustifolia Müll. Arg. in Mart., Fl. Bras. 6(5): 411. 1881. —Psychotria fluctuans var. angustifolia (Müll. Arg.) Standl., Publ. Field Columbian Mus., Bot. Ser. 7: 447. 1931. Ronabea latifolia var. hispidula Bremek., Recueil Trav. Bot. Néerl. 31: 291. 1934. Suffruticose herb, shrub, or small tree to 6 m tall, strigillose to glabrescent; leaves 8.5– 17 × 2.5–9.5 cm; petioles 7–13 mm long; stipules 1–7 mm long; peduncles 1–4 per axil, 2–20 mm long; calyx limb 1–2 mm long; corolla tube 3–4 mm long, the lobes 1.5–3 mm long; fruit 8–10 × 5–8 mm. Lowland to montane forests, 50–1500 m; Bolívar (basins of Río Caroní and Río Caura), Amazonas (basins of Río Casiquiare, Río Pasimoni, Río Negro, and Río Orinoco). Apure, Sucre, Zulia; Central America, Colombia, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 614.

73. RONDELETIA L., Sp. Pl. 172. 1753. by Charlotte M. Taylor and Julian A. Steyermark Shrubs or small trees, terrestrial, unarmed. Leaves opposite or infrequently ternate, sessile to petiolate, venation not lineolate; stipules interpetiolar, triangular to foliaceous, persistent or caducous, in bud generally valvate to imbricate. Inflorescence terminal or less often axillary, multiflowered, pedunculate to sessile, cymose, corymbose, paniculate or spiciform, bracteate. Flowers sessile to pedicellate, distylous, medium to rather large, often rather showy and fragrant. Hypanthium turbinate to subglobose. Calyx limb deeply lobed, the lobes 4 or 5, equal to often markedly unequal in size and shape on an individual flower, sometimes with 1–several

Rondeletia 803

white to pink calycophylls; corolla salverform, red, purple, roseate, yellow, orange, or white, glabrous internally except with the throat sometimes barbate or with a conspicuous enlarged annulus, the lobes 4 or 5, imbricate in bud, the margins often crisped. Stamens 4 or 5, inserted in middle or upper part of the corolla tube; anthers narrowly oblong, dorsifixed, included or exserted. Ovary 2-locular; ovules numerous in each locule, on axile placentas. Fruit capsular, subglobose to rather didymous, woody, loculicidal. Seeds fusiform, small, winged, flattened. Mexico, Central America, West Indies, Colombia, Venezuela, Ecuador, Peru; 140 species, 7 in Venezuela (as circumscribed by Steyermark, 1974), 1 of these in the flora area. The circumscription of Rondeletia has been controversial. This genus or complex of genera includes relatively wide variation in number of corolla and calyx lobes, corolla throat morphology, capsule dehiscence, seed morphology, and type of pubescence. Lorence (1991; 1994) has recognized a comprehensive Rondeletia, pointing out that nearly all of the character states that have been used to separate genera can be found in nearly all combinations, at least in Central America where this group is most speciose. In contrast, Steyermark (1967, 241–261; 1974) recognized two genera in this group: Rondeletia generally has five calyx and corolla lobes, corolla throats that are glabrous and have a thickened ring, loculicidal capsules, flattened winged seeds, simple trichomes, and species that are generally found in dry and seasonal vegetation; R. orinocensis belongs to this group. Arachnothryx Planch. generally has four calyx and corolla lobes, corolla throats that are barbate and sometimes also have a thickened ring, septicidal capsules, angled seeds, frequently arachnoid or pannose pubescence, and species that are generally found in wet and montane vegetation. Study of the relationships in this group using molecular sequence data is just beginning. Rondeletia orinocensis Steyerm., Mem. New York Bot. Gard. 17(7): 247. 1967. —Kehkoyó (Panare). Shrub or small tree to 8 m tall, sericeous; leaves 6.5–17.5 × 2.5–8 cm; secondary veins 5–8 pairs, persistent; petioles 0–3 mm long; stipules 2–3 mm long; inflorescences 4–11 × 3–14 cm; calyx lobes 5, 3–4 mm long, generally equal; corolla white, tube 9–12 mm long, with an annulus in the throat, lobes ca. 4 mm long; capsules 6–8 × 7–9 mm, loculicidal; seeds fusiform, 2–2.5 × 1 mm. Riparian forests, forests near the base of low mountains, 50–200 m; Bolívar (Cerro Negro in Río Caura basin, near Guaniamo, near Maniapure, Río Parguaza), Amazonas (Capibara). Endemic. ◆Fig. 615.

Fig. 615. Rondeletia orinocensis

804

R UBIACEAE

74. ROSENBERGIODENDRON Fagerl., Svensk. Bot. Tidskr. 42: 150. 1948. by Charlotte M. Taylor Shrubs or small trees, terrestrial, unarmed. Leaves opposite, often grouped on short lateral branches, subsessile to petiolate, venation not lineolate; stipules interpetiolar or shortly united around stem, triangular, persistent, in bud generally valvate to imbricate. Inflorescences terminal on short lateral shoots, with flowers solitary, sessile, bracts reduced or none. Flowers homostylous, protandrous, large and showy, nocturnal, fragrant. Hypanthium cylindrical to ellipsoid. Calyx limb with well-developed tube, lobes 5(6), without calycophylls; corolla salverform with a prolonged tube, white sometimes flushed with green externally, internally sericeous in upper part of tube, lobes 5(6), convolute in bud. Stamens 5, inserted in upper part of corolla tube; anthers subsessile, narrowly oblong, dorsifixed, partially exserted. Ovary 1- or 2-locular; ovules numerous, on parietal placentas. Fruits baccate, subglobose to cylindrical, smooth, when young green with white spots and longitudinal stripes, when mature yellow to orange, with endocarp woody. Seeds medium-sized, flattened, smooth, lenticular. Panama, Colombia, Venezuela, Trinidad-Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 3 species, all in Venezuela, 2 of these in the flora area. Rosenbergiodendron is sometimes confused with Sphinctanthus; the distinctions between them are outlined under Sphinctanthus. Rosenbergiodendron was formerly included in Randia, but more recently has been separated based on its lack of spines, bisexual flowers, and pollen in monads, versus armed, unisexual, and in tetrads in Randia. Rosenbergiodendron has been monographed by Gustafsson (1998), and his species circumscriptions, followed here, differ in several respects from those of Steyermark (1974). The third species of Rosenbergiodendron, R. longiflorum (Ruiz & Pav.) Fagerl. [syn. Randia ruiziana DC.], is widely distributed in northern South America. It is known from the Guianas and northern Brazil and might be expected in the flora area. Rosenbergiodendron longiflorum is distinguished from the other species in the genus by its corollas with tubes 12–30 cm long and its lobes 3–12 cm long, and its fruits 5–10 × 2–4 cm. It is found in wet evergreen forests. Key to the Species of Rosenbergiodendron 1. 1.

Corolla tube 3–6.5 cm long; corolla lobes 1.2–3 cm long; leaf blades obtuse to acute at apex .................................................................... R. densiflorum Corolla tube 6–12 cm long; corolla lobes 2.5–4.5 cm long; leaf blades acute at apex ..................................................................................... R. formosum

Rosenbergiodendron densiflorum (K. Schum.) Fagerl., Svensk. Bot. Tidskr. 42: 52. 1948. —Randia formosa var. densiflora K. Schum. in Mart., Fl. Bras. 6(6): 343. 1889. —Punteral. Shrub to 4 m tall; leaves 1–7 × 0.7–3 cm; stipules 2–3 mm long; calyx limb with tube 2–4 mm long, lobes 3–6 mm long; corolla with tube 3–6.5 cm long, lobes 1.2–3 cm long; fruit

1.5–3 × 1–2 cm. Deciduous forests, thickets bordering savannas, savannas on igneous outcrops, 50–200 m; Delta Amacuro (Capure), Bolívar (Río Botanamo basin, Río Caroní, Río Cuyuní), Amazonas (Río Orinoco south of Puerto Ayacucho). Colombia, Trinidad, Tobago, Guyana, Suriname, French Guiana, Brazil.

Rudgea 805

Rosenbergiodendron formosum (Jacq.) Fagerl., Svensk Bot. Tidskr. 42: 152. 1948. —Mussaenda formosa Jacq., Enum. Syst. Pl. 16. 1760. —Randia formosa (Jacq.) K. Schum. in Mart., Fl. Bras. 6(6): 342. 1889. Shrub or small tree. Panama, Colombia, Venezuela, Ecuador; 2 varieties, both in Venezuela, 1 in the flora area. The other variety, var. formosum, is found in northern and western Venezuela. R. formosum var. nitidum (K. Schum.) C. Gust., Brittonia 50: 452. 1998. —Randia formosa var. nitida K. Schum. in Mart., Fl. Bras. 6(6): 343. 1889. Randia orinocensis Rusby, Descr. S. Amer. Pl. 132. 1920. Usually a shrub 2–3 m tall, sometimes a small tree to 6 m tall; leaves 3.5–9 × 2–4 cm; stipules ca. 3 mm long; calyx limb with tube 4–6 mm long, lobes 4–11 mm long; corolla tube 6–12 cm long, lobes 2.5–4.5 cm long; fruit 3–4.5 × 1.5–2.5 cm. Riparian forests, gallery forests, 50–200 m; Delta Amacuro (lower Río Orinoco), Bolívar (basins of Río Botanamo and Río Orinoco), Amazonas (basins of Río Casiquiare and Río Orinoco). Apure, Portuguesa. ◆Fig. 616.

Fig. 616. Rosenbergiodendron formosum var. nitidum

75. RUDGEA Salisb., Trans. Linn. Soc. London 8: 327, t. 18–19. 1807. by Daniela Zappi and Julian A. Steyermark Shrubs, subshrubs, or medium-sized trees, sometimes reaching 20 m. Leaves opposite, petiolate or sessile, venation not lineolate, on lower surface often with foveolate domatia in the axils of the secondary veins; stipules interpetiolar, usually conspicuous, in bud generally valvate to imbricate, persistent or sometimes caducous, ovate to spathulate, entire or usually fringed or appendiculate with the appendages often glandular and/or caducous, or navicular and then dorsally appendiculate with 3–several bristles, these sometimes glandular and/or caducous. Inflorescences terminal, subterminal, or subaxillary, sessile or usually pedunculate, multiflowered, cymose, thyrsoidal, dichasioid, congested, rarely glomerulate, or rarely reduced to a solitary flower, bracteate or the bracts reduced. Flowers fragrant, distylous. Calyx tubular to campanulate, truncate to lobed, lobes 4–6, without calycophylls; corolla broadly funnelform to narrowly tubular, white, cream, rarely purplish, or sometimes yellowish internally, in texture delicate to fleshy, 0.5– 7 cm long, internally glabrous to densely pubescent, lobes 4–6, valvate in bud, frequently dorsally corniculate. Stamens 4–6, inserted in the corolla throat or the lower part of the corolla tube; anthers dorsifixed, narrowly oblong, exserted or included, blue or purple. Ovary 2-locular; ovules 1 in each locule, basal. Fruit drupaceous, subglobose to ellipsoid, fleshy, spongy or moderately hard, white, green, yellow, orange, brown, red, black, or rarely purple; pyrenes 2, 1-locular, hemi-

806

R UBIACEAE

spherical, rounded to elliptic in outline, moderately hard, smooth or ridged, with 2 marginal, 1–3(–5) abaxial, and occasionally 1 adaxial pre-germination slit. Seed rather small, ellipsoid, deeply furrowed adaxially. Mexico, West Indies, Colombia, Venezuela, Trinidad-Tobago, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina; 140 species, 18 in Venezuela, 11 in the flora area. Several species here treated in Psychotria may be confused with Rudgea (and some of them may belong in Rudgea); these are included in the key below. Psychotria can be distinguished by its stipules that are caducous leaving a fringe of persistent trichomes, or persistent and variously shaped but without glandular appendages and usually bilobed. Key to the Species of Rudgea and Similar Species of Psychotria 1.

Stipules spathulate, devoid of appendages; inflorescence subaxillary, appearing lateral, from an elongated peduncle .................... R. stipulacea 1. Stipules triangular to ovate or naviculate, with 3–several appendages or flat and conspicuously fringed or lacerate; inflorescence terminal ..... 2 2(1). Stipules flat and conspicuously fringed along the margin ....................... 3 2. Stipules usually naviculate with apical rather than marginal appendages ................................................................................................................ 5 3(2). Peduncle > 10 cm long; inflorescence racemose, lax; corollas to 3 mm long ................................................................................................. R. woronowii 3. Peduncle to 5 cm long; inflorescence congested to capitate above the peduncle; corollas 5.5–80 mm long ........................................................... 4 4(3). Inflorescences subcapitate to congested-cymose; corollas 50–80 mm long ................................................................................................ R. lanceifolia 4. Inflorescences capitate; corollas ca. 5.5 mm long ....................................... .......................................................................... Psychotria sphaerocephala 5(2). Leaves 16–30 cm long, velutinous on lower surface .................... R. cardonae 5. Leaves to 15 cm long, glabrous on lower surface ..................................... 6 6(5). Inflorescence sessile, or subsessile with the peduncle to 1 cm long; corolla tube > 3 cm long ............................................................................. R. klugii 6. Inflorescence pedunculate with the peduncle > 1 cm long; corolla tube < 1.2 cm long .......................................................................................... 7 7(6). Inflorescence congested above the peduncle, with bracts obscuring the calyx limb or entire flowers and fruits ................................................. 8 7. Inflorescence lax, clearly branched above the peduncle, the bracts not obscuring the calyx limbs, flowers, or fruits ............................................ 9 8(7). Leaves coriaceous with revolute margins and the reticulum of veins very prominent .......................................................................... R. phaneroneura 8. Leaves chartaceous with flat margins and the reticulum of veins visible but not prominent ................................................................... R. wurdackii 9(7). Lower surface of leaves with foveolate domatia ......................... R. cornifolia 9. Lower surface of leaves without foveolate domatia ............................... 10 10(9). Calyx limb 2–4 mm long, with the tube developed, covering the nectar disk, the disk not visible; fruits white, crowned by the remnants of the calyx tube ............................................................................... R. sclerocalyx

Rudgea 807

10.

11(10). 11. 12(11). 12. 13(12). 13. 14(13). 14.

Calyx limb 0.5–1.5 mm long, the tube short or lacking, the nectar disk visible; fruits yellow, red, purple, or black, with or without persisting calyx limb remnants ............................................................................ 11 Peduncles equal to petioles in length; inflorescence usually shorter than the lower 1/3 of the leaf ............................................................ R. crassiloba Peduncles at least 2 times as long as the petioles; inflorescence longer than leaves or at least reaching the lower 1/2 of leaf .......................... 12 Stipules 10–13 mm long ...................................................... R. hostmanniana Stipules 1–5 mm long .............................................................................. 13 Stipules with caducous glands inserted at apex on each side ................... ................................................................................ Psychotria ventuariana Stipules with a group of caducous glands inserted medially or near base on each side .......................................................................................... 14 Stipules 3–5 mm long; branched portion of inflorescence 5–10 × 3–8 cm; corolla tube ca. 4 mm long ................................... Psychotria coussareoides Stipules 1–2 mm long; branched portion of inflorescence 4–6 × 2.5–3 cm; corolla tube 6.5–7 mm long ................................. Psychotria venezuelensis

Rudgea cardonae Steyerm., Mem. New York Bot. Gard. 17(1): 403, fig. 39. 1967. Shrub or small tree to 5 m tall; leaves petiolate, chartaceous, dull, the lower surface velutinous especially along the secondary and tertiary veins; blades (16–)18–30 × (6–) 8–10 cm, lanceolate, cuneate at base, acute to attenuate at apex, the margin flat, without domatia; stipules 20–22 × 6–8 mm, naviculate, triangular, hirsute, with 4–7 apical appendages 4–8 mm long; inflorescences with 30 or more flowers, terminal; peduncle 3–5 cm long, pubescent; flowers congested at the apex of short secondary branches; bracts triangular, pubescent at the margins; calyx ca. 1.5 mm long, turbiniform, short-velutinous, the tube short, truncate; corolla 5–6.5 cm long, tubular, short-villous externally, the lobes ca. 1/8 the length of tube; fruits not known. Riparian forests, 100–200 m; Amazonas (Río Castanho and basin of Río Padauiri on Venezuela-Brazil border). Brazil (Roraima). ◆Fig. 622. Rudgea cornifolia (Kunth ex Roem. & Schult.) Standl., Publ. Field Columbian Mus., Bot. Ser. 7: 432. 1932. —Psychotria cornifolia Humb. & Bonpl. ex Roem. & Schult., Syst. Veg. 5: 191. 1819; Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 3: 362. 1818 [1819]. —Cafecillo. Psychotria fimbriata Benth., J. Bot. (Hooker) 3: 226. 1841. —Rudgea fimbriata (Benth.) Standl. in Standl. &

S. Calderón, Lista Pl. Salvador 274. 1925. —Strempelia fimbriata (Benth.) Bremek., Recueil Trav. Bot. Néerl. 31: 303. 1934. Psychotria inundata K. Krause, Verh. Bot. Vereins Prov. Brandenburg 50: 108. 1908, nom. illeg., not Psychotria inundata Benth. 1841. Understory shrub or treelet 1–6 m tall; leaves shortly petiolate, membranaceous, dull, discolorous; blades 5–15 × (1.5–)2.3–7.5 cm, lanceolate to broadly-elliptic, attenuate to abruptly truncate at base, attenuate to caudate at apex, the margin undulate to broadly crenate, domatia almost always present; stipules 3–7 × 3–6 mm, convex, ovate, smooth, fringed to dorsally appendiculate, the fringes or appendages to 2 mm long; inflorescences (6–)12–24-flowered, terminal, minutely pubescent; peduncle 0.5–2.5 cm long; secondary axes (2)3–6, umbellately arranged; calyx 2–3.5 mm long, shortly turbinate, coriaceous, glabrous, the tube expanded, 5-toothed; corolla 4–9 mm long, funnelform to tubular, the lobes ca. 1/2–2/3 as long as tube; fruit 5–6 mm diameter, ovoid, sessile or on a peduncle to 3 mm long; pericarp palegreen to white, smooth; persisting calyx remnants to 0.8 mm broad at apex, prominent, tubular. Riparian forests, gallery forests, evergreen lowland forests, 50–300 m; Delta Amacuro (north of Río Orinoco), Bolívar (Río Acanán, Río Caura, middle Río Orinoco, Río Parhueña), Amazonas (Río Casiquiare, Río Negro, Río Orinoco). Apure; Mexico, Central

808

R UBIACEAE

America, Amazonian Colombia, Guyana, Suriname, Amazonian Brazil. ◆Fig. 619. Rudgea crassiloba (Benth.) B.L. Rob., Proc. Amer. Acad. Arts 45(17): 408. 1910. —Coffea crassiloba Benth., J. Bot. (Hooker) 3: 233. 1841. —Cereza, Punteral, Tortolito, Totumito. Rudgea schomburgkiana Benth., Linnea 23: 459. 1850. Shrub or small tree 2–10 m tall; leaves petiolate, membranaceous, dull, discolorous, often drying yellow to olive green; blades 7–9 × 2–4.5 cm, lanceolate to obovate, acute to attenuate at base, attenuate to caudate at apex, the margin flat to slightly revolute; domatia almost always present; stipules 2–4 × 3 mm, convex, ovate, smooth, dorsally appendiculate, the fringes or appendages to 1 mm long; inflorescences 18–24-flowered, terminal; peduncle 0.5–1.5 cm long; secondary axes 2–5, dichasioid; calyx 1–1.5 mm long, shortly turbinate, coriaceous, glabrous, the tube absent, lobes triangular, narrow, acute; corolla 8–10 mm long, tubular, lobes ca. 1/4–1/2 as long as tube; fruit 6–7 mm diameter, ovoid, sessile; pericarp bright orange, smooth or ridged; persisting calyx remnants to 0.5 mm broad at apex, not tubular. Riparian forests and semideciduous to evergreen lowland to lower montane forests, 50–400 m; Bolívar (Río Coro Coro west of Cerro Yutajé, lower Río Orinoco, Río Paragua), Amazonas (Río Ventuari). Barinas, Portuguesa; Guyana, Brazil. ◆Fig. 620. Rudgea hostmanniana Benth., Linnaea 23: 459. 1850. Shrub or treelet; leaves petiolate, chartaceous; blades 4–14(–15) × 1.5–8 cm, variable in shape; domatia absent; stipules 10–13 × 6–7 mm, naviculate, triangular, with 3–7 apical appendages to 5 mm long; inflorescences with 20 or more flowers, terminal; peduncle (1.5–) 3–8 cm long; secondary axes in 1–3 levels, the flowers congested at branch apices; bracts triangular; calyx 1–1.5 mm long, turbiniform, glabrous, tube short, lobes 5, acute; corolla 6–8 mm long, short-tubular, lobes ca. 1/3 as long as tube, internally pubescent; fruit 4–10 × 4–8 mm, ovoid, sessile, ridged; pericarp yellow to orange-red or black when ripe. Evergreen or semideciduous gallery forests, savannas, 100–1100 m; Guyana,

Suriname, French Guiana, Brazil; 3 subspecies, all in Venezuela, 2 in the flora area. Rudgea hostmanniana subsp. freemanii (Sprague & R.O. Williams ex R.O. Williams & Cheesman) Steyerm. is distinguished from the other subspecies of R. hostmanniana by its densely appressed-pubescent hypanthium, externally pubescent calyx lobes, and shorter peduncle. It is found in the Coastal Cordillera of Venezuela. Key to the Subspecies of R. hostmanniana 1. Leaves dull above, the margin flat, the base acute to cuneate; fruit 7–10 mm long .................. subsp. hostmanniana 1. Leaves shiny above, the margin revolute in mature specimens, the base cuneate to subcordate; fruit 4–6.5 mm long ............................ subsp. maypurensis R. hostmanniana subsp. hostmanniana. —Cafecillo, Casco de mula, Comida de danta, Danta oreja de burro, Dantero. Rudgea bolivarensis Steyerm., Mem. New York Bot. Gard. 17(1): 415. 1967. Rudgea corocoroensis Steyerm., Ann. Missouri Bot. Gard. 75: 349. 1988. Shrub or small tree 1–15 m tall; leaves 8– 14(–15) × 4–8 cm, lanceolate to broadly ovate, dull, drying yellow to gray-green, with secondary veins prominent in both faces, acute to cuneate at base, acute to obtuse at apex, the margin flat; inflorescences with 50 or more flowers, terminal; peduncle 3.5–8 cm long; secondary axes in 2 or 3 levels; fruit 7– 10 × 6–8 mm, pericarp red or black. Evergreen or semideciduous gallery forests, 200– 1100 m; Delta Amacuro (Santa Catalina), Bolívar (Altiplanicie de Nuria, Gran Sabana, Río Botanamo, basin of Río Kukenán, Río Supamo, Santa Elena de Uairén, Serranía de Imataca, Upata), Amazonas (Cerro Yutajé). Anzoátegui, Barinas, Monagas; Guyana, Suriname, French Guiana, Brazil. ◆Fig. 623. R. hostmanniana subsp. maypurensis (Standl.) Zappi, comb. & stat. nov. —Rudgea maypurensis Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 7: 434. 1931; emend. Steyerm., Mem. New York Bot. Gard. 17: 411. 1967. Shrub 1–4 m tall; leaves 4–9 × 1.5–3.5 cm, lanceolate to oblong or rarely broadly ovate,

Rudgea 809

shiny above, drying yellow, with the secondary veins obscure below, cuneate, obtuse, rounded, or subcordate at base, slightly acute to obtuse at apex, the margins revolute to rolled in mature specimens; inflorescences 20–30-flowered, terminal; peduncle 1.5–6 cm long; secondary axes in 1 or 2 levels; fruit 4– 6.5 × 3–5 mm; pericarp orange to red. Scrub forests bordering igneous outcrops, savannas, riparian forests bordering igneous exposures, 100–200 m; Amazonas (region of Puerto Ayacucho, Sabana de Oso along upper Río Ventuari). Endemic. ◆Fig. 618. Rudgea klugii Standl., Field Mus. Nat. Hist., Bot. Ser. 13(6): 164. 1936. —Palo culebra. Rudgea standleyana Steyerm., Mem. New York Bot. Gard. 17(1): 409. 1967. Rudgea berryi Steyerm. & Dwyer, Pittieria 9: 12. 1981. Epiphytic, lianescent shrub to 5 m tall; leaves petiolate, chartaceous, moderately shiny, discolorous, gray-green to olive green when dry, new growth drying black; blades 4–15 × 1.4–6 cm, narrowly elliptic to obovate, attenuate at base, the apex 1–2 cm long, acuminate to caudate, the margin flat to slightly revolute, with the secondary veins prominent on both surfaces; venation with the reticulum fine and evident; domatia absent; stipules 2–2.2 × 1.5–3 mm, convex, broadly ovate, smooth, dorsally appendiculate, appendages to 1 mm long, sometimes deciduous; inflorescences 6–9-flowered, terminal, subsessile; peduncle absent or rarely to 1 cm long; bracts reduced; bracteoles linear-triangulate, acute, to 1 mm long; flowers sessile to subsessile; calyx 2–3.5 × 1.5–2 mm, cylindric, coriaceous, smooth, tube inconspicuous, lobes acute, to 1 mm long; corolla 32–47 mm long, narrowly tubular, glabrous, lobes ca. 1/6–1/5 as long as tube, with cornicula to 1.5 mm long; fruits ca. 10 mm long, ellipsoid; pericarp orange to brown. Evergreen lowland and lower montane forests, 100–400 m; Amazonas (Cerro Duida, San Carlos de Río Negro). Guyana, Suriname, French Guiana, Ecuador, Peru, northern Brazil. ◆Fig. 624. Rudgea lanceifolia Salisb., Trans. Linn. Soc. London 8: 327, t. 18. 1807. Rudgea ovalifolia Salisb., Trans. Linn. Soc. London 8: 328, t. 19. 1807.

Rudgea fissistipula Müll. Arg., Flora 59: 449, 460. 1876. Rudgea carolina Standl. & Steyerm., Fieldiana, Bot. 28: 614. 1953. Rudgea sipapoensis Steyerm., Mem. New York Bot. Gard. 17(1): 411. 1967. Shrub to large forest tree to 13 m tall; leaves petiolate, chartaceous, upper surface moderately shiny; blades (10–)15–24 × (6–) 8–17 cm, lanceolate to broadly ovate, cuneate to truncate at base, rounded to apiculate at apex, the margin flat, the lower surface with the secondary veins prominent; domatia absent; stipules 15–25(–35) × 10–15 mm, flat, margin deeply fringed to laciniate, the appendages 4–12(–20) mm long; inflorescences 50–multiflowered, terminal; peduncle 3–5 cm long, pubescent; flowers congested at the apex of the peduncle or on short secondary axes; bracts navicular, sometimes fringed; calyx 2–3 mm long, turbiniform, glabrous to short-velutinous, tube short to absent, lobes 5, lanceolate; corolla 5–8 cm long, tubular, short-villous externally, lobes ca. 1/10 as long as tube; fruit 15–25 × 12–17 mm, ovoid, sessile; pericarp yellow to orange-red when ripe, persisting calyx remnants forming a broad, subapical circular scar. Evergreen lowland to lower montane forests, 100–600 m; Amazonas (Cerro Sipapo, region of San Carlos de Río Negro). Colombia, Guyana, Suriname, French Guiana, Peru, Amazonian Brazil. ◆Fig. 625. Rudgea phaneroneura Steyerm., Mem. New York Bot. Gard. 17(1): 403, fig. 40. 1967. Small tree 3–4 m tall; leaves petiolate, coriaceous, shiny; blades (2–)3–9 × (1.5–)2–5 cm, broadly ovate to elliptic, rounded at base, rounded to apiculate or acute at apex, the margin revolute, with the vein reticulum prominent; domatia absent; stipules 5–8 × 4– 7 mm, flat or convex, ovate, pubescent, fringed or sometimes dorsally appendiculate, the appendages to 7 mm long; inflorescences 8–12-flowered, terminal; peduncle 0.5–2 cm long, pubescent; flowers congested, covered by long, triangular bracts; calyx 1.5–2 mm long, rotate, glabrous, tube absent, lobes 5, acute; corolla ca. 5 mm long, funnelform to tubular, lobes ca. 1/2 as along as tube; fruit ca. 15 × 10 mm, ovoid, sessile; pericarp green to brown. Tepui slope and summit, 1400–1900

810

R UBIACEAE

m; Bolívar (Cerro Venamo, Macizo del Chimantá, Matawi-tepui). Endemic. ◆Fig. 627. Rudgea sclerocalyx (Müll. Arg.) Zappi, comb. nov. —Mapouria sclerocalyx Müll. Arg., Flora 59: 496. 1876. Coffea laurifolia Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 3: 372. 1818 [1819], hom. illeg., not Coffea laurifolia Salisb. 1796, nom. illeg. superfl. —Rudgea laurifolia Steyerm., Mem. New York Bot. Gard. 17(1) 423. 1967. Mapouria spruceana Müll. Arg., Flora 59: 496. 1876. Rudgea morichensis Steyerm., Mem. New York. Bot. Gard. 17(1): 424. 1967. Shrub or treelet 2–8 m tall; leaves petiolate, chartaceous, shiny, discolorous; blades (5–)9–14 × (2–)2.5–6 cm, narrowly lanceolate to broadly elliptic, attenuate to rounded at base, attenuate to caudate at apex, margin undulate; domatia absent; stipules 2–3 × 3 mm, convex, truncate, smooth, dorsally appendiculate, the very short appendages sometimes deciduous; inflorescences 30–50flowered, terminal; peduncle 2–3.5 cm long; secondary axes flattened, 3–7, dichasioid; calyx 2–4 mm long, campanulate, glabrous, the tube expanded, 5-toothed to 5-lobed; corolla 11–12 mm long, funnelform to tubular, lobes ca. 1/4–1/3 as long as tube; fruit ca. 15 mm diameter, ovoid, sessile or on a peduncle to 3 mm long; pericarp pure white to pale green, the persisting calyx remnants to 2 mm broad at apex, prominent, tubular, yellow. Swampy or seasonally flooded forests, savanna formations, riparian forests, occasionally flooded scrub, 50–800 m; Amazonas (Cerro Moriche, Río Casiquiare, Río Negro basin, Río Orinoco). Amazonian Colombia, Brazil. ◆Fig. 628. Steyermark correctly determined that Coffea laurifolia Kunth is the oldest published name for this species, and attempted to publish a combination in Rudgea based on this name. However, Kunth’s name duplicates a previously published name, and thus is illegitimate and was not available to be used as a basionym by Steyermark. Consequently Steyermark’s name is not a combination and its publication date is 1967. Several other names for this species were published before Steyermark’s and thus have priority, and a new combination for one of these, Mapouria sclerocalyx, is made here. This name is chosen over M. spruceana because

the epithet of this latter name is quite similar to another Rudgea name, R. sprucei Standl., and could generate confusion. Rudgea stipulacea (DC.) Steyerm., Mem. New York Bot. Gard. 17 (1): 421. 1967. —Coffea stipulacea DC., Prodr. 4: 499. 1830. —Ají de gallineta, Cafecillo, Palo culebra, Simarjo. Ixora duidae Standl., Publ. Field Columbian Mus., Bot. Ser. 7: 412. 1931. —Rudgea duidae (Standl.) Steyerm., Mem. New York Bot. Gard. 17(1): 422. 1967. Psychotria maguirei Standl., Bull. Torrey Bot. Club 75: 578. 1948. Psychotria avia Standl. & Steyerm., Fieldiana, Bot. 28: 596. 1953. —Rudgea avia (Standl. & Steyerm.) Steyerm., Mem. New York Bot. Gard. 23: 405. 1972. Shrub or small tree 0.5–4 m tall; leaves petiolate, membranaceous, dull, discolorous; blades (16–)22–27 × (4.5–)6.5–9 cm, broadly lanceolate to obovate, acute to cuneate at base, attenuate, acute, or acuminate at apex, the margin flat; domatia absent; stipule (8–) 15–20 × 4–6 mm long, striate, entire to nearly entire at apex or erose with 1–4 subulate glandular-tipped teeth; inflorescences 50–multiflowered, axillary; peduncle 8–15 cm long, winged, glabrous; flowers borne on 3–6 secondary axes branches; bracts and bracteoles inconspicuous; calyx 1–1.5 mm long, rotate, glabrous, tube short, lobes 5, poorly defined; corolla ca. 5 mm long, tubular, lobes ca. 1/4 as long as tube; fruit 5–9 × 7– 9 mm, depressed-globose to ovoid, sessile; pericarp violet-black. Evergreen lowland to montane forests, gallery forests, 50–1100 m; Bolívar (base of Cerro Sarisariñama, south of El Dorado), Amazonas (Cerro Aracamuni, Cerro Duida, Cerro Huachamacari, Río Cataniapo, Río Cuao, Río Guainía, basin of Río Negro, Río Puruname, Río Siapa, Río Sipapo, Sierra de la Neblina, Tamatama). Amazonian Colombia, Guyana, Suriname, French Guiana, northern Brazil. ◆Fig. 621. Rudgea woronowii Standl., Publ. Field Columbian Mus., Bot. Ser. 7: 155. 1930; emend. Steyerm., Mem. New York Bot. Gard. 17: 406. 1967. Single-stemmed shrub 1–1.5 m tall; leaves petiolate, membranaceous to chartaceous,

Rudgea 811

moderately shiny; blades 20–30 × (8–)10–12 cm, broadly ovate, acute at base, rounded to apiculate at apex, the margin flat, with the secondary veins prominent; domatia absent; stipules 20–40 × 5–6 mm, flat, ridged, deeply fringed to laciniate, the appendages 6–10 mm long; inflorescences 50–multiflowered, terminal; peduncle 10–15 cm long; secondary axes racemose, numerous; calyx ca. 0.5 mm

long, rotate, glabrous, tube absent, lobes 5, acute; corolla ca. 1 mm long, tubular, lobes ca. 1/3 as long as tube; fruit ca. 5 × 4 mm, ovoid, sessile; pericarp green to brown. Evergreen lowland to lower montane forests, 100– 600 m; Bolívar (Río Abacapá, Río Hacha affluent of Río Caroní, Río Icabarú, Río Paramichí basin), Amazonas (Cerro Aracamuni, upper Río Orinoco, Suhinya on Río Siapa). Amazonian Colombia, Peru, and Brazil. ◆Fig. 617.

Fig. 617. Rudgea woronowii Fig. 618. Rudgea hostmanniana subsp. maypurensis

812

R UBIACEAE

Fig. 619. Rudgea cornifolia

Fig. 620. Rudgea crassiloba

Fig. 621. Rudgea stipulacea

Rudgea 813

Fig. 622. Rudgea cardonae

Fig. 623. Rudgea hostmanniana subsp. hostmanniana

814

R UBIACEAE

Fig. 624. Rudgea klugii

Fig. 625. Rudgea lanceifolia

Rudgea 815

Fig. 626. Rudgea wurdackii

Fig. 627. Rudgea phaneroneura

Fig. 628. Rudgea sclerocalyx

816

R UBIACEAE

Rudgea wurdackii Steyerm., Mem. New York Bot. Gard. 17(1): 413. 1967. Shrub to 2.5 m tall; leaves subsessile, chartaceous, shiny, slightly discolorous; blades 8– 14.5 × (2–)2.5–5 cm, narrowly lanceolate to oblong, rounded at base, attenuate to acute at apex, the margin flat; domatia absent; stipules 2–3 × 3 mm, convex, truncate, smooth, dorsally appendiculate, the very short appendages sometimes deciduous; inflorescences 15– 30-flowered, terminal; peduncle 2–4 cm long;

secondary axes none or very short, condensed above the peduncle; bracts triangular, to 3 mm long, covering the calyx; calyx 2–4 mm long, campanulate, glabrous, tube expanded, 5toothed; corolla ca. 12 mm long, funnelform, lobes ca. 1/3 as long as tube; fruit ca. 10 mm diameter, ovoid, sessile; pericarp white, with persisting calyx remnants to 2 mm broad at apex, prominent, tubular, yellow. River margins, 50–100 m; Amazonas (Río Atabapo, Río Orinoco). Endemic. ◆Fig. 626.

76. SABICEA Aubl., Hist. Pl. Guiane 192, t. 75, 76. 1775. by Charlotte M. Taylor and Julian A. Steyermark Suffrutescent climbers, less often shrubs, these sometimes from a xylopodium, terrestrial, unarmed, often with pannose, floccose, or arachnoid pubescence. Leaves opposite or verticillate, petiolate or sessile, venation not lineolate; stipules interpetiolar or sometimes shortly united around the stem, ligulate to ovate or triangular, persistent, frequently reflexed, in bud generally erect and flatly appressed or valvate. Inflorescences axillary, sessile to pedunculate, several- to multiflowered, glomerulate to capitate, cymose, or paniculate, bracteate. Flowers sessile to pedicellate, homostylous and protandrous or distylous, generally medium-sized. Hypanthium turbinate to ellipsoid. Calyx limb lobed, lobes 5, subequal to unequal but without calycophylls; corolla salverform to funnelform, white to pink, internally hirtellous at stamen insertion and sometimes in lower part of tube, lobes 5, valvate in bud. Stamens 5, inserted in corolla throat or upper part of corolla tube; anthers narrowly oblong, dorsifixed near middle, included or partially exserted. Ovary 5-locular; ovules numerous in each locule, on axile placentas. Fruit baccate, subglobose, fleshy to juicy, red to purple sometimes turning black. Seeds small, angled, reticulate. Mexico, Central America, Antilles, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Africa, Madagascar; 120–130 species, 16 in Venezuela, 12 of these in the flora area. Sabicea is often confused with Manettia, which is similar in habit. Their distinctions are discussed under Manettia. Steyermark (1967, 307–316; 1974) and other authors have considered the presence and distribution of arachnoid or pannose-floccose pubescence informative for circumscribing species of Sabicea. This character has not been studied in terms of its variability at a population level. Key to the Species of Sabicea 1. 1. 2(1). 2. 3(2).

Inflorescence cymose to capitate, these and/or flowers pedunculate and/or pedicellate .............................................................................................. 2 Inflorescence glomerulate, these and the flowers all sessile ................... 4 Plants strigose to strigillose, without arachnoid or pannose-floccose pubescence ..................................................................................... S. bariensis Plants with arachnoid or pannose-floccose pubescence at least on lower surfaces of leaves ................................................................................... 3 Calyx limb 1.5–2.5 mm long, the lobes ≤ 1/2 length of limb, externally densely strigose; leaves 8–19 × 3.5–9 cm; corolla lobes ca. 4 mm long

Sabicea 817

............................................................................................... S. grandifolia Calyx limb 2.5–3 mm long, almost completely lobed, externally arachnoid-pubescent; leaves 6.5–9.5 × 1.5–3 cm; corolla lobes 2–2.5 mm long ............................................................................................... S. morillorum 4(1). Lower surfaces of leaves glabrous to strigose, hirsute, or villous ........... 5 4. Lower surfaces of leaves arachnoid or pannose-floccose pubescent, the pubescence sometimes thinning with age ............................................ 7 5(4). Corolla tubes 4.5–5.5 mm long; calyx limbs 3–4.5 mm long, deeply lobed, the lobes reflexed after anthesis .................................................. S. villosa 5. Corolla tubes 9–15.5 mm long; calyx limbs 2–5 mm long, shallowly to deeply lobed, the lobes ascending or reflexed after anthesis ............... 6 6(5). Calyx limb divided for 1/2 or less its length, the lobes deltoid to triangular or broadly so, after anthesis ascending to somewhat spreading .............................................................................................. S. brachycalyx 6. Calyx limb divided for > 1/2 its length, the lobes narrowly triangular to linear, reflexed after anthesis .............................................. S. glabrescens 7(4). Upper surfaces of leaves arachnoid-pubescent ......................................... 8 7. Upper surfaces of leaves strigillose to strigose, hirsute, pilose, or pilosulous ....................................................................................................... 9 8(7). Calyx lobes 1.5–2.5 mm wide; corollas externally hirsute or pilose, i.e., with spreading trichomes ............................................................ S. cinerea 8. Calyx lobes 0.4–0.5 mm wide; corollas externally sericeous, i.e., with appressed trichomes ......................................................................... S. tillettii 9(7). Calyx limb 9–15 mm long; corolla tube 12–17 mm long ....... S. amazonensis 9. Calyx limb 2.5–7.5 mm long; corolla tube 5–11.5 mm long ................... 10 10(9). Corolla externally hirsute, the trichomes widely spreading; pubescence of lower surface of leaves rather tufted .................................. S. oblongifolia 10. Corolla externally sericeous to strigose, the trichomes appressed to a little spreading; pubescence of lower surface of leaves dense and even .............................................................................................................. 11 11(10). Calyx limb 4.5–7.5 mm long; corolla tube 8–11.5 mm long ......... S. velutina 11. Calyx limb 2.5–5 mm long; corolla tube 5–5.5 mm long ...... S. venezuelensis 3.

Sabicea amazonensis Wernham, Monogr. Sabicea 47, pl. 5, figs. 3, 4. 1914. Vine; stems hirsute; leaves 7–13 × 2–6 cm, adaxially pilosulous to hirsute, abaxially lanose; secondary veins 8–15 pairs; petioles 5–10 mm long; stipules 8–15 mm long; inflorescences glomerate, sessile; calyx limb 9–15 mm long, deeply lobed; corolla externally hirsute, tube 12–17 mm long, lobes 4–5 mm long; fruits ca. 1 cm diameter. Riparian forests, evergreen lowland forests, 100–200 m; Amazonas (basins of Río Atabapo, Río Guainia, and Río Yatúa). Amazonian Colombia, Peru, and Brazil. ◆Fig. 629. Sabicea romboutsii Bremek. of Suriname is similar to S. amazonensis and can be distinguished by its leaves ternate rather than opposite.

Sabicea bariensis Steyerm., Ann. Missouri Bot. Gard. 75: 350. 1988. Vine; stems strigose; leaves 11–13 × 5.5–7 cm, strigose; secondary veins 10–13 pairs; petioles 2–3.5 cm long; stipules 8–12 mm; inflorescences congested-cymose; peduncles 4– 10 mm long; pedicels 3–5 mm long; calyx limbs 3.5–5.5 mm long; corolla externally strigose, tube ca. 10.5 mm long, lobes ca. 4 mm long; fruits not seen. Riparian forests, 100–200 m; Amazonas (upper Río Baría, Río Mawarinuma, Río Pasimoni). Endemic. Several other Sabicea species of northern and western Venezuela also have simple pubescence and branched inflorescences, and probably at least some should be expected in the flora area. Sabicea panamensis Wernham also has appressed, strigose to strigillose

818

R UBIACEAE

pubescence, but can be distinguished by its narrow calyx lobes and corollas with tubes 8– 9 mm long externally with spreading pubescence; this species is widespread and weedy in Central America and western South America. Three species have spreading pilosulous to hirsute pubescence: S. liesneri Steyerm. of the Coastal Cordillera of Venezuela is distinguished by its ligulate to lanceolate calyx lobes 3–5 mm long, versus lanceolate to ovate and 6–8 mm long in S. novogranatensis K. Schum. and S. aristeguietae Steyerm. Sabicea brachycalyx Steyerm., Mem. New York Bot. Gard. 17(1): 313. 1967. —Carillo de perro, Chino. Vine; stems strigose or the trichomes sometimes spreading a bit; leaves 4.5–10.5 × 2.5–4.5 cm, on both surfaces strigillose to strigose; secondary veins 8–12 pairs; petioles 5–10 mm long; stipules 3–8 mm long; inflorescences glomerate, sessile; calyx limbs 2.5– 4 mm long, lobed to 1/2 their length; corolla externally hirsute, tube 11–15.5 mm long, lobes 3.5–4 mm long; fruits 4–6 mm long. Riparian forests, edges of Mauritia palm swamps, savanna borders, 100–500 m; Bolívar (basins of Río Caroní and Río Caura), Amazonas (Río Casiquiare, basin of Río Negro at confluence with Río Orinoco, Río Ventuari). Northern Brazil (Amazonas, Roraima). ◆Fig. 633. Sabicea cinerea Aubl., Hist. Pl. Guiane 193, t. 75. 1775. Vine; stems arachnoid-pubescent; leaves 5–11.5 × 2–6 cm, adaxially sparsely arachnoid-pubescent, abaxially densely arachnoid or lanose; secondary veins 11–15 pairs; petioles 5–10 mm long; stipules 5–9 mm long; inflorescences glomerate, sessile; calyx limbs 5–11 mm long, deeply lobed; corollas externally pilose to hirsute, tube 10–17 mm long, lobes 4–6 mm long; fruits 5–6 mm diameter. Lowland moist forests, expected in the flora area at 200–300 m. Sucre; Suriname, French Guiana. Sabicea cinerea is known from Sucre at 280 m and in Suriname and French Guiana down to nearly sea level. It might eventually be found in the flora area. See also comments under S. tillettii. Sabicea glabrescens (K. Schum.) Benth., J. Bot. (Hooker) 3: 219. 1841. —Sabicea

aspera var. glabrescens K. Schum. in Mart., Fl. Bras 6(6): 307. 1889. —Tabquillo. Vine or clambering shrub; stems strigose or the trichomes a bit spreading; leaves 7–13 × 2.5–5 cm, on both surfaces strigillose to strigose; secondary veins 8–13 pairs; petioles 3–15 mm long; stipules 5–10 mm long; inflorescences glomerate, sessile; calyx limbs 3–5 mm long, deeply lobed; corollas externally hirsute, tube 9–11.5 mm long, lobes 3–4 mm long; fruits 3–5 mm long. Riparian forests, 50–200 m; Delta Amacuro (Araguabia), Bolívar (lower and middle Río Caura, Río Cuyuní basin). Guyana, Suriname, French Guiana, Brazil. Sabicea grandifolia Steyerm. in Lasser & Steyerm., Fl. Venez. 9: 514. 1974. Vine; stems arachnoid-pubescent; leaves 8–19 × 3.5–9 cm, adaxially glabrescent to strigillose, abaxially densely arachnoid-pubescent and also sometimes strigillose; secondary veins 11 or 12 pairs; petioles 8–25 mm long; stipules 6–10 mm long; inflorescences congested-cymose; peduncles 1–12 mm long; pedicels 3–5 mm long; calyx limb 1.5–2.5 mm long, lobed for 1/2 or less; corolla externally strigose, tube ca. 11 mm long, lobes ca. 4 mm long; fruits ca. 5 mm diameter. Riparian forests, evergreen lowland forests on hummocks, 100–200 m; Amazonas (near Piedra de Cucuy, Río Guainía, Río Baría). Endemic. ◆Fig. 631. Sabicea morillorum Steyerm. in Lasser & Steyerm., Fl. Venez. 9: 511. 1974. Vine; stems arachnoid-pubescent; leaves 6.5–9.5 × 1.5–3 cm, adaxially puberulent, abaxially densely arachnoid-pubescent; secondary veins 11–13 pairs; petioles 8–12 mm long; stipules 6–8 mm long; inflorescences cymose; peduncles 5–7 mm long; pedicels 2.5–5 mm long; calyx limb 2.5–3 mm long, deeply lobed; corolla externally strigose, tube 8–10 mm long, lobes 2–2.5 mm long; fruits not seen. Río Negro caatinga, 50–200 m; Amazonas (basins of Río Guainía and Río Negro). Endemic. ◆Fig. 632. Sabicea oblongifolia (Miq.) Steyerm., Mem. New York Bot. Gard. 17(1): 316. 1967. —Sabicea velutina var. oblongifolia Miq., Linnaea 18: 615, 735. 1844. —Sabicea glabrescens var. oblongifolia

Sabicea 819

(Miq.) Sandw., Kew Bull. 1939: 12. 1939. Vine; stems hirsute; leaves 7–15 × 3–8.5 cm, adaxially pilosulous to strigillose, abaxially tufted-arachnoid; secondary veins 9–13 pairs; petioles 4–15 mm long; stipules 4–9 mm long; inflorescences glomerate, sessile; calyx limbs 1–3 mm long, deeply lobed; corolla externally hirsute, tube 9–10.5 mm long, lobes 2.5–3 mm long; fruits ca. 8 mm diameter. Lowland and riparian forests, 50–200 m; Delta Amacuro (Río Amacuro), Bolívar (Río Botanamo, Río Corumo), Amazonas (Río Negro basin). Apure, Barinas, Portuguesa; Trinidad, Guyana, Suriname, French Guiana. Sabicea tillettii Steyerm., Phytologia 31: 484. 1975. Vine; stems arachnoid- or lanose-pubescent; leaves 4.5–10 × 1.7–3.2 cm, adaxially arachnoid-pubescent, abaxially densely lanose; secondary veins 15 or 16 pairs; petioles 4–10 mm long; stipules 8–9 mm long; inflorescences glomerate, sessile; calyx limbs 9.5– 10 mm long, deeply lobed; corolla externally sericeous, tube ca. 12 mm long, lobes ca. 4 mm long; fruits not seen. Riparian forests bordering igneous outcrops, 100–200 m; Amazonas (near Caño Chamuchina in Río Atabapo basin). Endemic. Sabicea velutina Benth., J. Bot. (Hooker) 3: 219. 1841. —Sabicea aspera var. velutina (Benth.) K. Schum. in Mart., Fl. Bras. 6(6): 307. 1889. Sabicea guianensis Wernham, Monogr. Sabicea 52. 1914, nom. illeg., non (Aubl.) Baill. 1880. Sabicea leucotricha K. Krause, Notizbl. Bot. Gart. Berlin-Dahlem 6: 202. 1914. Vine; stems arachnoid-pubescent and pilose to hirsute; leaves 5–12 × 2–5.5 cm, adaxially pilosulous to pilose, abaxially densely lanose; secondary veins 10–15 pairs; petioles 3–15 mm long; stipules 8–14 mm long; inflorescences glomerate, sessile; calyx limb 4.5–7.5 mm long, deeply lobed; corolla externally sericeous, tube 8–11.5 mm long, lobes 3–4 mm long; fruits 5–8 mm diameter. Savannas, rocky slopes, 400–1800 m. Venezuela, Trinidad, Guyana, French Guiana, Brazil (Roraima: Rio Branco); 3 subspecies, all in the flora area. Steyermark’s subspecies of Sabicea velutina are of questionable separation and taxo-

nomic value, but little material was seen so they are provisionally treated here. Sabicea velutina is similar to S. brasiliensis Wernham of central Brazil, and these should be compared critically. Key to the Subspecies of S. velutina 1. Petioles mainly 3–6 mm long, rarely to 15 mm; leaf blades rounded, subcordate, or broadly obtuse at base, mainly 1.3–2.7 (–3) times as long as broad; heads usually 10–16-flowered ... subsp. velutina 1. Petioles mainly 8–15 mm long; leaf blades acute to obtuse at base, generally 2.2– 3.3 times as long as broad; heads usually 16–25-flowered .................................. 2 2. Corolla 15–15.5 mm long, the tube 11.5 mm long; heads 20–25-flowered; anthers ca. 2 mm long .... subsp. chimantensis 2. Corolla 12–12.5 mm long, the tube 9 mm long; heads 16–20-flowered; anthers 1.2–1.3 mm long ..... subsp. duidensis S. velutina subsp. chimantensis Steyerm., Mem. New York Bot. Gard. 17(1): 315. 1967. Bolívar (Macizo del Chimantá). Endemic. S. velutina subsp. duidensis Steyerm., Mem. New York Bot. Gard. 17(1): 316. 1967. Amazonas (Cerro Duida, Río Matacuni, upper Río Orinoco, Simarawochi). Endemic. S. velutina subsp. velutina Bolívar (Auyán-tepui, Cerro Toribio, Gran Sabana, Kurún-tepui), Amazonas (Sierra Parima). Táchira; Trinidad, Guyana, French Guiana, Brazil. ◆Fig. 630. Sabicea venezuelensis Steyerm., Mem. New York Bot. Gard. 17(1): 309, fig. 35. 1967. —Bejuco salvia. Vine; stems strigose to hirsute; leaves 4– 13 × 1.5–4 cm, adaxially hirtellous to strigillose, abaxially densely arachnoid; secondary veins 5–10 pairs; petioles 3–6 mm long; stipules 4–8 mm long; inflorescences glomerate, sessile; calyx limb 2.5–5 mm long, deeply lobed; corolla externally strigose, tube 5–5.5 mm long, lobes 2–2.5 mm long; fruits not seen. Savannas, gallery forests, borders of Mauritia palm swamps, scrub savannas, 50– 300 m; Bolívar (Río Caroní, basin of Río

820

R UBIACEAE

Fig. 629. Sabicea amazonensis

Fig. 630. Sabicea velutina subsp. velutina

Fig. 631. Sabicea grandifolia

Sabicea 821

Fig. 632. Sabicea morillorum

Fig. 633. Sabicea brachycalyx

Fig. 634. Sabicea venezuelensis

822

R UBIACEAE

Orinoco, Río Paragua), Amazonas (near Puerto Ayacucho). Apure, Guárico; Colombia. ◆Fig. 634. Sabicea venezuelensis may not be distinct from Sabicea camporum Sprague of eastern Colombia, but too little information is available to evaluate this. Steyermark separated S. camporum based on its generally erect habit, obtuse leaves, and more acute calyx lobes, but in the specimens now available of S. venezuelensis there is greater morphological variation than he saw. Sabicea villosa Willd. ex Roem. & Schult., Syst. Veg. 5: 265. 1819. Sabicea hirsuta Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 3: 417. 1818 [1820]. Vine or clambering shrub; stems strigose to hirsute; leaves 7–13.5 × 2.5–7 cm, on both surfaces strigose to hirsute; secondary veins 7–14 pairs; petioles 5–15 mm long; stipules 5–8 mm long; inflorescences glomerulate, sessile; calyx limb 3–4.5 mm long, deeply lobed; corolla externally strigose to hirsute, tube 4.5–5.5 mm long, lobes 1.5–2 mm long; fruits 4–5 mm diameter. Roadsides, cut-over and secondary forests, riparian banks, shrubby savannas, 50–800 m. Southern Mexico, Central America, Colombia, Ecuador, Peru, Brazil, Bolivia. Although the two varieties below differ only in pubescence and are both generally widespread, they were recognized by Andersson and Ståhl (1999), who listed dis-

tributional differences in these varieties in Ecuador. In the flora area, the plants assignable to var. adpressa are generally (though not exclusively) found in the northern and western portions, while those assignable to var. villosa are found in the southern and eastern regions of the Guianas. Key to the Varieties of S. villosa 1. Pubescence of stems and lower surfaces of the leaves strigose, appressed ................ ....................................... var. adpressa 1. Pubescence of stems and lower surfaces of the leaves hirsute, spreading .................. .......................................... var. villosa S. villosa var. adpressa (Wernham) Standl., Publ. Field Columbian Mus., Bot. Ser. 7: 52. 1930. —Sabicea hirsuta var. adpressa Wernham, Monogr. Sabicea 55. 1914. Delta Amacuro (Caño Araguao), Bolívar (near Icabarú, basin of Río Cuchivero), Amazonas (basins of Río Negro and Río Orinoco). Barinas, Monagas, Zulia; throughout the range of the species. S. villosa var. villosa Delta Amacuro (Río Cuyubini), Bolívar (near Santa Elena), Amazonas (Mavaca, upper Río Orinoco, Sierra Parima). Apure, Mérida, Zulia; throughout the range of the species.

77. SCHRADERA Vahl, Eclog. Amer. 1: 35, pl. 5. 1796 [1797]. Fuchsia Sw., Prodr. 62. 1788, nom. illeg., non L. 1753. Urceolaria Willd. in Cothen., Dispos. Veg. 10. 1790. Lucinaea DC., Prodr. 4: 368. 1830. by Charlotte M. Taylor and Julian A. Steyermark Epiphytic shrubs or lianas, generally succulent and glabrous, unarmed, stems often with adventitious roots. Leaves opposite, petiolate, venation not lineolate and usually not visible; stipules interpetiolar or united around the stem, generally erect and appressed in bud, caducous, ligulate to obovate. Inflorescences terminal and/or axillary, several- to multiflowered, capitate or the flowers solitary on each peduncle, the heads or solitary flowers surrounded by a reduced to well-developed, green, truncate involucre, otherwise bracts reduced, the peduncles solitary or 2–7 and fasciculate. Flowers sessile, medium-sized to rather large, fragrant, nocturnal, fleshy, frequently distylous. Hypanthium turbinate to subglobose. Calyx limb truncate to unduluate, without calycophylls; corolla salverform, white, internally pubescent in upper part of tube and in throat, the lobes 5 or 6(–10), valvate in bud, often fleshy and triangular in cross section and prolonged at apex into an appendage. Stamens 5

Schradera 823

or 6(–10), inserted in upper part of corolla tube; anthers narrowly oblong, dorsifixed, included or partially exserted. Ovary 2(–4)-locular; ovules numerous in each locule, axile. Fruit baccate, subglobose, fleshy, mature color unknown. Seeds small, suborbicular, flattened or angled. Mexico, Central America, the Antilles, Colombia, Venezuela, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia, southeast Asia to New Guinea; 65 species, 12 in Venezuela, 8 of these in the flora area. Schradera is similar to Hillia; for their distinctions see the discussion under Hillia. Puff et al. (1993) concluded that the paleotropical genus Lucinaea DC. is synonymous with Schradera. Schradera is poorly known, rather infrequently collected, and only rarely found with mature flowers. Species distinctions and the morphological variation within species in this genus are both poorly understood. Steyermark (1964) placed great weight on patterns of pubescence to separate species, but this character has not been well evaluated in this group. Juvenile and flowering stems are frequently heteromorphic: the juvenile stems have smaller, thinner-textured leaves and adhere to their supporting substrate by adventitious roots, while the flowering stems are frequently spreading, have tougher-textured, usually larger leaves, and lack adventitious roots. These flowering stems may be several meters long and are sometimes pendulous. Key to the Species of Schradera 1. 1. 2(1). 2. 3(2). 3. 4(3). 4. 5(4). 5. 6(5). 6. 7(5). 7.

Apex of peduncle and/or involucre moderately to densely pilosulous .................................................................................................... S. nilssonii Apex of peduncle and involucre glabrous or sparsely puberulous ........... 2 Peduncles 2 or more at stem apex ....................................... S. bipedunculata Peduncles solitary at stem apex ................................................................ 3 Peduncle 0.5–1 cm long; leaves 3.5–5 × 1–2.5 cm ....................... S. maguirei Peduncle 2–14 cm long; leaves 6.5–13 × 1.5–6.5 cm ................................ 4 Outer basal half of stipule puberulent; stigmas 3 ..................... S. yutajensis Stipule completely glabrous; stigmas 2 .................................................... 5 Leaf blades generally obtuse to rounded at the apex ............................... 6 Leaf blades acute to acuminate at the apex ............................................. 7 Calyx and hypanthium ca. 10 mm long; corolla tube ca. 6 mm long ......................................................................................... S. marahuacensis Calyx and hypanthium 7–8.5 mm long; corolla tube 7–11 mm long ......... .............................................................................................. S. polycephala Leaves with secondary veins 9–10 pairs and blades rounded or obtuse at base; corolla ca. 10 mm long ...................................................... S. brevipes Leaves with secondary veins 7 or 8 pairs or not visible and blades subacute to acute at the base; corolla 25–26 mm long ................ S. hilliifolia

Schradera bipendunculata Steyerm., Mem. New York Bot. Gard. 10(5): 263. 1963. Plant climbing high; leaves 13–15.5 × 5–8 cm; petioles 20–27 mm long; stipules not known; heads 2 at stem apex, 7–10-flowered; peduncles 5.5–8.5 cm long; involucres 6–7 mm long; flowers not known; fruits 12–16 × 8–10 mm with persistent calyx limb 3–4 mm

long. Montane forests, 500–600 m; Bolívar (Río Uiri-yuk in Río Cuyuní basin). Endemic. Other species of Schradera in northeastern South America with several fasciculate peduncles are S. suaveolens (H. Karst.) Steyerm. of the Venezuelan Coastal Cordillera, with small flower heads, 12–15 × 17–20 mm; S. ternata Steyerm. of Guyana, with 8 corolla lobes; and S. polycephala DC., which

824

R UBIACEAE

is only known to have solitary heads in the flora region but frequently has 2 or 3 heads in the Guianas and Brazil. Schradera brevipes Steyerm., Mem. New York Bot. Gard. 10(5): 263. 1963. Vine-like epiphyte; leaves 9–13.5 × 4.5–9 cm; petioles 1–2 cm long; stipules 15–20 mm Fig. 635. Schradera polycephala

Fig. 637. Schradera nilssonii

long; heads solitary, terminal, 9- or 10-flowered; peduncles 2–3.5 cm long; involucres 7– 10 mm long; calyx limb ca. 7 mm long; corolla with tube ca. 5 mm long, lobes 6, ca. 5 mm long; fruits not known. Upper tepui slope forests, ca. 1400 m; Amazonas (Cerro Sipapo). Endemic.

Fig. 636. Schradera hilliifolia

Simira 825

Schradera hilliifolia Steyerm., Mem. New York Bot. Gard. 10(5): 269. 1963. —Matapalo. Climbing epiphyte; leaves 6.5–11.5 × 3.5– 5.5 cm; petioles 1–1.5 cm long; stipules 1.5–2 cm long; heads solitary, terminal, 7–15-flowered; peduncles 4–5.5 cm long; involucres 10–16 mm long; calyx limb 8–12 mm long; corolla with tube ca. 16 mm long, lobes 5, ca. 10 mm long; fruits not known. Evergreen lowland forests, 100–200 m; Amazonas (Maroa, base of Sierra de la Neblina). Endemic. ◆Fig. 636. Schradera maguirei Steyerm., Mem. New York Bot. Gard. 10(5): 27. 1963. Shrubby plant; leaves 3.5–5 × 1–2.5 cm; petioles 4–5 mm long; stipules 5–7 mm long; heads solitary, terminal, 3–5-flowered; peduncles 0.5–1 cm long; involucres 2–3 mm long; calyx limb ca. 7 mm long; corolla with tube ca. 6 mm long, lobes 5, ca. 6 mm long; fruits not known. Tepui summit forests, ca. 1700 m; Amazonas (Cerro Guaiquinima). Endemic. Schradera marahuacensis Steyerm., Ann. Missouri Bot. Gard. 74: 114. 1987. Liana; leaves 9.5–13 × 2.5–5.5 cm; petioles 5–16 mm long; stipules ca. 1 cm long; heads solitary, terminal, 10–20-flowered; peduncles 3.5–6 cm long; involucres 7–8 mm long; calyx limb ca. 7 mm long; corolla tube ca. 6 mm long, lobes 5, ca. 5 mm long; fruits not known. Tepui slope forests, 1100–1300 m; Amazonas (Cerro Marahuaka). Endemic. Schradera nilssonii Steyerm., Bol. Soc. Venez. Ci. Nat. 23: 83. 1962. Liana, climbing to 20 m high; leaves 13–

15 × 6–6.5 cm; petioles 1–2 cm long; stipules 15–20 mm long; heads solitary, terminal, 20– 40-flowered; peduncles 5–10.5 cm long; involucres 7–10 mm long; calyx limb ca. 5 mm long; corolla with tube ca. 7.5 mm long, lobes 5, ca. 4.5 mm long; fruits not seen. Tepui slope forests, 700–1600 m; Bolívar (Macizo del Chimantá, Río Cuyuní basin). Guyana. ◆Fig. 637. Schradera polycephala A. DC. in DC., Prodr. 4: 444. 1830. Liana; leaves 5–12 × 1.5–5.5 cm; petioles 5–20 mm long; stipules 10–16 mm long; heads solitary (1 or 3 in the Guianas), terminal, 8–20-flowered; peduncles 2.5–7 cm long; involucres 3–10 mm long; calyx limb 3.5–5 mm long; corolla with tube 7–11 mm long, lobes 5 or 6, 4–11 mm long; fruits not seen. Elfin mossy forests, montane forests, tall forests along streams, usually overlying sandstone or quartzitic substrate at the higher altitudes, rarely at low altitudes, 200–1400 m; Bolívar (upper Río Orinoco, Sierra Parima). Guyana, Suriname, French Guiana, Amazonian Brazil. ◆Fig. 635. Schradera yutajensis Steyerm., Mem. New York Bot. Gard. 10(5): 276, fig. 70g– h. 1963. Shrubby plant; leaves 7–11.5 × 3–6.5 cm; petioles 10–14 mm long; stipules 12–20 mm long; heads solitary, terminal, 9–12-flowered; peduncles 3–5 cm long; involucres ca. 8 mm long; calyx limb ca. 8 mm long; corolla with tube ca. 10.5 mm long, lobes 5, ca. 6 mm long; fruits not known. Seasonally dry tepui slope forests and scrub, ca. 1500 m; Amazonas (Cerro Yutajé). Endemic.

78. SIMIRA Aubl., Hist. Pl. Guiane 170, pl. 65. 1775, not Raf. 1836. Sickingia Willd., Ges. Naturf. Freunde Berlin Neue Schriften 3: 445. 1801. by Charlotte M. Taylor and Julian A. Steyermark Shrubs or trees, terrestrial, unarmed, with the internal tissues often turning red or rose-purple when exposed to air. Leaves opposite, petiolate to subsessile, venation not lineolate; stipules interpetiolar, persistent or caducous, triangular, erect, twisted in bud. Inflorescence terminal and in the axils of the uppermost leaves, multiflowered, pedunculate to sessile, paniculate to congested-cymose or subcapitate, bracteate with the bracts sometimes reduced. Flowers often very fragrant, homostylous, protandrous, subsessile to pedicellate, sometimes a little zygomorphic. Hypanthium turbinate to cupuliform. Calyx limb truncate or usually 4-or 5(6)-lobed, in some flowers of some species 1 lobe enlarged into a pink, red, white, or

826

R UBIACEAE

cream petaloid calycophyll; corolla funnelform to campanulate, white, pale green, or pale yellow, internally glabrous or often with a pubescent ring at or below stamen insertion, lobes 4 or 5(6), imbricate or rarely open in bud. Stamens 4 or 5(6), inserted below middle of corolla tube; anthers narrowly oblong, dorsifixed near the middle, exserted. Ovary 2-locular; ovules numerous in each locule, on axile placentas. Fruit capsular, ellipsoid to globose, loculicidally dehiscent from apex, woody. Seeds large, lunulate to oblong, flattened, chartaceous to papery, winged. Southern Mexico, Central America, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; ca. 35 species, 8 in Venezuela, 3 of these in the flora region. This genus was long treated under the name Sickingia, and its recent change to Simira for purely nomenclatural reasons has caused some confusion. Most specimens from northern South America have been called Simira rubescens or S. cordifolia, but Simira is poorly known taxonomically and these names are undoubtedly applied too broadly. Steyermark considered pubescence details, particularly those of the abaxial leaf surface, to be taxonomically informative; however with more collections now available this character seems to vary widely within species. Steyermark (1974) reported an additional species from outside the flora area in eastern Venezuela, Simira myriantha (Standl.) Steyerm., but that species has been more recently treated as Elaeagia myriantha (Standl.) C.M. Taylor & Hammel. Key to the Species of Simira 1. 1. 2(1). 2.

Leaves oblanceolate, narrowed to an acute or cuneate base .. S. pisoniiformis Leaves elliptic to ovate or obovate, at least shortly obtuse to rounded or cordate at the base ................................................................................. 2 Inflorescences paniculate, branched to several orders, with the flowers well separated on developed axes; calyx limb ca. 1 mm long ... S. ignicola Inflorescences subcapitate to congested-cymose, unbranched or branched to 1 order, with the flowers congested, hardly separated; calyx limb 2.5–3 mm long .......................................................................... S. rubescens

Simira ignicola Steyerm., Ann. Missouri Bot. Gard. 75: 1086. 1988. Tree to 15 m tall; leaves 17–19 × 11–11.5 cm; secondary veins 13–16 pairs; petioles 1– 1.5 cm long; stipules caducous, not known; inflorescence pyramidal, 5–6 × 5–8 cm; calyx limb ca. 1 mm long, lobed, without calycophylls; mature corolla and capsules not known. Gallery forests along streams on igneous substrate, 50–200 m; Bolívar (Quebrada la Flore in Río Parguaza basin). Endemic. Simira pisoniiformis (Baill.) Steyerm., Mem. New York Bot. Gard. 72: 305. 1972. —Sickingia pisoniiformis Baill., Adansonia 12: 307. 1879. Ixora podocarpa Benth. in Benth. & Hook. f., Gen. Pl. 2: 113. 1873. Tree to 8 m tall; leaves 15–30 × 3.5–10 cm; secondary veins 15–19 pairs; petioles 0.5–3 cm long; stipules 5–6 mm long, persistent; in-

florescences pyramidal, 6–12 × 5–10 cm; calyx limb ca. 1 mm long, lobed, without calycophylls; corolla white, tube 5–7 mm long, lobes 5, 0.5–1 mm long; capsules obpyriform, 1.3–2 × 1.5–2 cm, stipitate; seeds 8–10 mm long. Seasonally flooded and riparian forests, 100–200 m; Amazonas (Río Casiquiare, basin of Río Negro). Northern Brazil. ◆Fig. 639. Simira pisoniiformis is remarkably similar to S. longifolia (Willd.) Bremek. of the Venezuelan Coastal Cordillera. Simira longifolia differs in its leaves with more numerous secondary veins (22–25 pairs), the tertiary venation impressed adaxially, and its corollas with tubes 4–4.5 mm long and lobes 1–1.5 mm long. Simira rubescens (Benth.) Bremek. ex Steyerm., Mem. New York Bot. Gard. 23: 301. 1972. —Sprucea rubescens Benth., Hooker’s J. Bot. Kew Gard. Misc. 5: 230.

Simira 827

Fig. 638. Simira rubescens

Fig. 639. Simira pisoniiformis

1853. —Cafecillo, Paraguatán, Paraguatán blanco. Simira erythroxylon var. saxicola Steyerm., Acta Bot. Venez. 6: 112. 1971. Shrub or tree to 10 m tall; leaves 9–19 × 5–13 cm; secondary veins 6–12 pairs; petioles 6–12 mm long; stipules 5–25 mm long; inflorescences rounded, 1.5–2 × 2–3.5 cm; calyx limb 2.5–3 mm long, lobed, without calycophylls; corolla 4–5 mm long, lobes 5(6), 2.5–3 mm long; capsules subglobose, 2.5–4 × 3–4.5 cm; seeds 1.8–2 cm long. Riparian forests, sa-

vannas, shrubby vegetation on igneous outcrops (lajas), 50–200 m; northern Bolívar (widespread), Amazonas (near Puerto Ayacucho, Santa Barbara del Orinoco, and near Brazil border at Río Padauiri). Amazonian Brazil, probably also in the Amazon basin of Colombia and Peru. ◆Fig. 638. Simira rubescens has been mistakenly called S. tinctoria Aubl. by some authors, but that name applies to a tree of wet forests that is currently known only from French Guiana. Simira rubescens is circumscribed differently

828

R UBIACEAE

here than by Steyermark (1972; 1974). With more collections now available this species is clearly variable in leaf size and shape and the presence versus absence of pubescence on the lower leaf surfaces. Pubescence was the only character Steyermark used to separate S. erythroxylon var. saxicola from S. rubescens and combine it with S. erythroxylon, a

tree of wet premontane forests; consequently S. erythroxylon var. saxicola, a tree of seasonally dry lowland forests, is here treated as a synonym of S. rubescens. Simira rubescens appears to be quite common in the flora area, but very few flowering collections are available, and more than one species may be included here.

79. SIPANEA Aubl., Hist. Pl. Guiane 147. 1775. by Piero G. Delprete and Julian A. Steyermark Annual or perennial herbs, terrestrial, unarmed; stems erect, spreading, or creeping. Leaves opposite, sessile to petiolate, venation not lineolate; stipules interpetiolar, persistent, in bud generally imbricate to valvate, reduced or well developed, rounded to triangular, subulate, or divided. Inflorescence axillary and/or terminal, few-flowered to multiflowered, sessile to pedunculate, bracteate but without floral bracts, cymose or reduced to 1–3 flowers, frequently branched 1 or 2 times with the subsequent axes monochasial. Flowers sessile or shortly pedicellate, small to medium-sized, homostylous, protandrous. Hypanthium turbinate to ellipsoid. Calyx limb deeply lobed, lobes 4 or 5, well developed, narrow, without calycophylls. Corolla salverform, rose or roseate to white, internally barbate in the mouth with white or yellow trichomes, these included or exserted, lobes 5, convolute in bud. Stamens 5, inserted below the middle of the corolla tube or rarely in its upper part; anthers linear, dorsifixed, included or partially exserted. Ovary 2-locular; ovules numerous in each locule, on axile placentas. Fruit capsular, ellipsoid, chartaceous, loculicidal from the apex. Seeds small, angulate; testa foveolate-reticulate. Central America, Colombia, Venezuela, Guyana, Suriname, French Guiana, Peru, Brazil, Bolivia, Paraguay; about 17 species, 9 in Venezuela, all in the flora area. Key to the Species of Sipanea 1. 1. 2(1). 2. 3(2). 3. 4(3).

4.

5(1).

Stems trailing, sprawling, or rooting at the nodes ................................... 2 Stems mainly erect or ascending .... .......................................................... 5 Corolla ca. 7.5 mm long, the tube ca. 4 mm long, the lobes ca. 3.5 mm long, moderately pubescent externally ....................................... S. setacea Corolla 14–22 mm long, the tube 8–15 mm long, the lobes 4.5–11 mm long, glabrous externally ....................................................................... 3 Stipules triangular-lanceolate, 2–3 mm long ............................... S. carrenoi Stipules suborbicular, lunulate, or broadly triangular, sometimes depressed or split at the center, 0.1–1 mm long ....................................... 4 Orifice of the corolla white-villous within, the trichomes not exserted; 1 glandular appendage present in each sinus between the calyx lobes; leaf blades 8–26 mm wide; petiole 4–18 mm long ....................... S. biflora Orifice of the corolla densely yellow-villous within, the trichomes exserted; 2 glandular appendages present in each sinus between the calyx lobes, one on each side of the sinus; leaf blades 2–17 mm wide; petiole 0.5–6 mm long ............................................................................ S. veris Orifice of the corolla glabrous or at least not yellow-villous ...................... ......................................................................................... S. wilson-brownei

Sipanea 829

5. 6(5).

6.

7(6).

7.

8(7).

8.

Orifice of the corolla densely yellow-villous ............................................. 6 Corolla white; outermost bracts enveloping or surrounding the inflorescence, broadened or rounded at the base, often irregularly lobed or lacerate ........................................................................................ S. glomerata Corolla pale to dark pink or rarely white; outermost bracts not enveloping or surrounding the inflorescence, narrowed at the base, not lobed, lacerate, or cleft ..................................................................................... 7 Leaves linear to linear-lanceolate, 5–8 times as long as broad, with the secondary veins faint, 2 or 3 on each side, not manifest on upper surface; stems mostly simple; corolla tube 15–18 mm long ......... S. galioides Leaves linear-lanceolate to elliptic or elliptic-, ovate-, or oblong-lanceolate, 3.5–7 times as long as broad, with the secondary veins mostly manifest, 3–7 on each side, usually evident on the upper surface; stems usually branched; corolla tube 6–15 mm long ...................................... 8 Corolla lobes rounded to truncate; stems and leaves densely and conspicuously hirsute or villous; capsules densely hispid with spreadingascending trichomes ....................................................................S. hispida Corolla lobes acute or apiculate; stems and leaves glabrescent to densely pubescent, the trichomes when present appressed to ascending or rarely sparsely spreading; capsules glabrescent or sparsely to densely pubescent with erect, appressed, or ascending trichomes ...... S. pratensis

Sipanea biflora (L. f.) Cham. & Schltdl., Linnaea 4: 168. 1829. —Virecta biflora L. f., Suppl. 134. 1781 [1782]. Sipanea radicans Endl., Atakt. Bot. 7, fig. 7. 1833. Herb, trailing, decumbent at terminal nodes; stems to 0.5 m long, rooting at nodes; leaves short- to long-petiolate, blades 1.5–4.8 × 0.8–2.6 cm; stipules entire or bifid, 0.5–1 mm long; calyx lobes 2.5–4.5 mm long; corolla tube 10–16 mm long, the lobes rounded or obtuse at apex, 4.5–8 × 2–2.5 mm; capsules 4 × 3–4 mm. Riparian forests, semideciduous to evergreen lowland to lower montane forests, 50–800 m; Delta Amacuro (Serranía de Imataca), Bolívar (Altiplanicie de Nuria, Cerro Paují, Río Cuyuní). Guyana, Suriname, French Guiana, Brazil. ◆Fig. 642. Sipanea carrenoi Steyerm., Ann. Missouri Bot. Gard. 71: 332. 1984. Herb, trailing or sprawling; stems to 0.5 m long, rooting at nodes; leaves subsessile to short-petiolate, blades 0.9–1.8 × 0.5–0.9 cm; stipules entire, narrowly triangular-lanceolate, 2–3 mm long; calyx lobes ca. 4.5 mm long; corolla tube 10–13 mm long, the lobes rounded or obtuse at apex, 6–6.5 × 5–7 mm; capsules 9 × 2 mm. Low places in savannas bordering gallery forests, moist sandstone slopes below waterfall bordering savannas,

400–1200 m; Bolívar (Gran Sabana, affluents of Río Caroní). Endemic. ◆Fig. 646. Sipanea galioides Wernham, J. Bot. 55: 172. 1917. Perennial herb; stems ascending or prostrate, 0.1–0.6 m long; leaves subsessile, blades 2–4 × 0.3–2 cm; stipules entire, triangular, 1.5–2 mm long; calyx lobes 5–8 mm long; corolla tube 15–18 mm long, the lobes acute to obtuse at apex, 6–10.5 × 5–6 mm; capsules 5–7.5 × 3–4 mm. Dry sandy savannas, bare lateritic slopes of savannas, borders of dwarf forests, riparian forests, open tepui slopes, 700–1800 m; Bolívar (Gran Sabana, upper Río Caroní). Guyana. ◆Fig. 641. Sipanea glomerata Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 3: 398. 1818 [1820]. Sipanea glomerata var. paucinervia Steyerm., Mem. New York Bot. Gard. 17(1): 282. 1967. Herb, erect, much branched, 0.3–1.5 m tall; leaves subsessile to short-petiolate, blades 3–13 × 1–4.5 cm; stipules entire, broadly triangular, 4–11 mm long; calyx lobes 6–13 mm long; corolla tube 12–14 mm long, the lobes rounded or obtuse at apex, 7– 8 × 3.5–5.5 mm; capsules 7–9 × 3.5–5.5 mm. Forested and open igneous outcrops, gravelly savannas, igneous outcrops along streams,

830

R UBIACEAE

50–200 m; Amazonas (basin of Río Orinoco between Río Sipapo and Río Cataniapo). Colombia, Amazonian Brazil. ◆Fig. 643. Sipanea hispida Benth. ex Wernham, J. Bot. 55: 173. 1917. Colombia, Venezuela, Peru, Brazil, Bolivia, Paraguay; 2 varieties, 1 in Venezuela. The second variety of this species, var. major (Hassl.) Steyerm., is found in Paraguay. S. hispida var. hispida Herb, erect, sparsely branched, 0.1–0.8 m tall; leaves short-petiolate, blades 1.2–7.3 × 0.6–3 cm; stipules subulate, 4–13 mm long; calyx lobes 4–6 mm long; corolla tube 6–10 mm long, the lobes rounded to truncate at apex, 2.5–5 × 1.5–3 mm; capsules 6–11 × 3–5 mm. Hilly savannas, savannas bordering Mauritia palm swamps, 50–500 m; Bolívar (middle Río Orinoco), Amazonas (Río Orinoco). Carabobo, Distrito Federal, Miranda; Colombia, Peru, Brazil, Bolivia, Paraguay. ◆Fig. 645. Sipanea pratensis Aubl., Hist. Pl. Guiane 147. 1775. Colombia, Venezuela, Trinidad, Guyana, Suriname, French Guiana, Brazil; 2 varieties, 1 in Venezuela. Sipanea pratensis is highly variable morphologically. Its second variety, var. pratensis, is found in the Guianas and Brazil. S. pratensis var. dichotoma (Kunth) Steyerm., Mem. New York Bot. Gard. 17(1): 273. 1967. —Sipanea dichotoma Kunth in H.B.K., Nov. Gen. Sp. (quarto ed.) 3: 397. 1818 [1820]. —Conejo rebalsero, Lazo, Rebalsera, Sardina, Siempre viva, Yurima. Sipanea colombiana Wernham, J. Bot. 55: 174. 1917. Sipanea pratensis var. dichotoma f. brachycarpa Steyerm., Mem. New York Bot. Gard. 17(1): 276. 1967. Sipanea pratensis var. dichotoma f. glabriloba Steyerm., Mem. New York Bot. Gard. 17(1): 276. 1967. Sipanea pratensis var. dichotoma f. glabrior Steyerm., Mem. New York Bot. Gard. 17(1): 277. 1967. Sipanea pratensis var. dichotoma f. breviflora Steyerm., Mem. New York Bot. Gard. 17(1): 277. 1967.

Sipanea pubinoda Steyerm., Mem. New York Bot. Gard. 17(1): 278. 1967. Herb, erect, sparsely branched, 0.2–1 m tall; leaves sessile to short-petiolate, blades 1.7–7.5 × 0.5–2.5 cm; stipules entire, triangular-lanceolate or narrowly triangular, 2–6 mm long; calyx lobes 2–6 mm long; corolla tube (8–)10–17 mm long, the lobes acute or apiculate at apex, 4–11 × 3–7 mm; capsules 5–8 × 2–3 mm. Marshy and semiflooded savannas, scrub savannas on white sand, savannas bordering lakes or Mauritia palm swamps, Trachypogon savannas with Curatella trees, 50–500 m; Delta Amacuro (Serranía de Imataca), Bolívar (Altiplanicie de Nuria, Río Caroní, Río Cuyuní, Río Yuruaní, Serranía de Imataca), Amazonas (Río Guainía, Río Negro, Río Ocamo, Río Orinoco, Río Pacimoni). Guyana. ◆Fig. 644. Sipanea setacea Steyerm., Ann. Missouri Bot. Gard. 74: 114. 1987. Herb, trailing; stems 0.1–0.15 m long; leaves short-petiolate, blades 1.2–2 × 0.9–1.2 cm; stipules setaceous, 2–3 mm long; calyx lobes 2.2–2.5 mm long; corolla tube ca. 4 mm long, the lobes obtuse at apex, ca. 3.5 × 1.2 mm; capsules unknown. Open scrub on tepui summits, ca. 1300 m; Bolívar (summit of Cerro Venado). Endemic. Sipanea veris S. Moore, Trans. Linn. Soc. London, Bot. 4: 368. 1895. Sipanea acinifolia Spruce ex Sprague, Trans. & Proc. Bot. Soc. Edinburgh 22: 433. 1905. Sipanea spraguei Wernham, J. Bot. 55: 172. 1917. Herb, trailing; stems rooting at nodes, 0.1–0.7 m long; leaves sessile or short-petiolate, blades 0.5–2.6 × 0.2–1.7 cm; stipules broadly triangular to lunate, 0.1–0.5 mm long; calyx lobes 2–6 mm long; corolla tube 8– 15 mm long, the lobes acute to obtuse at apex, 5–11 × 2.5–8 mm; capsules ca. 7 × 3 mm. Moist stream banks, margins of lakes, along streams, borders of Mauritia palm swamps, low periodically flooded sites, edges of gallery forests, wet savannas, 50–200 m; Bolívar (Río Botanamo, lower Río Caura, Río Cuchivero, middle Río Orinoco), Amazonas (Río Orinoco, Río Ventuari). Anzoátegui, Guárico; Colombia, Brazil. ◆Fig. 647. Sipanea wilson-brownei Brittonia 7: 412. 1952.

R.S.

Cowan,

Sipanea 831

Fig. 640. Sipanea wilson-brownei

Fig. 641. Sipanea galioides

Bertiera palustris A. Rich in DC., Prodr. 4: 392. 1830. —Sipanea palustris (A. Rich.) J.H. Kirkbr., Brittonia 49: 360. 1997, hom. illeg., not Sipanea palustris Seem. [=Limnosipanea palustris (Seem.) Hook. f.]. Herb, erect, sparsely to densely branched, 0.2–1 m tall; leaves short- to long-petiolate, blades 2–7 × 0.9–3.5 cm; stipules entire, broadly triangular, 2–5 mm long; calyx lobes 3.5–8 mm long; corolla tube 9–15 mm long, the lobes obtuse at apex, 4–8 × 2.5–5 mm; capsules 4.5–6.5 × 5.5–6 mm. Lower montane to montane forests, usually most common among igneous outcrops and over sandstone, 500–1300 m; Bolívar (basin of Río Cuyuní). Guyana, French Guiana, Brazil (Pará: Oriximiná). ◆Fig. 640.

Fig. 642. Sipanea biflora

832

R UBIACEAE

Fig. 643. Sipanea glomerata

Fig. 645. Sipanea hispida var. hispida

Fig. 646. Sipanea carrenoi

Fig. 644. Sipanea pratensis var. dichotoma

Fig. 647. Sipanea veris

Sipaneopsis 833

80. SIPANEOPSIS Steyerm., Mem. New York Bot. Gard. 17(1): 284. 1967. by Piero G. Delprete and Julian A. Steyermark Shrubs or subshrubs, terrestrial, unarmed; stems erect or prostrate. Leaves opposite, subsessile to petiolate, venation not lineolate; stipules interpetiolar, persistent, in bud apparently valvate to imbricate, triangular to deeply bifid or multifid. Inflorescence terminal, cymose, few-flowered to multiflowered, sessile to pedunculate, bracteate. Flowers small to medium-sized, sessile to pedicellate, homostylous, protandrous. Hypanthium ellipsoid. Calyx limb lobed, lobes 5 or 6, equal to a little unequal, without calycophylls; corolla salverform, roseate, pink, or reddish, in the mouth densely barbate and with 5 triangular appendages, the lobes 5 or 6, imbricate in bud. Stamens 5, inserted in the upper part of the corolla tube; anthers included, linear, dorsifixed near base. Ovary 2-locular; ovules several to numerous in each locule, on axile placentas. Fruit dry, subglobose to ellipsoid, apparently indehiscent, chartaceous. Seeds small, globose, foveolate. Amazonian Colombia, Venezuela, Amazonian Brazil; 6 species, all in the flora area. Sipaneopsis is endemic to white-sand areas of the basins of Río Orinoco and Río Amazon. Key to the Species of Sipaneopsis 1. 1. 2(1). 2. 3(2). 3.

4(1). 4. 5(4).

5.

Some or all of the stipules bifid to multifid, split to the base; corolla tube 6–11 mm long; plants 0.3–2 m tall ........................................................ 2 All the stipules entire; corolla tube mainly 15–22 mm long; plants 0.1–0.4 m tall ...................................................................................................... 4 Leaf blades glabrous on both surfaces or the midrib and lateral veins strigose below .................................................................................. S. rupicola Leaf blades pubescent on both surfaces .................................................... 3 Stems strigose, the trichomes appressed; peduncle sericeous with appressed trichomes ......................................................................... S. huberi Stems densely villosulous, the trichomes loosely spreading or ascending; peduncle hirsutulous with spreading to ascending trichomes .............................................................................................. S. morichensis Lower surface of leaves glabrate to sparsely strigillose ............. S. maguirei Lower surface of leaves densely strigose or hispid ................................... 5 Upper and lower surfaces of leaves densely strigose or hispid; leaf blades 3–7.5 cm long; petioles 2–4 mm long; secondary leaf veins 6–10 on each side; corolla lobes 3.5–6.5 mm long, strigose-hispidulous adaxially .................................................................................................... S. foldatsii Upper surface of leaves glabrescent except the midrib strigillose, lower surface of leaves strigose; leaf blades 1.5–3 cm long; petioles 1–2 mm long; secondary leaf veins 3–5 each side; corolla lobes 8–9.5 mm long, glabrous adaxially ............................................................ S. pacimoniensis

Sipaneopsis foldatsii Steyerm., Mem. New York Bot. Gard. 17(1): 288. 1967. Subshrub 0.2–0.5 m tall; stems erect or spreading; stipules entire, triangular, ca. 4 mm long; leaves 3–7.5 × 0.8–3 cm; calyx lobes 1.5–2.5 mm long; corolla tube 13–16 mm

long, the lobes 3.5–6.5 × 3–4 mm; capsules 3– 3.5 × 3 mm. White-sand savannas, periodically flooded low, dwarf evergreen forests overlying white sand, 100–200 m; Amazonas (Caño Caname, Caño Yagua, Río Atacavi, Río Guayapo). Endemic. ◆Fig. 648.

834

R UBIACEAE

Fig. 648. Sipaneopsis foldatsii

Fig. 650. Sipaneopsis rupicola

Fig. 649. Sipaneopsis maguirei

Sphinctanthus 835

Sipaneopsis huberi Steyerm., Ernstia 23: 37, fig. 11. 1984. Subshrub 0.5–1.5 m tall; stems erect, sparsely branched; stipules 2–4 mm long, divided, with 2 or 3 lobes; leaves 2.5–6 × 1–1.5 cm; calyx lobes 1–1.5 mm long; corolla tube 9–11 mm long, the lobes 4–6 × 1.5–2.5 mm; capsules 2–3 × 2–2.5 mm. Granitic lajas and hilly savannas on sandstone, 200–500 m; Amazonas (Cerro Aracamuni, Cerro Vinilla, Río Siapa). Colombia (Caquetá).

Sipaneopsis pacimoniensis Steyerm., Mem. New York Bot. Gard. 17(1): 289. 1967. Subshrub 0.1–0.2 m tall; stems unbranched; stipules entire, triangular, 2.5–3 mm long; leaves 1.5–3 × 0.7–1.3 cm; calyx lobes 1.7–2 mm long; corolla tube ca. 17 mm long, the lobes 8–9.5 × 4.5–5 mm; capsules unknown. White-sand savannas bordering streams, 100–200 m; Amazonas (Río Pacimoni). Endemic.

Sipaneopsis maguirei Steyerm., Mem. New York Bot. Gard. 17(1): 287. 1967. Sipaneopsis wurdackii Steyerm., Mem. New York Bot. Gard. 17(1): 288. 1967. Subshrub 0.1–0.5 m tall; stems erect or prostrate; stipules entire, triangular, 2–3.5 mm long; leaves 1–7.5 × 0.4–2.8 cm; calyx lobes 2–3 mm long; corolla tube 15–22 mm long, the lobes 4.5–8 × 3–4.2 mm; capsules 3.5–4 × 3.5 mm. White-sand savannas subject to flooding, Río Negro caatinga, 50–200 m; Amazonas (Río Atabapo along affluent Río Caname, Río Autana, Río Cotúa, basin of Río Guainía, Río Guayapo, Río Orinoco along affluents Río Yagua, Río Ventuari). Adjacent Colombia. ◆Fig. 649.

Sipaneopsis rupicola (Spruce ex K. Schum.) Steyerm., Mem. New York Bot. Gard. 17(1): 285, fig. 29G–M. 1967. —Rondeletia rupicola Spruce ex K. Schum. in Mart., Fl. Bras. 6(6): 222. 1889. —Sipanea rupicola Spruce ex K. Schum. in Mart., Fl. Bras. 6(6): 222. 1889, nom. nud. pro syn. Rondeletia rupicola var. chiribiquetana R.E. Schult., Bot. Mus. Leafl. 14: 46. 1949. Subshrubs 0.3–2 m tall; stems erect, sparsely branched; stipules 3–5 mm long, divided and with 2 or 3 lobes or fimbriate and with 5–7 setae; leaves 3.8–6.5 × 0.8–1.5 cm; calyx lobes 1–1.5 mm long; corolla tube 8–8.5 mm long, the lobes 3–3.5 × 1.7–2 mm; immature capsules to 3 × 2.5 mm. Igneous outcrops along streams, borders of riparian forests, 100–200 m; Amazonas (Piedra Arauicaua, Piedra Tururumeri, basins of Río Guainía, Río Pacimoni, and Río Yatúa). Colombia (Vaupés: Cerro Chiribiquete). ◆Fig. 650.

Sipaneopsis morichensis Steyerm., Mem. New York Bot. Gard. 17(1): 286. 1967. Subshrub 1–2 m tall; stems erect, sparsely branched; stipules fimbriate, 8–10 mm long, with 3 or 4 setae; leaves 4.5–8.5 × 1.5–2.7 cm; calyx lobes ca. 2 mm long; corolla tube 6.7–11 mm long, the lobes 4.5–6 × 2–3.5 mm; capsules unknown. Savannas on tepui slopes, ca. 800 m; Amazonas (Cerro Moriche). Endemic.

81. SPHINCTANTHUS Benth., J. Bot. (Hooker) 3: 212. 1841. Conosiphon Poepp. in Endl., Gen. Pl. Suppl. 2: 54. 1842. by Charlotte M. Taylor and Julian A. Steyermark Shrubs or small trees, unarmed. Leaves opposite, petiolate, venation not lineolate; stipules interpetiolar and free or shortly united intrapetiolarly, persistent, ovate to triangular. Flowers medium to large, terminal, solitary or 2–4 and fasciculate, pedunculate, bracteate, bisexual, homostylous. Hypanthium cylindrical to narrowly turbinate. Calyx limb with well-developed tube, subtruncate or lobes 5 or 6, without calycophylls; corolla yellow, orange, green, or white and often with purple to brown spots on limb, salverform, externally densely sericeous to puberulous, tube sometimes constricted at throat, internally with a pilose ring (annulus of Steyermark) at or below the middle, lobes 5 or 6, convolute in bud. Stamens 5 or 6, inserted in upper part of corolla tube; anthers subsessile, narrowly oblong,

836

R UBIACEAE

Fig. 651. Sphinctanthus striiflorus

dorsifixed, included or situated in corolla throat. Ovary 2-locular; ovules numerous on axile placentas. Fruits baccate, subglobose to ovoid, smooth, thin-walled, yellow. Seeds flattened, smooth, subcircular to oblong. Colombia, Venezuela, Ecuador, Peru, Brazil; 5 or 6 species, 2 in Venezuela, 1 of these in the flora area. Sphinctanthus is similar to several other South America genera, and their delimitation is in need of reconsideration. With similar bisexual flowers and similar fruits are: Rosenbergiodendron, with the large white nocturnal flowers borne on short shoots, pollen in tetrads, and ovary placentation parietal; Tocoyena, with large yellow flowers borne in cymose groups and larger, thick-walled fruits; Genipa, trees with white to pale yellow flowers and thick-walled fruits; and Posoqueria, with large white nocturnal flowers with the corolla lobes imbricated in bud. Several other genera are similar to Sphinctanthus but can be separated by their unisexual flowers. Probably the most often confused with Sphinctanthus has been Ibetralia surinamensis Bremek. of Suriname, French Guiana, and northeastern Brazil; Ibetralia differs in its unisexual flowers with the staminate several and sessile and the pistillate pedunculate but solitary.

Stachyarrhena 837

Steyermark (1974) separated Sphinctanthus from Tocoyena based in part on stipule arrangement, which he described as shortly united intrapetiolarly in Tocoyena but free there, and thus interpetiolar, in Sphinctanthus. However Sphinctanthus actually varies in this regard, and does sometimes have stipules that are also partially fused intrapetiolarly (e.g., Steyermark 1974, p. 656, fig. 103, S. striiflorus; Vásquez & Jaramillo 9854, MO, S. maculatus Spruce ex K. Schum.). Sphinctanthus striiflorus (DC.) Hook. f. in Benth. & Hook. f., Gen. Pl. 2: 84. 1873. —Genipa (?) striiflora DC., Prodr. 4: 378. 1830. Sphinctanthus rupestris Benth., J. Bot. (Hooker) 3: 212. 1841. Conosiphon aureus Poepp. & Endl., Nov. Gen. Sp. Pl. 3: 27, pl. 233. 1845. Shrub or small tree to 3(–12) m tall; leaves 5.5–23 × 2–9 cm; stipules 1.5–5 mm long; ca-

lyx limb 3–5 mm long, shallowly lobed; corollas yellow to orange often with brown spots on limb, tube 10–20 mm long, lobes 10–12 mm long; fruit 2–3 cm diameter; seeds 5–6 mm long. Riparian forests, gallery forests, rocky places along streams, 100–500 m; Bolívar (middle Río Caura, Río Parhueña, Río Suapure), Amazonas (Río Manapiare, Río Parucito). Northeastern Peru, northern Brazil. ◆Fig. 651.

82. STACHYARRHENA Hook. f. in Hook., Icon. Pl. 11: 54, t. 1068. 1870. by Julian A. Steyermark and Claes Persson Trees of small to medium stature, glabrous, dioecious, unarmed, terrestrial. Leaves opposite, coriaceous, petiolate, venation brochidodromous, tertiary venation not lineolate; stipules chiefly connate, united around the stem into a tubular sheath to interpetiolar, persistent, mostly short-suborbicular, obtuse to truncate, rarely acute, in bud generally imbricated to valvate. Inflorescence terminal, bracteate, pedunculate, the staminate fasciculate, spicate or paniculate, the pistillate with flowers solitary. Flowers small, sessile to pedicellate, unisexual. Staminate flowers: hypanthium reduced; calyx limb cup-shaped, truncate to slightly repand-subundulate or obscurely 5-lobulate; corolla broadly infundibuliform or broadly campanulate, white or sometimes pink, glabrous externally, densely villous in upper portion internally, lobes 5, contorted in bud; stamens 5, inserted near the top of the corolla tube; anthers exserted, narrowly oblong, dorsifixed, sessile; pollen 3(4)-colporate; pistillode present. Pistillate flowers: hypanthium globose to ellipsoid; calyx and corolla similar to the staminate flowers or sometimes larger; ovary 4- or 5-locular; ovules numerous in each locule, placenta axial. Fruit baccate, globose or ellipsoid, glossy, sessile or long-pedicellate, pericarp thin but hard and woody, brown to black. Seeds usually horizontal, compressed; testa firm, of elongated cells with secondary thickenings in the radial walls. Panama, Colombia, Venezuela, Guyana, Ecuador, Peru, Brazil; 10 species, 4 in Venezuela, all in the flora area. Stachyarrhena is similar to Botryarrhena, from which it principally differs in being dioecious with solitary pistillate flowers (versus hermaphroditic with numerous flowers) and its 4- or 5-locular ovaries with numerous ovules (versus 2-locular with 2 or 4 ovules in each locule). Although Stachyarrhena is a fairly distinct genus, its species are in great need of revision. Most of the species are based on one or a few specimens. However, as more material has accumulated the delimitation of some species has become doubtful. A further complication is that most species descriptions are based on staminate specimens only. This is reflected in the key, which thus is of little help in identifying

838

R UBIACEAE

specimens with female flowers and fruits. However, pending a detailed study the traditional species circumscriptions are maintained here. Key to the Species of Stachyarrhena 1. 1. 2(1). 2. 3(1). 3.

Staminate inflorescence erect ................................................................... 2 Staminate inflorescence pendent .............................................................. 3 Leaves acuminate or subacuminate at the apex ............................. S. duckei Leaves obtuse or rounded at the apex ............................................ S. spicata Leaves obtuse to acuminate at the apex, with tertiary venation not reticulate and midrib terminating at the apex of the blade ........ S. penduliflora Leaves rounded or truncate at the apex, with tertiary venation prominently and loosely reticulate on both sides and midrib ending abruptly 6–8 mm below the apex of the blade ....................................... S. reticulata

Stachyarrhena duckei Standl., Field Mus. Nat. Hist., Bot. Ser. 22: 124. 1940. Small tree to 5 m tall; staminate corolla cream-colored or white, 5–6 mm long in bud. Seasonally flooded forests, riparian forests, among igneous boulders by streams, 100–300 m; Amazonas (Río Asisa, Río Cunucunuma, Río Negro). Amazonian Brazil. ◆Fig. 652. Stachyarrhena penduliflora K. Schum. in Mart., Fl. Bras. 6(6): 370. 1889. —Carutillo, Caruto. Shrub or small tree, 4–22 m tall; staminate corolla white or greenish white, 7.5–11 mm long. Rain forests, non-flooded gallery forests, Río Negro caatinga forests, 50–400 m; Bolívar (middle Río Orinoco at Río Pargueni and Río Horeda in Distrito Cedeño, Río Parguaza), Amazonas (Caño Yagua, Río Casiquiare, Río Cunucunuma, Río Manapiare, basin of Río Orinoco at Río Cataniapo, Río Ventuari). Amazonian Brazil.

50–300 m; Amazonas (Río Atabapo at Río Atacavi, Río Caname, Río Guiania, basin of Río Orinoco between Río Sipapo and Río Sanariapo, San Fernando de Atabapo). Endemic. Stachyarrhena spicata Hook. f. in Hook., Icon. Pl. 11: 54–55, t. 1068. 1870. —Schradera spicata Spruce ex Hook. f. in Hook., Icon Pl. 11: 45. 1870, nom. nud., pro syn. Stachyarrhena longifolia Hook. f. in Hook., Icon. Pl. 11: 55. 1870.

Stachyarrhena reticulata Steyerm., Mem. New York Bot. Gard. 12(3): 220, fig. 33, A–D. 1965. —Caruto, Pendare, Saranda. Shrub to 5 m tall; staminate corolla white, broadly campanulate, ca. 6.5 mm long. Riparian forests, flooded forests along streams,

Fig. 652. Stachyarrhena duckei

Tobagoa 839

Stachyarrhena spicata var. multinervis K. Schum. in Mart., Fl. Bras. 6(6): 370. 1889. Shrub or tree 2.5–8 m tall; staminate corolla white, broadly tubular, ca. 8 mm long.

Lowland and gallery forests, 100–300 m; Amazonas (basin of Río Casiquiare and Río Orinoco near Santa Bárbara del Orinoco). Apure, Guárico; Guyana (upper Río Mazaruni), Brazil (Amazonas: Rio Negro).

83. TOBAGOA Urb., Repert. Spec. Nov. Regni Veg. 14: 341. 1915. by Julian A. Steyermark and Charlotte M. Taylor Perennial, sprawling to prostrate herbs, terrestrial, unarmed, usually foulsmelling. Leaves opposite, subsessile to petiolate, venation not lineolate; stipules interpetiolar and united to petiole bases, persistent, truncate to broadly rounded, setose, in bud generally erect and valvate. Inflorescence axillary or sometimes at several nodes produced on alternating sides of stem, sessile, several-flowered, glomerulate, bracteate, the bracts sometimes setose. Flowers small, distylous, sessile or subsessile. Hypanthium turbinate. Calyx limb deeply lobed, lobes 4, without calycophylls; corolla rotate-funnelform, white, densely pubescent internally, lobes 4, valvate in bud. Stamens 4, inserted in corolla throat; anthers dorsifixed, oval-oblong, included. Ovary 2-locular; ovules solitary in each locule, axile. Fruit dry, indehiscent, ellipsoid, dicoccous but the cocci not separating. Seeds small, obovoid, ventrally excavated, finely reticulated. Panama, Antilles, northern Colombia, northern Venezuela, Tobago; 1 species. Tobagoa is similar to some species of Borreria and consequently often overlooked, although it is distinctive when alive due to its foul sulphurous odor. Because of this frequent confusion, Tobagoa is included in the key to the species of Borreria and similar genera. Tobagoa maleolens Urb., Repert. Spec. Nov. Regni Veg. 14: 343. 1916. —Peo. Diodia ottonis K. Schum. ex Pittier, Man. Pl. Usual. Venez. 338. 1926. Plant to 1 m tall, drying black; leaves 4–11 × 1–3 cm; secondary veins 5 or 6 pairs; petioles 1–10 mm long; stipules with sheath 0.5– 1 mm long, setae 3–5, 1.5–4 mm long; glomerules 3–5 mm diameter; calyx limb ca. 0.5 mm long; corolla with tube 1–1.5 mm long, lobes ca. 1 mm long; fruits oblong, 1.5–2 mm long. Abandoned coffee plantations, deciduous forests, 400–500 m; Bolívar (Cerro Tomasote). Distrito Federal, Miranda, Sucre; Antilles, Panama, Colombia, Tobago. ◆Fig. 653.

Fig. 653. Tobagoa maleolens

840

R UBIACEAE

84. TOCOYENA Aubl., Hist. Pl. Guiane 131, t. 50. 1775. by Charlotte M. Taylor and Julian A. Steyermark Trees or shrubs, terrestrial, unarmed, often turning black when dried. Leaves opposite or ternate, petiolate, venation not lineolate; stipules interpetiolar and often shortly intrapetiolar, triangular to ligulate, persistent, generally imbricate to valvate in bud. Inflorescence terminal, several- to multiflowered, cymose to corymbose, bracts reduced or absent, sessile to pedunculate. Flowers sessile to pedicellate, large, showy, fragrant, homostylous, protandrous. Hypanthium turbinate to ellipsoid or cylindrical. Calyx limb lobed, the lobes 5 or 6, without calycophylls; corolla salverform with a prolonged tube, white or yellow, internally glabrous except papillose to puberulent in throat, lobes 5 or 6, convolute in bud. Stamens 5 or 6, inserted in corolla throat; anthers dorsifixed, subsessile, narrowly oblong, partially exserted. Ovary 2-locular, ovules numerous in each locule, on axile placentas. Fruit baccate, ovoid to ellipsoid or subglobose, smooth, apparently black or brown, the pericarp coriaceous to woody. Seeds ellipsoid to ovoid, rather large, smooth, subcompressed. Central America, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay; 22 species, 6 in Venezuela, 5 of these in the flora area. Tocoyena is similar to Posoqueria and Sphinctanthus; their distinctions are discussed under each of these two latter genera. The root of Tocoyena is said to be used as a substitute for ipecac [Psychotria ipecacuanha (Brot.) Stokes]. Tocoyena has been studied by Luiza Kinoshita and Anajde Limes do Prado (personal communication; Universidade de Campinas, São Paulo), who have generously provided information on their view of this genus. Key to the Species of Tocoyena 1. 1. 2(1). 2. 3(2). 3.

4(2). 4. 5(4).

5.

Leaves rounded to broadly obtuse at apex; peduncles 20–30 mm long ................................................................................................. T. pendulina Leaves acute to acuminate at apex; inflorescences subsessile, peduncles to 5 mm long .......................................................................................... 2 Leaves glabrescent to sparsely appressed-pubescent abaxially .............. 3 Leaves pubescent abaxially with spreading trichomes ............................ 4 Leaves membranaceous, 17–36 × 8–13 cm, sparsely to densely pubescent; petioles 1–3.5 cm long; corolla pale yellow ............................ T. guianensis Leaves papyraceous to chartaceous, glabrous or sometimes shortly pubescent in the abaxial vein axils, 7.5–22 × 3.5–15 cm; petioles 0.5– 1.2 cm long; corolla white ...................................................... T. orinocensis Leaves 17–36 × 8–13 cm; corolla tubes 13–25 cm long ............. T. guianensis Leaves 5.5–13.5 × 2.5–8.5 cm; corolla tubes 6.5–10.2 cm long ................ 5 Lower surface of leaves with trichomes straight to somewhat crisped, moderately grouped, erect and not obscuring the surface; calyx teeth 0.3–1 mm long; Amazonas ........................................................ T. brevifolia Lower surface of leaves with trichomes crisped, densely grouped to intertwined, and usually obscuring the surface; calyx teeth 1–4 mm long; Bolívar ......................................................................................... T. neglecta

Tocoyena 841

Tocoyena brevifolia Steyerm., Mem. New York Bot. Gard. 12(3): 196. 1965. Shrub or small tree to 3(15) m tall; leaves 5.5–13.5 × 2.5–8 cm; secondary veins 4–10 pairs; petioles 2–10 mm long; stipules 2–4 mm long; inflorescences subsessile, subcapitate to congested-cymose; calyx limb with tube 1–1.5 mm long, lobes 0.3–1 mm long; corolla pale yellow, tube 8–10.2 cm long, lobes 20–26 mm long; fruits ca. 2.5 cm diameter; seeds 4–5 mm long. Igneous outcrops, low forests on igneous substrate bordering streams, 100–200 m; Amazonas (near Puerto Ayacucho, Raudal de Atures, Río Orinoco). Endemic. ◆Fig. 655. See comments under Tocoyena neglecta, below. As in that species, a larger range of measurements for leaves and corollas is given here than by Steyermark (1974). Tocoyena guianensis K. Schum. in Mart., Fl. Bras. 6(6): 346. 1889. Tocoyena guianensis var. communis Steyerm., Mem. New York Bot. Gard. 12(3): 194. 1965. Tocoyena guianensis var. glabriuscula Steyerm., Mem. New York Bot. Gard. 12(3): 195. 1965. Shrub or small tree to 3 m tall; leaves 17– 36 × 8–13 cm; secondary veins 10–14 pairs; petioles 1–3.5 cm long; stipules 3–8 mm long; inflorescences subsessile, congested-cymose; calyx limb ca. 4 mm long, lobed for ca. 1/2 its length; corolla pale yellow, tube 13–15 cm long, lobes 30–35 mm long; fruit ca. 3 cm diameter. Riparian forests and forests at bases of igneous topography, 50–300 m; Amazonas (Río Casiquiare basin, Río Manapiare, Río Negro at Piedra Cucuy, Río Pasimoni, Río Siapa). French Guiana, Brazil. ◆Fig. 654. Steyermark recognized three varieties of Tocoyena guianensis: the typical variety, known only from French Guiana and Brazil, with the leaves densely pubescent on both surfaces and corolla tubes 17–27 cm long; var. communis Steyerm. of French Guiana, Brazil, and Amazonas, Venezuela, with the leaves densely pubescent abaxially and corolla tubes 17–27 cm long; and var. glabriuscula Steyerm. of Amazonas, Venezuela, with the leaves only sparsely pubescent and corolla tubes ca. 15 cm long. However,

recent collections have made separation of these varieties based on these characteristics problematic. In particular Liesner 4057 (MO) has leaves densely pubescent on both surfaces as in var. guianensis but corolla tubes 15 cm long as in var. glabriuscula. Thus either there is clinal variation in corolla length or more than one species is involved here, as suggested by Steyermark (1965). In either case his varieties cannot be sustained as he delimited them, and they are not recognized here. Tocoyena neglecta N.E. Brown, Trans. Linn. Soc., Bot. 6: 35. 1901. Shrub to 4 m tall; leaves 7–13 × 3–8.5 cm; secondary veins 6–8 pairs; petioles 3–5 mm long; stipules 3–4 mm long; inflorescence subsessile, congested-fasciculate to shortly cymose; calyx limb with tube ca. 2 mm long and lobes 1–4 mm long; corolla white, tube 6.5–9 cm long, lobes 1.5–2.8(–3.8) cm long; fruit subglobose, 2.5–3 cm diameter, glabrescent; seeds ca. 5 mm long. Open savannas, gallery forests, igneous outcrops (lajas), rocky margins of cascades and stream borders, 200–500 m; Bolívar (basins of Río Caroní, middle Río Orinoco, and Río Paragua). Guyana. Tocoyena neglecta is similar to T. brevifolia, and in general replaces it in the eastern part of the flora area. The measurements given here, in particular for leaf size and corolla tube length, include a wider range than those presented by Steyermark (1965; 1974) and are based on more collections than he examined. Tocoyena orinocensis Standl. & Steyerm., Fieldiana, Bot. 28: 617. 1953. Shrub to 3 m tall; leaves 7.5–22 × 3.5–15 cm; secondary veins 8–11 pairs; petioles 5–12 mm long; stipules 3–6 mm long; inflorescences subsessile, congested-cymose; calyx limb ca. 2 mm long, lobed for ca. 1/2 its length; corolla white, tube 8–15 mm long, lobes 20– 30 mm long; fruits subglobose, ca. 2 cm diameter; seeds not seen. Igneous outcrops, wet savannas, 50–200 m; Bolívar (Agua Amena near Caicara), Amazonas (east of Isla Ratón, near Puerto Ayacucho). Eastern Colombia. ◆Fig. 656. The relationship between Tocoyena orino-

842

R UBIACEAE

Fig. 654. Tocoyena guianensis

censis and T. brevifolia probably deserves reexamination. These species differ primarily in their densely pubescent (T. brevifolia) versus glabrous (T. orinocensis) vegetative and reproductive structures. Tocoyena orinocensis is circumscribed somewhat differently here than by Steyermark (1965); in particular a number of specimens called T. guian-

ensis by him are here included in T. orinocensis, extending the range of corolla tube length in T. orinocensis. Tocoyena pendulina Spruce ex Standl., Publ. Field Columbian Mus., Bot. Ser. 7: 391. 1931. —Guarisicuarejue (Curripaco).

Tocoyena 843

Fig. 655. Tocoyena brevifolia

Fig. 656. Tocoyena orinocensis

Tree to 15 m tall; leaves 10–17 × 5.5–8.5 cm; secondary veins 6–8 pairs; petioles 5–15 mm long; stipules 4–6 mm long; inflorescence cymose, with peduncle 2.5–3 cm long; calyx limb with tube 3–4 mm long, lobes ca. 0.5 mm long; corolla white, tube 4–4.5 cm long, lobes 15–17 mm long; fruits ellipsoid, 4–5 × 2.5–3.5

cm; seeds not seen. Savannas, flooded margins of riparian forests, 100–200 m; Amazonas (Río Guainía basin). Endemic. Tocoyena pendulina is the only species of Tocoyena in the flora area with rounded corolla lobes. It may eventually be found in adjacent Colombia.

844

R UBIACEAE

85. UNCARIA Schreb., Gen. Pl. 1: 125. 1789, nom. cons. Ourouparia Aubl., Hist. Pl. Guiane 177, t. 68. 1775, nom. rejic. by Charlotte M. Taylor and Julian A. Steyermark Woody shrubs or lianas, terrestrial, climbing to the canopy by recurved axillary spines. Leaves opposite, petiolate, venation not lineolate; stipules interpetiolar, ligulate to bilobed, in bud erect and flatly appressed, persistent to caducous. Inflorescence axillary or terminal, multiflowered, capitate, the heads pedunculate, solitary or several in a cyme, ebracteate. Flowers rather small, sessile or pedicellate, homostylous, protandrous. Hypanthium fusiform to turbinate. Calyx limb lobed, lobes 5, without calycophylls; corolla narrowly funnelform, greenish white, creamcolored, pale yellow, or sometimes orange, internally glabrous, lobes 5, valvate in bud. Stamens 5, inserted in the corolla throat; anthers dorsifixed, oblong or linear, partially exserted. Ovary 2-locular; ovules numerous in each locule, axile. Fruit capsular, fusiform, septicidal, chartaceous, smooth. Seeds small, flattened, fusiform, marginally winged. Central America, Colombia, Venezuela, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia, Africa, Asia; 34 species, 2 in Venezuela, both in the flora area. The spines are formed by modified peduncles and infrequently bear inflorescences. Neotropical Uncaria is thought to have anti-cancer properties; though this has not been tested scientifically it is a popular herbal remedy. Some Old World species are important sources of chemicals for tanning and dyeing. Key to the Species of Uncaria 1. 1.

Lower surface of leaves and petioles glabrous; flowers and fruits pedicellate .......................................................................................... U. guianensis Lower surface of leaves, including the midvein and petioles, puberulent or tomentose; flowers and fruits sessile ................................. U. tomentosa

Uncaria guianensis (Aubl.) J.F. Gmel., Syst. Nat. 2: 370. 1791. —Ourouparia guianensis Aubl., Hist. Pl. Guiane 177, t. 68. 1775. —Racimo de cambur, Uña de gavilán. Liana; spines 1.5–2 cm long; leaves 7–13.5 × 2.5–7 cm; stipules 5–15 mm long; heads of inflorescence 1–9, each 2–3 cm diameter; pedicels 1–3 mm long; calyx limb 2–3 mm long; corollas with tubes 4.5–7 mm long, lobes 1.5–3 mm long; capsules 15–25 × 4–7 mm, with pedicels to 5 mm long; seeds 6.5– 8.5 mm long. Riparian forests, semideciduous to evergreen lowland and lower montane forests, savannas, 50–500 m; Delta Amacuro (widespread), Bolívar (common in basin of Río Botanamo, Río Caroní, Río Caura, and Río Paragua), Amazonas (Río Siapa). Distrito Federal, Zulia; Colombia, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 657.

Uncaria tomentosa (Willd. ex Roem. & Schult.) DC., Prodr. 4: 349. 1830. —Nauclea tomentosa Willd. ex Roem. & Schult., Syst. Veg. 5: 221. 1819. —Ourouparia tomentosa (Willd. ex Roem. & Schult.) K. Schum. in Mart., Fl. Bras. 6(6): 132. 1889. Liana; spines 1–2 cm long; leaves 6–15 × 2.5–9 cm; stipules 1–1.5 cm long; heads of inflorescence 3–5, each 1.2–2.5 cm diameter; calyx limb 0.5–1 mm long; corollas with tubes 3.5–6 mm long, lobes 1–1.5 mm long; capsules 7–15 × 4 mm; seeds 2–3 mm long. Semideciduous to evergreen lowland forests, near sea level to 300 m; Delta Amacuro (Pedernales), Amazonas (basin of Río Guainía at base of Sierra de la Neblina). Anzoátegui, Apure, Barinas, perhaps Zulia; Central America, Colombia, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 658.

Uncaria 845

Fig. 657. Uncaria guianensis

Fig. 658. Uncaria tomentosa

846

R UBIACEAE

86. WARSZEWICZIA Klotzsch, Monatsber. Königl. Preuss. Akad. Wiss. Berlin 496. 1853. by Charlotte M. Taylor and Julian A. Steyermark Trees or shrubs. Leaves opposite, petiolate, venation not lineolate; stipules interpetiolar, triangular, erect, caducous. Inflorescence terminal and sometimes in the uppermost leaf axils, paniculate to racemiform, the flowers in small cymules borne along spiciform axes, bracteate though bracts often reduced. Flowers rather small, homostylous, protogynous, usually subsessile. Hypanthium turbinate to ellipsoid. Calyx limb sinuate to 5-lobed, the lobes equal or in some flowers of some plants (usually the terminal flower of the cyme) one of them expanded into a petaloid, cream to pink or red calycophyll; corolla white to orange, campanulate to funnelform, internally barbate in throat, lobes 5, imbricate in bud. Stamens 5, inserted in the corolla throat; anthers dorsifixed, exserted. Ovary 2-locular; ovules numerous, on axile placentas. Fruits capsular, ellipsoid to obconic, septicidal, woody, smooth, with calycophylls generally persistent. Seeds small, angled to flattened, reticulate. Mexico, Central America, South America south to Bolivia; 2–6 species, 3 in Venezuela, all in the flora area. The expanded, petaloid calyx lobes are often mistaken for bracts. Key to the Species of Warszewiczia 1.

1.

2(1). 2.

Secondary leaf veins in 15–40 pairs; inflorescences usually unbranched, with the flower cymes borne directly from the primary axis; petaloid calyx lobes yellow or orange to usually deep red; corolla yellow or orange-yellow, 6.5–10 mm long ................................................. W. coccinea Secondary leaf veins in 10–15 pairs; inflorescences usually branched to several orders; petaloid calyx lobes white or greenish white; corolla white, 1–1.5 mm long ............................................................................ 2 Leaves elliptic to ovate, 4.5–11 cm wide; corolla 1.2–1.5 mm long; capsules 1.2–2 mm long ....................................................................... W. elata Leaves elliptic to ovate or elliptic-oblong, 12–20 cm wide; corolla ca. 2 mm long; capsules ca. 2 mm long .................................................. W. schwackei

Warszewiczia coccinea (Vahl) Klotzsch, Monatsber. Königl. Preuss. Akad. Wiss. Berlin 497. 1853. —Macrocnemum coccineum Vahl, Symb. Bot. 2: 38. 1791. —Barba gallo, Clavellino, Papagayo, Pescuezo de pavo, Rabo de guaca. Warszewiczia schomburgkiana Klotsch, Monatsber. Königl. Preuss. Akad. Wiss. Berlin 497. 1853. Tree or shrub to 12 m tall; leaves 15–50 × 5–18 cm; secondary veins 15–40 pairs; petioles 5–25 mm long; stipules 10–25 mm long; inflorescences racemiform, 30–60 × 1.5–5 cm; calyx limb 1.5–2 mm long, calycophylls yellow or orange to deep red, 9–15 × 2–4 cm; corolla yellow or orange-yellow, tube 4–6 mm long, lobes 2.5–4 mm long; capsules 4–5 × 3–

4 mm; seeds 0.2–0.5 mm long. Semideciduous to evergreen lowland forests, forest edges, 100–200 m; Amazonas (basins of Río Casiquiare and Río Orinoco from Río Sipapo and Río Cuao to Caño Yapacana). Anzoátegui, Barinas, Mérida, Monagas, Sucre, Táchira, Trujillo, Zulia; Mexico, Central America, Colombia, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia. ◆Fig. 659. Warszewiczia coccinea is striking in flower and is commonly collected. It is the national tree of Trinidad and Tobago. This widespread tree apparently occupies a variety of habitats. The name Warszewiczia cordata Spruce ex K. Schum. has been applied to trees from the western and southwestern Amazon basin that have truncate to cordate leaf bases (ver-

Warszewiczia 847

Fig. 659. Warszewiczia coccinea

sus acute to cuneate in W. coccinea), and appear to be restricted to seasonally flooded river banks. Warszewiczia elata Ducke, Arch. Jard. Bot. Rio Janeiro 3: 254. 1922. Tree to 20 m tall; leaves 10–20 × 4.5–11 cm; secondary veins 10–13 pairs; petioles 5– 20 mm long; stipules 5–20 mm long; inflorescences 10–20 × 8–15 cm; calyx limb ca. 0.8 mm long, calycophylls white, 3.5–6 × 0.8–2 cm; corolla white to pale green, 1.2–1.5 mm long, lobes minute; capsules 1.2–2 × 1–2 mm. Evergreen lowland forests, 50–100 m; Amazonas (basins of Río Orinoco, Río Cataniapo, and Río Coromoto south of Puerto Ayacucho). Peru, Brazil.

Warszewiczia schwackei K. Schum. in Mart., Fl. Bras. 6(6): 219, t. 115. 1889. Tree to 16 m tall; leaves 20–32 × 12–20 cm; secondary veins 12–18 pairs; petioles 3– 4.5 cm long; stipules 12–15 mm long; inflorescences 18–30 × 18–25 cm; calyx limb ca. 0.5 mm long, calycophylls white, 3–10 × 1–3 cm; corolla white to pale green, tube ca. 1.5 mm long, lobes ca. 0.5 mm long; capsules ca. 2 × 1.5 mm. Evergreen lowland forests, 100– 300 m; Amazonas (Río Cunucunuma northeast of Cerro Huachamacari, Río Mawarinuma at base of Sierra de la Neblina). Brazil. The distinctions between Warszewiczia schwackei and W. elata are subtle, and these species need further consideration. The leaves of W. schwackei are usually densely pilosulous abaxially, though they are glabrescent in some specimens (e.g., A. Fernández 7522, MO); the leaves of W. elata are usually glabrous abaxially. Also, the inflorescences of W. schwackei usually have several well-developed calycophylls, while those of W. elata usually have no or only 1 or 2 calycophylls.

Appendix List of new names and emendations published in this volume

Alibertia edulis var. obtusiuscula (Steyerm.) Delprete & C. Perss., comb. nov. .......................................................... Cordiera myrciifolia (Spruce ex K. Schum.) C. Perss. & Delprete, comb. nov. .......................................................... Notopleura sandwithiana (Steyerm.) C.M. Taylor, comb. nov. . . . . . . . . . . . . . . Remijia aracamuniensis (Steyerm.) C.M. Taylor, comb. nov. . . . . . . . . . . . . . . . Remijia globosa (Steyerm.) C.M. Taylor, comb. nov. . . . . . . . . . . . . . . . . . . . . . . Retiniphyllum concolor (Spruce ex Benth.) Müll. Arg., emend. Cortés . . . . . . . . Rudgea hostmanniana subsp. maypurensis (Standl.) Zappi, comb. & stat. nov. .......................................................... Rudgea sclerocalyx (Müll. Arg.) Zappi, comb. nov. . . . . . . . . . . . . . . . . . . . . . . . .

848

514 559 663 781 782 794 808 810

Index Compiled by George Thornburgh

Entries in Roman type = accepted names of taxa and vernacular names Entries in italics = synonyms Page numbers in bold = illustrations

—A— Acicarpa sacchariflora, 85 Acosta aculeata, 324 Acroceras, 15 excavatum, 15 zizanioides, 15, 16 Adájate, 339 Adenostephanus, 385 Aegopogon, 17 cenchroides, 16, 17 Agrosticula, 273 muralis, 274 Agrostis brasiliensis, 251 compressa, 51 minutiflora, 274 radiata, 72 Aguacatillo, 704 Aira laxa, 174 Ají de agojó, 401 Ají de gallineta, 810 Ají de gato, 663 Ají de morocoto, 365 Ají de morrocoy, 610, 794 Ají de paloma, 671 Ají de paloma, 670 Alatrique peludo, 735

Alatriquillo, 743 Aledodö, 355 Alerón, 352 Alibertia, 512 acuminata, 513 var. obtusiuscula, 514 benensis, 560 bertierifolia, 513, 514 davidsei, 514 edulis, 513 var. edulis, 513 var. obtusiuscula, 514 granulosa, 514 hexagyna, 513 latifolia, 514 var. pargueniana, 514 var. parvifolia, 514 longistipulata, 513 myrciifolia, 559 nitidula, 514 panamensis, 513 steinbachii, 560 stenantha, 513 tenuifolia, 560 tobagensis, 514 triflora, 560 triloba, 559 trinitatis, 514 tutumilla, 513 uniflora, 559 utilis, 513 Alseis, 515

849

labatioides, 515, 516 leiantha, 516 trichocarpa, 516 Amaioua, 516 corymbosa, 517, 517 fusifera, 581 genipoides, 581 guianensis, 517 var. guianensis, 518 var. macrantha, 518 peruviana, 517 saccifera, 583 velutina, 587 Amarilla, 544 Ameu, 802 Ammianthus, 792 Ampelozizyphus, 475 amazonicus, 474, 475 Amphidasya, 518 neblinae, 518, 519 Amphiphyllum, 416 rigidum, 416, 417 schomburgkii, 429 Anagallis, 383 pumila, 384, 384 Anastrophus, 45 capillaris, 50 platycaulis, 51 Anatherum holcoides, 106 Andriapetalum, 386 rubescens, 389

850

I NDEX

[Andriapetalum] sessilifolium, 389 Andropogon, 17 adustus, 94 angustatus, 19, 21 angustifolius, 288 barbatus, 72 bicornis, 19 var. bicornis, 19 var. burchellii, 19 bracteatus, 121 brevifolius, 262 canescens, 288 carinatus, 19 condensatus, 263 contortus, 117 crassus, 20 crucianus, 20 dactyloides, 289 dissolutus, 289 diuturnus, 19, 22 fasciculatum, 72 fastigiatus, 19, 21 hirtiflorus, 265 hispidus, 40 hypogynus, 19, 23 indetonsus, 20 insolitus, 20 insularis, 85 latifolius, 137 leucostachyus, 20, 22 subsp. selloanus, 20 ligularis, 289 longiramosus, 20 macroglossus, 289 microstachyum, 263 mollis, 288 montufarii, 288 perdignus, 26 plumosus, 288 polydactylon, 72 riedelii, 265 secundus, 289 segetum, 137 selloanus, 20 semiberbis, 265 setosus, 269 sorghum, 272 spicatus, 288 stipoides, 289 tener, 265 trichospirus, 291 truncatus, 289

vestitus, 289 vetus, 20, 24 virgatus, 25, 26 Anisomeris, 547 caurensis, 548 malaneoides, 548 polyantha, 549 tenuiflora, 549 volubilis, 550 Anoe, 514 Anoeyo, 514 Anthaenantia, 147 lanata, 147 Anthephora, 26 hermaphrodita, 26, 27 Antigonon, 348 leptopus, 349, 349 Antirhea surinamensis, 615 Aphanocarpus, 519 steyermarkii, 520, 520 f. elongatus, 520 f. glabrior, 520 Apinagia, 303 brevicaulis, 304, 306 corymbosa, 304 var. corymbosa, 304 exilis, 304, 306 guyanensis, 304 kochii, 304 longifolia, 305, 306 multibranchiata, 305 perpusilla, 311 richardiana, 305, 306 ruppioides, 305, 306 staheliana, 305 Apodanthes, 409 caseariae, 410, 410 flacourtiae, 410 roraimae, 410 Aporuh-yaré, 542 Appunia, 656 angulata, 657 tenuiflora, 657 var. leiophylla, 657 venezuelensis, 659 Arabica coffee, 557 Aragueque, 353 Arahueque, 353 Arahuequito, 354 Arajueque montañero, 355 Araña, 665 Arberella, 28

aff. bahiensis, 28 venezuelae, 28, 29 Arisi, 176 Aristida, 28 adscensionis, 30 var. bromoides, 30 subsp. bromoides, 30 var. coarctata, 30 americana, 63 arenaria, 35 bromoides, 30 capillacea, 31, 32 coarctata, 30 dispersa var. humilis, 31 doelliana, 35 elegans, 31 gibbosa, 31 humilis, 31 implexa var. aequa, 31 longifolia, 31, 34 marginalis, 31 moritzii, 31, 33 neesiana, 31 orizabensis, 31 pittieri, 31 planifolia, 31 recurvata, 31, 33 riedeliana, 31 riparia, 31, 32 scabra, 35 setifolia, 31, 33 var. arenaria, 35 var. grandiflora, 35 var. intermedia, 35 tarapotana, 35 tenuis, 35 ternipes, 35 tincta, 35 torta, 32, 35 Aristopsis, 28 Arizo, 355 Arroz, 176 Arthratherum, 28 Arthrostylidium, 35 cacuminis, 36 pubescens, 36 racemiflorum, 252 scandens, 36 schomburgkii, 36, 37 steyermarkii, 42 subpectinatum, 45

I NDEX

venezuelae, 38 sp. A, 38 sp. B, 38 sp. C, 38 sp. D, 38 Arundinaria deflexa, 42 schomburgkii, 38 Arundinella, 38 confinis, 40 hispida, 39, 40 Arundo altissima, 245 australis, 245 phragmites, 245 roraimensis, 78 Aspidanthera, 600 Atractantha, 40 amazonica, 40, 41 Aulonemia, 40 chimantaensis, 42, 43 deflexa, 42, 44 jauaensis, 42, 43 steyermarkii, 42 aff. subpectinata, 45 sp. A, 45 sp. B, 45 sp. C, 45 Axonopus, 45 affinis, 52 sect. Anastrophus, 45 anceps, 48, 56 anomalus, 53 appendiculatus, 51 arundinaceus, 49 ater, 52 aturensis, 49 aureus, 49, 57 var. pilosus, 50 sect. Cabrera, 45 canescens, 49 var. psilachne, 49 capillaris, 49, 58 caracarahyensis, 49 carinato-vaginatus, 49 var. sprucei, 49 casiquiarensis, 50 var. casiquiarensis, 50 var. A, 50 caulescens, 50, 59 chimantensis, 50 chrysites, 49 chrysoblepharis, 50

chrysodactylus, 49 compressus, 50, 60 var. affinis, 52 var. australis, 51 var. macropodius, 51 cuatrecasasii, 51 eminens, 51 equitans, 51 erectus, 49 exasperatus, 49 excavatus, 51, 57 extenuatus, 50 fissifolius, 52, 60 var. coronatus, 52 flabelliformis, 52, 59 var. camporum, 52 var. decipiens, 52 flexilis, 53 fockei, 54 gentilis, 51 gracilis, 52 hirsutus, 52 hitchcockii, 53 immersus, 50 iridifolius, 52 kaietukensis, 52 laxus, 50 leptostachyus, 52, 61 longispicus, 53 macrostachyus, 53 magallanesiae, 53 maguirei, 52 minutus, 49 multipes, 51 paranaensis, 53 paucisetosus, 49 pennellii, 53 piccae, 52 pruinosus, 48 pulcher, 49 purpurellus, 52 purpusii, 53 ramosus, 53 rivularis, 53 schultesii, 53, 61 sprucei, 49 var. glabratus, 49 steyermarkii, 54, 62 stragulus, 52 suffultiformis, 54 var. suffultiformis, 54, 62 var. A, 54

851

surinamensis, 54 tamayonis, 52 triglochinoides, 54, 58 villosus, 54 yutajensis, 54 sp. A, 54 sp. B, 55 sp. C, 55 —B— Babandi, 545 Bada, 783 Bambusa guadua, 111 latifolia, 111 venezuelae, 111 Bambusillo, 111 Baquerito, 354 Barba de tigre, 779 Barba gallo, 846 Barka-makata-purai, 582 Barnhartia, 317 floribunda, 318, 318 Basanacantha, 776 dioica, 778 hebecarpa, 778 spinosa, 777 f. macrophylla, 777 Bathysa, 521 bathysoides, 521, 521 Bejuco carena, 319 Bejuco colorado, 319 Bejuco cuadrado, 339 Bejuco cuarenta días, 340, 341 Bejuco culebrón, 340 Bejuco de bocachico, 319 Bejuco de curare, 320 Bejuco de jabón, 341 Bejuco jabón, 339, 340, 341, 347 Bejuco negro, 339 Bejuco parásito, 326 Bejuco puguacari, 319 Bejuco pujajui, 475 Bejuco reuma, 478 Bejuco reumo, 478 Bejuco salvia, 819 Bellardia repens, 555 tontanea, 555 Bellisima, 349

852

I NDEX

Bellynkxia, 656 angulata, 657 Bernardino, 475 Bertiera, 522 diversiramea, 522 guianensis, 523 palustris, 831 parviflora, 523, 524 Billardiera paniculata, 565 Biyw mamo ka hi, 369 Blochmannia weigeltiana, 370 Boca de sapo, 619 Bonita de noche, 568 Boquilla, 747 Bora, 372 Borojoa, 512 lanceolata, 513 Borrajón, 795 Borreria, 524 alata, 528, 534 assurgens, 532 bolivarensis, 528, 535 capitata, 528, 529 var. capitata, 529 f. ferruginea, 529 f. glabra, 529 var. tenella, 529, 535 cataractarum, 529, 535 confertifolia, 529 densiflora, 530 exilis, 532 ferruginea, 529 fockeana, 530 hispida, 530, 536 var. glabrescens, 530 hyssopifolia, 575 intricata, 530 jangouxii, 530, 534 kappleriana, 529 latifolia, 530, 536 var. fockeana, 530 var. latifolia f. fockeana, 530 f. minor, 531 var. minor, 531 macrocephala, 531, 534 multiflora, 576 ocymifolia, 576 oligodonta, 533 prostrata, 531 pygmaea, 531, 536

remota, 532 repens, 532, 536 spicata, 576 tenella, 529 var. platyphylla, 529 umbellata, 589 verticillata, 533, 534 wurdackii, 533, 535 Botón de soldado, 379 Botryarrhena, 537 pendula, 538 venezuelensis, 537, 538 Bouteloua, 63 americana, 63, 63 Boyuyo, 742 Brachiaria, 64 arrecta, 64 decumbens, 64 fasciculata, 65, 66 humidicola, 67 mollis, 67 mutica, 65, 67 plantaginea, 66, 67 Bredemeyera, 318 altissima, 319 densiflora, 319 var. glabra, 319 floribunda, 319, 320 lucida, 319 myrtifolia, 319, 320 parviflora, 320 sp. A, 320 Briza, 67 minor, 68, 68 Bromus spicatus, 293 Brusquillo blanco, 354 Buchia plantaginea, 697 Buena, 562 Bujiyuju, 763 Buju, 489 —C— Cabrera, 45 chrysoblepharis, 50 Cabrito negro, 568 Cachete de vieja, 618 Cachicamo, 565 Cachira-huaro, 671 Cachito, 779 Cacho de bagre, 615

Cacho de venado, 517, 518 Cadajo-como-mamu-jei-du, 763 Café, 557, 565 Café criollo, 557 Café montañero, 704 Café negro, 625 Café orillera, 565 Café robusta, 557 Cafecillo, 594, 595, 596, 615, 657, 704, 722, 745, 763, 807, 808, 810, 827 Cafecillo de danta, 594 Cafecito, 593, 594, 778 Caicará, 124 Callicocca, 708 alba, 760 purpurea, 761 tomentosa, 753 Calycophyllum, 538 obovatum, 539, 539 venezuelense, 540 Camaguari, 532 Camilla de locho, 389 Campderia, 349 Camuco-anajoru, 363 Caña brava, 113 Canalete, 517 Canaotera, 121 Canilla de tintín, 330 Canilla de venado, 355, 514, 518 Capa rosa, 690 Capim melas, 151 Capirona, 540 decorticans, 540, 541 leiophloea, 540 wurdackii, 540 Capriola, 78 dactylon, 78 Carapichea, 708 kappleri, 743 Caricillo, 168 Carillo de perro, 818 Carinta, 608 Carisillo, 627, 756 Cari-yarena, 106 Carne asada, 391 Carricillo, 137 Carrisillo, 596, 721 Carriso, 688

I NDEX

Carriso de picure, 688, 762 Carriso morado, 721 Carriso sortijo, 688 Carrizo, 111, 168, 684 Carrizo de picure, 684, 721, 724 Carrutillo morichalero, 625 Cartancillo, 475 Cartera, 618 Carurú, 304 Carutilla, 513, 514 Carutillo, 516, 518, 544, 581, 582, 785, 838 Carutillo blanco, 522 Carutillo rebalsero, 514, 581, 582 Caruto, 607, 627, 628, 753, 782, 838 Caruto montañero, 607 Caryochloa bahiensis, 149 Cascarilla roraimae, 784 Cascarón, 488 Casco de mula, 808 Cashék, 341 Caspadillo, 400 Cassipourea, 486 guianensis, 485, 486 Cassupa, 624 verrucosa, 628 Catocoma, 318 altissima, 319 floribunda, 319 lucida, 319 Cazabe chiquito, 514 Cedui-chu, 124 Celastrus myrtifolius, 495 Cenchrus, 68 brownii, 69, 71 ciliaris, 70, 71 echinatus, 69, 71 granularis, 115 inflexus, 92 parviflorus, 266 setosus, 240 Centunculus pentandrus, 384 pumilus, 384 Cephaelis, 708 alba, 760

amoena, 733 axillaris, 724, 802 barcellana, 754 blepharophylla, 747 bolivarensis, 726 bracteocardia, 728 colorata, 733 dichotoma, 740, 752 ernestii, 759 fanshawei, 738 hemicephaelis, 740 hirsuta, 753 hoffmannseggiana, 740 humboldtiana, 741 var. caudata, 741 var. ornata, 741 kappleri, 743 microcephala, 740, 754 oblonga, 748 paraensis, 761 prunifolia, 754 pubescens, 728 purpurea, 761 rosea, 755 rubra, 740 sandwithiana, 663 sororiella, 760 sphaerocephala, 757 stipulosa, 758 surinamensis, 761 tatei, 734 tepuiensis, 759 tomentosa, 753 violacea, 724 Cephalodendron, 779 aracamuniensis, 781 globosum, 782 Cephalostemon, 416 affinis, 418, 419 cyperaceoides, 421 flavus, 421 junciformis, 421 microglochin, 419 squarrosus, 421 Cereza, 808 Chaetaria, 28 adscensionis, 30 bromoides, 30 capillacea, 31 coarctata, 30 gibbosa, 31 humilis, 31 recurvata, 31

853

setifolia, 35 torta, 35 Chalepophyllum, 541 coriaceum, 643 guianense, 542, 542 var. cuneatum, 542 latifolium, 644 longilobum, 542 speciosum, 642 tatei, 643 Chaparrillo rebalsero, 625 Chaparro de agua, 368, 389 Chaparro de sabana, 700 Chaparro rebalsero, 355 Chimarrhis, 542 bathysoides, 521 brevipes, 543, 543 longistipulata, 544 microcarpa, 544 subg. Pseudochimarrhis, 542 Chinák, 319 Chinay-yek, 353 Chino, 818 Chiococca, 544 alba, 545 subsp. parvifolia, 545 var micrantha, 545 f. pilosa, 545 auyantepuiensis, 545 brachiata, 545 erubescens, 546 lucens, 545 micrantha, 545 nitida, 545 var. amazonica, 546 var. chimantensis, 546 var. nitida, 546, 546 parvifolia, 545 pubescens, 546 racemosa, 545 Chi-ví-du, 733 Chiyaya, 530 Chloris, 71 barbata, 72, 72 dandyana, 72 elata, 72 foliosa, 112 inflata, 72 mollis, 97 polydactyla, 72 radiata, 72

854

I NDEX

Chomelia, 547 barbellata, 550 caurensis, 548 delascioi, 548 glabricalyx, 548 malaneoides, 548, 550 monachinoi, 549 polyantha, 549 stergiosii, 549 tenuiflora, 549 volubilis, 550 Chondrococcus, 552 laevis, 552 Choori yo’, 530 Chusquea, 73 linearis, 73, 74 venezuelae, 38 sp. A, 73 Chuta, 763 Chytropsia, 708 astrellantha, 724 Cinchona amazonica, 552 brasiliensis, 640 firmula, 782 grandiflora, 562 lambertiana, 636 magnifolia, 636 oblongifolia, 636 pedunculata, 784 roraimae, 784 Cinchonopsis, 551 amazonica, 551, 552 Ciruela de playa, 657 Ciutica, 629 Clavellino, 846 Clematis, 411 caracasana, 412 caripensis, 412 floribunda, 413 goudotiana, 413 guadeloupae, 411 var. guadeloupae, 412, 412 populifolia, 413 Coccochondra, 552 laevis, 552 subsp. laevis, 553, 553 subsp. maigualidae, 553 Coccocypselum, 553 aureum, 554, 556 var. capitatum, 554 brevipetiolatum, 556

canescens, 556 condalia, 555 croatii, 555 guianense, 555, 556 var. patens, 555 hirsutum, 555 huberi, 556 lanceolatum, 556 tontanea, 555 Coccoloba, 349 acuminata, 352 ascendens, 352, 357 bolivarana, 354 bracteolosa, 353 caracasana, 352 caribaea, 353 caurana, 354 charitostachya, 353 coronata, 353 cyclophylla, 352 declinata, 353, 356 densifrons, 353 dugandiana, 353, 358 excelsa, 353, 359 fallax, 353, 360 grandis, 354 guianensis, 354 gymnorrhachis, 354 latifolia, 354 llewelynii, 354, 360 lucidula, 354 marginata, 354, 359 micropunctata, 353 mollis, 354 novogranatensis, 353 obtusifolia, 354, 361 ochreolata, 355 orinocana, 355, 357 ovata, 355 paraensis, 353 parimensis, 353 var. schomburgkii, 353 pittieri, 355 polystachya, 354 sagotii, 354 schomburgkii, 355, 361 spruceana, 355, 362 striata, 355, 356 trinitatis, 354 uvifera, 355 wurdackii, 355 sp. A, 355 sp. B, 356

sp. C, 356 sp. D, 356 Coccolobis, 349 Coco de mono, 397 Coco de mono pequeño, 397 Coelorachis, 73 aurita, 75, 76 loricata, 256 Coffea, 556 arabica, 557, 558 crassiloba, 808 laurifolia, 810 liberica, 557 sessilis, 762 stipulacea, 810 subsessilis, 802 tenuiflora, 657 Coix, 76 lacryma-jobi, 76, 77 Cola de pava, 401 Cola de pava chiquita, 401 Cola de pava negra, 401 Coloradito, 401, 723 Colubrina, 475 glandulosa, 475, 476 Comida de danta, 808 Commianthus, 792 concolor, 794 discolor, 795 pilosus, 795 schomburgkii, 796 Condalia, 553 lanceolata, 556 repens, 555 Coneja sabanera, 532 Conejilla, 800 Conejo rebalsero, 830 Conosiphon, 835 aureus, 837 Conserva, 580 Cordiera, 558 acuminata, 513 bertierifolia, 513 edulis, 513 hexagyna, 513 latifolia, 514 myrciifolia, 559, 559 var. tepuiensis, 560 triflora, 560 trifolia, 560 Coronilla, 349 Cortaderia, 78 roraimensis, 78, 79

I NDEX

Coryphothamnus, 560 auyantepuiensis, 560, 561 Cosmibuena, 562 grandiflora, 561, 562 Coupoui micrantha, 582 Coussarea, 562 brevicaulis, 564 evoluta, 564 grandis, 564 hirticalyx, 564 lasseri, 565 leptoloba, 565 leptophragma, 565, 567 liesneri, 565 ovalis, 565 paniculata, 565, 567 revoluta, 565, 566 schomburgkiana, 566 violacea, 566 sp. A, 567 Coutarea, 568 campanilla, 568 hexandra, 568, 569 var. campanilla, 568 var. hexandra f. pubescens, 568 var. pubescens, 568 lindeniana, 568 pubescens, 568 speciosa, 568 Cowicowinae, 354 Crataegus obtusifolia, 491 Cruceta, 549 Cruceta de la reina, 545 Cruceta real, 778 Cruceta, 779 Cruceto, 545, 594, 778 Cryptostachys, 273 Cuentica, 733, 741 Cupi, 669 Cupi banero, 800 Cupi hoja fina, 800 Cura, 38 Curata, 38 Curello, 564 Cymbopogon bracteatus, 121 Cynodon, 78 dactylon, 78, 80 Cynosurus aegyptius, 81

domingensis, 145 indicus, 94 virgatus, 145 —D— Dactyloctenium, 80 aegyptium, 81, 81 Danta oreja de burro, 808 Dantero, 808 De-beu-ni, 36 Declieuxia, 569 alfredii, 570 brasiliensis, 570 chiococcoides, 570 var. guyanensis, 570 fruticosa, 570, 570 var. guyanensis, 570 var. mexicana, 570 mexicana, 570 roraimensis, 730 tenuiflora, 570 Dendrosipanea, 571 revoluta, 571, 572 spigelioides, 571 wurdackii, 571 Despretzia, 294 Diachyrium, 273 Diandrochloa, 97 diplachnoides, 100 glomerata, 100 japonica, 100 Dichanthelium, 81 aequivaginatum, 82 davidsei, 82 hebotes, 82 pycnoclados, 82, 83 sciurotoides, 82 telmatum, 83 Dichromena squarrosa, 421 Diclidanthera, 321 bolivarensis, 322, 322 octandra, 322 orinocensis, 322 wurdackiana, 321, 322 sp. A, 323 Dicrobotryum divaricatum, 615 Didymochlamys, 572 connellii, 572, 573 Diectomis, 17 angustata, 19

855

fastigiata, 19 laxa, 19 Digitaria, 84 aurea, 49 ciliaris, 85 fuscescens, 85, 86 glabriculmis, 86 horizontalis, 85, 87 insularis, 85, 88 nervalis, 85 nuda, 85 platycaulis, 51 sacchariflora, 85 sanguinalis, 85 tenuis, 85 uniflora, 51 venezuelae, 86 violascens, 86, 86 Dimorphostachys, 214 Diodella rigida, 575 teres, 577 Diodia, 573 apiculata, 575, 577 hyssopifolia, 575, 577 var. hyssopifolia f. psiloclada, 575 var. linearis, 575 f. glabriuscula, 575 kuntzei, 575 multiflora, 576 ocymifolia, 576 ottonis, 839 prostrata, 577 radula, 576 subsp. venezuelensis, 576 rigida, 575 sarmentosa, 576 setigera, 575 spicata, 576 teres, 577 subsp. angustata, 577 var. angustata, 577 f. latior, 577 subsp. prostrata, 577 var. prostrata f. latifolia, 577 f. leiocarpa, 577 Dorona, 784 Duckea, 419 cyperaceoidea, 421 flava, 420, 421

856

I NDEX

[Duckea] junciformis, 420, 421 squarrosa, 421, 421 Dufourea, 314 trifaria, 315 Duidania, 578 montana, 578, 578 Dukuadi-jodu, 354 Duroia, 579 bolivarensis, 580 eriopila, 580 var. eriopila, 581, 584 f. glabra, 581 fusifera, 581 genipoides, 581, 585 gransabanensis, 581 kotchubaeoides, 581, 586 maguirei, 582 var. maguirei, 582 var. patentinervia, 582 micrantha, 582 nitida, 582 paruensis, 582 retrorsipila, 583 saccifera, 583, 586 sprucei, 582 strigosa, 583 velutina, 583, 587 —E— Echepen yo’, 76 Echinochloa, 89 colona, 89 crus-pavonis, 89, 91 polystachya, 89, 90 Echinolaena, 92 gracilis, 92, 93 inflexa, 92, 93 Echipipin, 329 Eichhornia, 371 azurea, 372, 373 crassipes, 372, 373 diversifolia, 372 heterosperma, 372 venezuelensis, 372 Elaeagia, 587 maguirei, 587 var. maguirei, 587, 588 var. pubens, 588 Eleusine, 92 indica, 94, 95 Elsota, 337

Elionurus, 94 adustus, 94 muticus, 94, 96 planifolius, 94 Emmeorhiza, 588 umbellata, 589 subsp. septentrionalis, 589, 589 Endlichera, 588 Enteropogon, 94 mollis, 96, 97 Epidryos, 422 guayanensis, 422, 423 sp. A, 422, 424 Epiphyton, 422 Eragrostis, 97 acuminata, 100 acutiflora, 98, 103 var. humilior, 98 amabilis, 98 var. plumosa, 99 amoena, 100 atrovirens, 99 aturensis, 100 barbata, 278 chariis, 99 ciliaris, 99 var. ciliaris, 99 var. patens, 99 conferta, 100 diplachnoides, 100 elegantula, 99 filiformis, 101 flamignii, 99 floridana, 101 fragilis, 101 gangetica, 99 glomerata, 100 guianensis, 99, 102 hapalantha, 100 hypnoides, 99 interrupta var. diplachnoides, 100 var. laxiflora, 100 var. parviflora, 100 japonica, 100, 103 lindeniana, 100 linkii, 101 maypurensis, 100 var. densiuscula, 100 var. meratiana, 100 meratiana, 100 namaquensis

var. diplachnoides, 100 pallida, 100 panamensis, 100 pectinacea, 101 var. pectinacea, 101 pilosa, 101 plumosa, 99 polytricha, 101 var. glabrior, 101 var. hirsutior, 101 purpusii, 101 reptans var. contracta, 99 var. laxior, 99 var. pygmaea, 99 stenoclada, 99 tenella, 99 var. japonica, 100 var. plumosa, 99 var. viscosa, 101 trichocolea, 101 var. floridana, 101 unioloides, 101, 102 vahlii, 100 var. polyantha, 100 viscosa, 101, 103 var. pilosissima, 101 warmingii, 102 Erianthus, 256 asper, 259 saccharoides var. ß trinii, 259 trinii, 259 Eriochloa, 104 distachya, 104, 105 punctata, 104 Eriochrysis, 104 cayennensis, 106 aff. holcoides, 105, 106 Erosion, 97 hypnoides, 99 Espuelito montañero, 549 Etuburrucuaja, 354 Euplassa, 385 chimantensis, 386 venezuelana, 386, 386 —F— Faramea, 589 angustifolia, 592 anisocalyx, 592 var. pulchella, 592

I NDEX

berryi, 592 boomii, 593 capillipes, 593, 597 cardonae, 593, 598 crassifolia, 593 egregia, 594, 599 leptoloba, 565 maguirei, 594 morilloi, 594 multiflora, 594 var. epedunculata, 594 neblinae, 594 occidentalis, 594 subsp. occidentalis var. meridionalis, 595 orinocensis, 595 pachydictyon, 596 paludicola, 596 parvibractea, 595 pulchella, 592 schomburgkiana, 566 sessilifolia, 595, 599 stenopetala, 596 tamberlikiana, 596 subsp. tamberlikiana, 596 torquata, 596, 597 yavitensis, 596 yutajensis, 596 Ferdinandea, 600 Ferdinandusa, 600 boomii, 601 goudotiana, 601 var. eciliata, 601, 602 var. goudotiana, 601 var. psilocarpa, 601 guainiae, 601, 603 neblinensis, 601, 602 sprucei, 601 uaupensis, 601, 603 Festuca filiformis, 145 Flor de Canaripó, 449 Flor de Inírida, 449 Flor de Inírida real, 426 Flor de Maroa, 426 Flor de rana, 513 Flor del Atabapo, 449 Francisco Javier, 545 Froesia, 394 gereauana, 396 tricarpa, 395, 396 Fruta de pajarito, 741

Fruto de macaco, 326 Fuchsia, 822 —G— Galium, 604 croceum, 604 hypocarpium, 604 var. hypocarpium, 605, 605 sect. Relbunium, 604 Gamotopea alba, 761 purpurea, 761 surinamensis, 761 Garapatica, 558 edulis, 513 Gardenia armata, 777 edulis, 513 Gardeniola, 558 Garrapata, 671 Gaspadillo negro, 401 Genipa, 605 americana, 606 var. americana, 607 var. caruto, 607 caruto, 607 edulis, 513 spruceana, 607, 608 striiflora, 837 Genipella, 512 Geocardia, 608 Geophila, 608 cordifolia, 609 var. cordifolia, 609, 609 orbicularis, 609 var. neblinae, 609, 610 var. orbicularis, 610 paraensis, 761 picta, 761 repens, 609, 610 tenuis, 610 Gleasonia, 610 duidana, 611 var. duidana, 611, 611 var. latifolia, 611 var. oblanceolata, 611 Gomphosia, 600 Gonzalagunia, 612 dicocca, 612 subsp. dicocca var. guianensis, 612

857

subsp. venezuelensis, 612 surinamensis, 613, 613 Gouania, 477 blanchetiana, 478 colurnifolia, 478 cornifolia, 478 discolor, 478 frangulifolia, 478 hypochroa, 478 polygama, 476, 479 pubescens, 479 tomentosa, 479 ulei, 478 wurdackii, 479 Gouinia, 106 latifolia, 107, 107 Grumilea, 708 Guacamaya, 426 superba, 425, 426 Guacharaca, 549 Guácimo cimarrón, 475 Guácimo negro, 368 Guaco, 618, 619 Guadua, 108 angustifolia, 111 ciliata, 109, 111 fascicularis, 111 aff. glomerata, 111 latifolia, 111 venezuelae, 110, 111 Guaiabara, 349 Guambe, 339 Guanabanillo, 495 Guarataro, 595 Guaricha, 594 Guatacare amarillo, 568 Guayaba de monte, 513 Guayaba rebalsera, 514 Guayabilla morada, 513 Guayabillo, 539 Guayabita, 657, 741 Guayabito, 513 Guayapapón, 354, 355 Guaya-papón sabanero, 354 Guayapón, 352 Guettarda, 614 acreana, 615 divaricata, 615, 616 leiantha, 615 macrantha, 615 malacophylla, 615 spruceana, 616 ulei, 615

858

I NDEX

Guianaguiarra, 646 Gymnopogon, 111 fastigiatus, 112 foliosus, 112, 112 mollis, 97 spicatus, 112 Gynerium, 113 saccharoides, 113 sagittatum, 113, 114 —H— Hackelochloa, 113 granularis, 115, 115 Hamelia, 616 axillaris, 617, 617 lutea, 617 patens, 617 Hedyotis lancifolia, 665 Hemidiodia, 573 ocymifolia, 576 Henriquezia, 617 aturensis, 701 nitida, 618 var. nitida, 618, 619 var. oblonga, 618 oblonga, 618 verticillata, 619 Hesperomeles, 491 heterophylla, 491 obtusifolia, 491 var. obtusifolia, 491, 492 Heteranthera, 374 diversifolia, 372 multiflora, 374, 374 reniformis var. multiflora, 374 Heteropogon, 117 contortus, 116, 117 secundus, 289 stipoides, 289 truncatus, 289 Heteropsychotria, 755 Hiel de pescado, 401 Hillia, 619 foldatsii, 620 illustris, 620, 621 parasitica, 621, 622 psammophila, 622 rivalis, 621, 622 tubiflora, 620

Hoja de baba, 783 Hoja de danto, 704 Hoja de guacuraya, 755 Hoja de mono, 475 Holcus bicolor, 272 sorghum, 272 striatus, 260 Holstianthus, 623 barbigularis, 623, 624 Homolepis, 117 aturensis, 118, 119 glutinosa, 118, 119 isocalycia, 118, 119 Homopogon, 287 Horca, 391 Horca mandingo, 391 Houstonia fruticosa, 570 Huitillo, 608 Huonay-key, 355 Hymenachne, 118 amplexicaulis, 121 donacifolia, 120, 121 Hyparrhenia, 121 bracteata, 121, 122 rufa, 123 Hypogynium, 17 virgatum, 26 —I— Icayek, 401 Ichnanthus, 123 acuminatus, 124 angustus, 125 axillaris, 125 breviscrobs, 124, 126 calvescens, 124, 127 chaseae, 124 duidensis, 125 ephemeroblepharis, 125, 128 hispidus, 124 lancifolius, 125 var. weberbaueri, 125, 129 longifolius, 125 neblinaensis, 125 nemoralis, 125 nemorosus, 125 nubilis, 125 pallens, 125

panicoides, 125, 127 procurrens, 130, 131 ruprechtii, 131 sabulosus, 131 serratus, 125 sucrensis, 131 tamayonis, 131 tectus, 125 tenuifolius, 125 tenuis, 128, 131 venezuelanus, 131 vestitus, 124 weberbaueri, 125 Imperata, 131 brasiliensis, 132, 133 contracta, 133 flexuosa, 133 Ipecacuanha, 708 Isachne, 133 arundinacea, 135 ligulata, 135 polygonoides, 133, 134 rigens, 134, 135 Ischaemopogon, 135 latifolius, 137 Ischaemum, 135 arenosum, 136, 137 guianense, 136, 137 var. schomburgkii, 137 latifolium, 137 subsp. hirtivaginum, 137 var. minus, 137 rugosum, 137 var. segetum, 137 segetum, 137 Isertia, 624 hypoleuca, 625, 626 parviflora, 625, 628 var. hirta, 625 rosea, 626, 627 verrucosa, 627, 628 Ixora, 628 acuminatissima, 629, 630 angulata, 657 duidae, 810 intropilosa, 630 occidentalis, 594 panurensis, 630 podocarpa, 826 schomburgkiana, 630 ulei, 630 yavitensis, 630

I NDEX

—J— Jaboncillo, 478 Jagua, 607 Jakí-ko, 397 Jariso, 355 Jarizo, 354 Jasmín de estrella, 704 Jasmín de monte, 615 Jasmín montañero, 596 Jazmín de monte, 777 Jazmín montañero, 594 Jenmaniella, 307 ceratophylla, 307 var. parva, 307, 307 Jiba, 131 Jibaju, 38 Jovillo, 796 Juiqui-Juiqui, 332 —K— Kabadi-wochi, 280 Karaw-karaw, 353, 354 Karaw-yek, 353 Karoráy, 495 Kasemani, 489 Kawadi-jodedü, 389 Kehkoyó, 803 Kitak-Yek, 796 Knoxia brasiliensis, 570 Kowa-waka-anamahu, 754 Krombholzia, 294 Kunhardtia, 426 radiata, 428 rhodantha, 427, 428 Ku’pe kwamen, 330 Kurawa-dek, 318 Kurucosadíyu, 688 Kutchubaea, 631 longiloba, 632, 632 micrantha, 632 morilloi, 632, 634 neblinensis, 633, 634 sericantha, 633, 634 —L— Lacunaria, 396 crenata, 397, 398 jenmanii, 397, 398 macrostachya, 397

oppositifolia, 397 spruceana, 397 Ladenbergia, 634 amazonensis, 635, 636 lambertiana, 636, 637 lucens, 636 magnifolia, 636 oblongifolia, 636 puberula, 636 roraimae, 784 schomburgkii, 636 venamoensis, 636 sp. A, 638 Lappagopsis, 45 Lasiacis, 137 acuminata, 143 anomala, 139, 140 compacta, 139 glabra, 139 guaraniticum, 143 liebmanniana, 139 ligulata, 139, 140 nigra, 139 patentiflora, 143 procerrima, 139, 141 ruscifolia, 139 scabrior, 139, 142 sloanei, 139, 142 sorghoidea, 139 var. patentiflora, 143 swartziana, 143 Lata, 353, 355 Lazo, 830 Lechuga de agua, 372 Leersia, 143 distichophylla, 143 grandiflora, 143 hexandra, 143 ligularis, 143, 144 var. glabriflora, 144 var. grandiflora, 143 Lengua de vaca, 516 Leptochloa, 145 domingensis, 145 filiformis, 145 scabra, 145, 146 virgata, 145, 146 Leptocoryphium, 147 lanatum, 147, 148 Lepturopsis, 255 Let’dété rowe, 405 Ligea richardiana, 305

859

var. corymbosa, 304 var. exilis, 304 Limnosipanea, 638 guaricensis, 639 kuntzei, 639 palustris, 638, 639 schomburgkii, 638 spruceana, 639, 639 ternifolia, 639 Limoncillo, 484 Lipostoma, 553 Lirio de agua, 372 Lithachne, 147 pauciflora, 147, 148 Lonicera alba, 545 Lophogyne capillacea, 308 Lucinaea, 822 Luñek, 38 Luse, 38 Luziola, 149 bahiensis, 149 doelliana, 149, 150 pittieri, 149 spruceana, 149 subintegra, 149, 150 Lycurus muticus, 94 —M— Machaonia, 639 brasiliensis, 640, 640 Macho de puya, 401 Macollo, 549 Macrocnemum coccineum, 846 Maguireocharis, 640 neblinae, 641, 642 Maguireothamnus, 642 jauaensis, 643 var. breweri, 643 speciosus, 642 subsp. jauaensis, 643, 643 subsp. speciosus, 643, 643 tatei, 643, 643 var. latifolius, 644 Majada, 493 Malanea, 644 angustifolia, 647

860

I NDEX

[Malanea] auyantepuiensis, 646 chimantensis, 646 gabrielensis, 646, 648 guaiquinimensis, 646 hypoleuca, 646 jauaensis, 646 macrophylla, 646 microphylla, 647, 648 obovata, 647, 648 ptariensis, 647 roraimensis, 647 sarmentosa, 647, 649 f. tomentosa, 647 setulosa, 647 sipapoensis, 647 ueiensis, 648 ursina, 648 Manáse-yek, 625 Mandingo, 391 Manettia, 649 alba, 650, 651 calycosa, 650 var. karsteniana, 650 coccinea, 650 reclinata, 651, 651 uniflora, 651 Mangle, 354, 355, 488 Mangle blanco, 488 Mangle de burro, 488 Mangle negro, 488 Mangle rojo, 488, 489 Manisuris aurita, 76 guianensis, 256 loricata, 256 Manteco de agua, 405 Manteco rebalsero, 368 Manzanillo, 704 Mapouria, 719 alba, 722 borjensis, 727 cupularis, 735 fockeana, 731 guianensis, 746 herbacea var. orbicularis, 609 microdon, 747 podocephala, 753 remota, 755 sclerocalyx, 810 spruceana, 810 subsessilis, 802

var. angustifolia, 802 tenuis, 610 umbrosa, 727 vasivensis, 762 Mara, 484 Marahuacaea, 429 schomburgkii, 428, 429 Marathrum, 307 aeruginosum, 308, 309 capillaceum, 308 squamosum, 308 var. spruceanum, 308 var. squamosum, 308 utile, 309, 309 Margaritopsis astrellantha, 724 Maria, 370 Marmelada, 513 Maspara, 704 Masure, 372 Matapalo, 825 Matotoma de rebalse, 514 Mattuschkea galioides, 696 hispida, 696 Mazano, 704 Medi-wadi, 753 Megastachya, 97 acutiflora, 98 breviflora, 99 corymbifera, 99 gouinii, 99 hypnoides, 99 maypurensis, 100 Melanopsidium, 512 Melero, 355 Melinis, 151 minutiflora, 151, 151 repens, 255 Menta-berti, 493 Mentol, 330 Meoschium, 135 rugosum, 137 Mergo, 517 Mermo, 582 Merostachys, 152 maguireorum, 152, 153 retrorsa, 152, 154 Merumea, 651 coccocypseloides, 652, 652 Mesosetum, 152 cardonum, 155 cayennense, 155, 157

chaseae, 155, 156 chlorostachyum, 155, 158 filifolium, 155, 157 loliiforme, 156 rottboellioides, 156, 158 Mespilus heterophylla, 491 Micropyxis pumila, 384 Miel de pajarito, 320 Milium compressum, 50 punctatum, 104 Mitë, 172 Mitracarpus, 653 diffusus, 654, 655 frigidus, 654 var. orinocensis, 654, 655 hirtus, 654, 655 microspermus, 654 parvulus, 655, 655 villosus, 654 sp. A, 655 Mnasium, 434 paludosum, 436 unilaterale, 451 Mnesithea aurita, 76 subgibbosa, 256 Molleja de gallineta, 397 Molleja de gallineta banera, 397 Molleja de paují, 397 Monnina, 323 cacumina, 323, 323 var. duidae, 323 duidae, 323 uaipanensis, 323 Monotrema, 429 aemulans, 430, 431 ×affine, 430, 431 bracteatum, 430 subsp. bracteatum, 430, 432 subsp. major, 430 xyridoides, 430, 432 Moorea, 78 Morinda, 656 longipedunculata, 657 peduncularis, 657, 658 tenuiflora, 657, 659

I NDEX

var. leiophylla, 657 venezuelensis, 659 Mosori, 372 Mourera, 309 fluviatilis, 309, 310 Moutabea, 324 aculeata, 324 aff. chodatiana, 325 guianensis, 325, 326 longifolia, 324 sp. A, 326 sp. B, 326 Mulunyek, 401 Murundek, 400 Mussaenda formosa, 805 spinosa, 777 Mutis, 636 Myriocladus, 159 affinis, 161 cardonae, 160, 163 churunensis, 160 confertus, 160 distantiflorus, 160 exsertus, 160 gracilis, 161 grandifolius, 161 involutus, 161, 162 longiramosus, 161 maguirei, 161 neblinaensis, 161 paludicolus, 161, 164 paraquensis, 161 paruensis, 161 purpureus, 160 simplex, 161, 163 steyermarkii, 161 variabilis, 160 virgatus, 161 wurdackii, 160 sp. A, 162 Myristiphyllum, 719 —N— Nabajoso, 555 Nacibea alba, 650 coccinea, 650 Nairu-cuaja, 354 Naletonia, 708 Naranjillo, 565 Nauclea

tomentosa, 844 Neblinathamnus, 659 aracamunianus, 660 argyreus, 660, 660 brasiliensis, 660 glabratus, 660 Neeragrostis, 97 hypnoides, 99 Neolacis corymbosa, 304 var. exilis, 304 Neurolepis, 165 angusta, 165 densiflora, 167 diversiglumis, 167 glomerata, 166, 167 nigra, 167 pittieri, 167 sp. A, 167 Nirgua, 484 Nonatelia, 708 grandiflora, 686 longiflora, 687 macrophylla, 686 officinalis, 749 paniculata, 750 racemosa, 755 secundiflora, 796 Notopleura, 661 aligera, 662 crassa, 662 multiramosa, 662 perpapillifera, 662 sandwithiana, 663 tapajozensis, 663, 663 thesceloantha, 663 uliginosa, 663 —O— Oenone guyanensis, 304 kochii, 304 longifolia, 305 multibranchiata, 305 staheliana, 305 Ohwe wale ‘empa, 582 Oido de picure, 565 Ojo de gavilán, 405 Ojo de venado, 324, 325, 326 Ojo de zamuro, 326, 339 Oldenlandia, 664

861

corymbosa, 664, 665 filicaulis, 665 lancifolia, 665, 665 tenuis, 665, 666 Olyra, 167 caudata, 168, 169 ciliatifolia, 168, 170 cordifolia, 168 ecaudata, 168 fasciculata, 172 lateralis, 212 latifolia, 168 longifolia, 168, 170 luetzelburgii, 214 micrantha, 168, 171 nana, 248 pauciflora, 147 sarmentosa, 212 standleyi, 172 sympodica, 248 wurdackii, 172 Oplismenus, 172 burmannii, 172, 173 crus-pavonis, 89 hirtellus, 173, 174 polystachyus, 89 tenuis, 131 Orégano, 532 Oreja de picure, 688, 741, 754 Oreja de rabo pelado, 754 Oribasia racemosa, 755 Oroma-Jijote, 397 Orthoclada, 174 laxa, 174, 175 rariflora, 174 var. lanceolata, 174 var. sesquiflora, 174 Oryza, 174 alta, 176 latifolia, 176, 177 monandra var. grandiflora, 143 perennis, 176 rufipogon, 176, 177 sativa, 176 Oserya, 310 perpusilla, 311, 311 Osíbu-ajuca, 365 Otachyrium, 176 grandiflorum, 179 versicolor, 178, 179

862

I NDEX

Ourouparia, 844 guianensis, 844 tomentosa, 844 —P— Pagamea, 666 anisophylla, 668, 674 auyantepuiensis, 560 brevipedunculata, 669 capitata, 669 subsp. capitata, 669, 674 f. breviloba, 669 subsp. conferta, 669 conferta, 669 coriacea, 669, 677 var. pubescens, 669 diceras, 670, 676 duidana, 670, 677 garryoides, 679 guianensis, 670, 675 hirsuta, 670 jauaensis, 671, 673 magniflora, 671, 675 montana, 671, 675 pauciflora, 671, 676 plicata, 671, 678 var. multinervia, 671 plicatiformis, 672 reducta, 672 sessiliflora, 672, 678 spadicea, 756 standleyana, 672, 677 steyermarkii, 520 thyrsiflora, 672, 676 uniflora, 672 velutina, 673, 678 Pagameopsis, 678 garryoides, 679, 680 maguirei, 679, 680 subsp. neblinensis, 679 var. angustifolius, 679 subsp. pusillus, 679 var. glabrus, 679 Paja aguja, 665 Paja canilla teu teu, 115 Paja carrizo, 168 Paja cepillo, 266 Paja cortaderia, 280 Paja del Brasil, 123 Paja gordura, 151

Paja imán, 266 Paja macaguera, 282 Paja pelua, 89 Paja perro de agua, 168 Pajarito, 778 Palicourea, 680 aculeifera, 688 bracteosa, 684 calophylla, 684 cardonae, 684 chrysorrhachis, 688 corymbifera, 684, 692 crocea, 685, 693 var. riparia, 685 f. heterodoxa, 685 croceoides, 685 elliptica, 737 expetens, 684 fastigiata, 685 foldatsii, 686 glabriflora, 686 grandiflora, 686, 695 grandifolia, 686, 693 var. sprucei, 686 guianensis, 687 subsp. guianensis, 687, 692 subsp. occidentalis, 687 var. occidentalis f. glabra, 687 huberi, 687 lancigera, 687 longiflora, 687 longistipulata, 688 subsp. chrysorrhachis, 688 subsp. longistipulata, 688, 694 nitidella, 688 obtusata, 689, 691 ottohuberi, 689 pensilis, 689 perquadrangularis, 689 var. guayanensis, 689 quadrifolia, 690 subsp. quadrifolia, 690 rigida, 690 subsp. rigida var. hirtibacca, 690 riparia, 685 roraimae, 748 sprucei, 686 tepuicola, 690, 694

triphylla, 690, 694 wurdackiana, 691 sp. A, 691 Palo culebra, 809, 810 Palo de agua, 366 Palo de agujón, 783 Palo de baba, 583 Palo de bachaco, 686 Palo de boya, 514 Palo de cogollo de piedra, 794 Palo de culebra, 475 Palo de gallineta, 794 Palo de paloma, 401 Palo de peluza, 795 Palo de pereze, 564 Palo de sortija, 686, 755 Palo de tema, 596 Palo hinchazón, 783 Palo Maria, 370 Palo ojo de picure, 794 Palo perro de agua, 353, 368 Panicum, 179 aequivaginatum, 82 agglutinans, 143 albociliatum, 82 altum, 183 amplexicaule, 121 angulosum, 188 angustissimum, 260 appressifolium, 82 arctum, 183 arrectum, 64 asperifolium, 282 aturense, 118 auyanense, 185 axillare, 125 sect. Brachiaria, 64 burmannii, 172 sect. Cabrera, 45 calvescens, 124 caricoides, 183, 190 caroniense, 276 cayennense, 183 cervicatum, 184, 191 chlorostachyum, 155 chnoodes, 184, 192 chrysites, 49 chrysoblephare, 50 chrysodactylon, 49 churunense, 184 ciliare, 85 coenosum, 188

I NDEX

colonum, 89 compactum, 139 cordovense, 184, 193 cowanii, 184 curvifolium, 184 cyanescens, 184, 194 dactylon, 78 davidsei, 82 deciduum, 184, 195 decipiens, 276 densifolium, 186 subg. Dichanthelium, 81 dichotomiflorum, 184, 196 discrepans, 184, 192 distichum var. lancifolium, 188 divaricatum var. lanatum, 143 var. latifolium, 143 donacifolium, 121 elephantipes, 184, 196 eligulatum, 185 esenbeckii, 248 exasperatum, 49 excavatum, 15 fasciculatum, 66 fistulosum, 184 fontanale, 185 fonticolum, 185 funckianum, 187 fuscum, 66 geminatum, 214 geniculatum, 266 glutinosum, 118, 143 gracillissimum, 188 grande, 185, 197 graniticum, 187 granuliferum, 185, 194 guaraniticum, 143 guianense, 186 haenkeanum, 185, 198 hebotes, 82 hirsutum, 185, 199 hirtellum, 174 hirtum, 185, 200 hispidifolium, 186, 195 humidicolum, 67 hylaeicum, 186, 201 ichunense, 186 immersum, 50 inversum, 184 isocalycium, 118 jauanum, 186, 202

kaietukense, 187 kavanayense, 184 lanatum, 143 var. sorghoideum, 143 sect. Lasiacis, 137 laterale, 212 var. ß, 248 latifolium, 139 laxum, 276 liebmannianum, 139 ligulare, 186 loliiforme, 156 luticola, 276 maguirei, 184 manacalensis, 189 mauryi, 186 maximum, 186, 194 mertensii, 186, 203 micranthum, 186, 193 var. hirtum, 186 milleflorum, 188 millegrana, 187, 191 mirandum, 82 molle, 67 muticum, 67 myuros, 260 nemorale, 125 nemorosum, 125 nervosum, 187 obovatum, 188 olyroides, 187, 204 orinocanum, 187, 200 pallens, 125 pandum, 187 pantrichum, 187 parvifolium, 187 petrense, 187 petrosum, 287 pilisparsum, 188 pilosum, 188 var. lancifolium, 188 plantagineum, 67 platycaulon, 51 poiretianum, 266 politii, 185 polycomum, 188, 198 polygonatum, 188, 201 polygonoides, 133 polystachion, 240 praegnans, 143 procerrimum, 139 procurrens, 131 pulchellum, 188

863

purpurascens, 67 pycnoclados, 82 pyrularium, 188, 203 quetameense, 188 rariflorum, 174 rigens, 135 rivale, 188 rottboellioides, 156 rudgei, 189 ruscifolium, 139 savannarum, 50, 184 scabridum, 189 sciurotoides, 82 sellowii, 189 siccaneum, 188 sipapoense, 189 sloanei, 139 sorghoideum, 139 spissifolium, 185 stenodes, 189, 190 stenodoides, 183 stenophyllum, 276 steyermarkii, 189, 205 stoloniferum, 189, 206 subcordatum, 185 subfalcatum, 226 subinclusum, 185 supernum, 186 surinamense, 54 swartzianum, 143 tamayonis, 188 tatei, 184 telmatum, 83 tenax, 266 tenuissimum, 274 tepuianum, 189 tiricaense, 82 tiricaoides, 82 trichoides, 205, 208 tricholaenoides, 207, 208 tropidoblephare, 184 vannum, 184 versicolor, 179 vigoratum, 183 vilvoides, 260 vinaceum, 184 vulpisetum, 266 yavitaense, 198, 208 zizanioides, 15 Pankecho, 355 Panopsis, 386 cuaensis, 389 ornatinervia, 387

864

I NDEX

[Panopsis] parimensis, 387 ptariana, 387 rubescens, 388, 389 var. simulans, 389 sessilifolia, 388, 389 tepuiana, 389 Papagayo, 846 Pappophorum, 208 mucronulatum, 208 pappiferum, 208, 209 Paraguatán, 514, 827 Paraguatán blanco, 827 Para-ma-yek, 389 Para-pájaro, 368 Paraparillo, 745 Paraparo, 704 Paratheria, 208 prostrata, 209, 210 Pareu-rei-yek, 422 Pariana, 210 angustifolia, 278 obtusa, 210 pallida, 210 radiciflora, 210, 211 trichosticha, 210 violascens, 210 vulgaris, 210 zingiberina, 210 Pari-co-guác-ariskú, 331 Parikuá-muranó, 330 Paripari, 565, 756 Pariwá-muranó, 329 Parodiolyra, 212 lateralis, 212, 213 luetzelburgii, 213, 214 micrantha, 172 Pashomacasi, 484 Pashonema-casiji-ajuji, 484 Paspalidium, 214 geminatum, 214, 215 Paspalum, 214 abstrusum, 223 altsonii, 220, 227 ammodes, 221 sect. Anastrophus, 45 anceps, 48 apiculatum, 221, 228 appendiculatum, 51 arenarium, 221 aspidiotes, 221, 229 atabapense, 221 aureolatum, 223

aureum, 49 axillare, 283 sect. Cabrera, 45 canum, 221 capillare, 49 carinato-vaginatum, 49 carinatum, 221, 229 caulescens, 50 chaffanjonii, 221 chrysites, 49 chrysoblephare, 50 chrysodactylon, 49 var. glabratum, 49 var. psilachne, 49 var. villosum, 49 clavuliferum, 221 compressum, 51 var. arenarium, 52 conjugatum, 221 var. subcordatum, 51 conspersum, 222 contractum, 223 convexum, 222, 230 corcovadense, 222 coryphaeum, 222, 231 decumbens, 222, 232 delicatum, 222 densum, 222, 233 depressum, 51 dispar var. marahuacense, 225 eminens, 51 erectifolium, 221 excavatum, 51 extenuatum, 50 familiare, 222 fasciculatum, 223, 231 filostachyum, 51 fissifolium, 52 flexile, 53 fockei, 54 furcatum var. fissum, 51 var. parviflorum, 51 fuscescens, 85 gardnerianum, 223, 228 gnaphalioideum, 50 gossypinum, 223 guadaloupense, 51 hyalinum, 223, 234 immersum, 50 var. pilosum, 50 indutum, 222

intermedium, 223 iridifolium, 52 lanatum, 147 lanciflorum, 223, 235 laticulmum, 51 leptostachyum, 53 longispicum, 53 macropodium, 51 maculosum, 223, 236 melanospermum, 223, 230 millegrana, 224, 233 minutum, 50 morichalense, 224 multicaule, 224 nudatum, 224, 237 nutans, 224 orbiculatum, 224, 236 paniculatum, 224 parviflorum, 224, 237 pectinatum, 225, 235 petilum, 225, 232 pictum, 225, 234 piligerum, 223 pilosum, 225, 232 platycaulon, 51 var. gracilius, 53 var. parviflorum, 53 plicatulum, 225 polychaetum, 223 pulchellum, 225, 236 pulchrum, 49 purpusii, 53 raunkiaerii, 51 repens, 225, 238 saccharoides, 226, 238 scoparium var. parviflorum, 48 setiglume, 221 stellatum, 226 subciliatum, 226 subfalcatum, 226, 227 tillettii, 226 triglochinoides, 54 tristachyon, 51 vaginatum, 226 virgatum, 226, 239 wagenerianum, 226 Pasto chiguirera, 89 Pastora, 618 Pata de grillo, 330, 332 Pata de grulla, 593, 594, 596

I NDEX

Pata de picure, 755 Pata de tintín, 735 Pata gallina, 72 Pata morrocoy, 742 Patabea alba, 745 Patima laxiflora, 795 Pegadera, 266 Peludita, 742 Pendare, 838 Pene del diablo, 355 Pennisetum, 240 polystachion, 240, 241 purpureum, 240 setosum, 240 Pentro, 339, 340 Perama, 695 dichotoma, 696 var. dichotoma, 696, 698 var. hirsutula, 696 var. monocephala, 696 var. scaposa, 696 galioides, 696, 698 var. densipila, 696 var. galioides f. macrocephala, 697 f. quaternata, 697 var. hispida, 696 f. cinera, 697 var. intermedia, 697 var. longiflora, 697 hirsuta, 697, 698 var. stricta, 697 plantaginea, 697, 698 scaposa, 696 stricta, 697 Peramán de vieja, 794 Peramancito, 796 Perro de agua, 405, 629 Persicaria, 363 Pescuezo de pavo, 846 Pharus, 240 glaber, 242 lappulaceus, 242, 243 latifolius, 242, 243 mezii, 242 parvifolius, 245 subsp. parvifolius, 245 virescens, 244, 245 Phelpsiella, 433 ptericaulis, 433, 433

Phragmites, 245 australis, 245, 246 communis, 245 Picatón, 700 Piptatherum confine, 40 Piresia, 245 sympodica, 247, 248 Pitillo, 125, 168, 242 Planotia, 165 Platanote, 700 Platycarpum, 699 decipiens, 700 duckei, 700 maguirei, 700 negrense, 700 var. glaucum, 700 orinocense, 700 var. grandiflorum, 701 var. orinocense, 701, 702 rhododactylum, 701 rugosum, 701 schultesii, 701 var. zarucchii, 701 Poa acutiflora, 98 amabilis, 98 atrovirens, 99 aturensis, 100 chariis, 99 ciliaris, 99 conferta, 100 elegantula, 99 gangetica, 99 glomerata, 100 hypnoides, 99 japonica, 100 linkii, 101 maypurensis, 100 meratiana, 100 pectinacea, 101 pilosa, 101 plumosa, 99 polytricha, 101 racemosa, 100 tenella, 99 unioloides, 101 vahlii, 100 viscosa, 101 var. pilosissima, 101 Poaceae, 1 Poa-wip, 352

865

Podocarpaceae, 297 Podocarpus, 298 acuminatus, 299 aracensis, 299 brasiliensis, 299 buchholzii, 299 celatus, 299 magnifolius, 299 roraimae, 299, 300 steyermarkii, 299, 300 tepuiensis, 299, 300 Podostemaceae, 301 Podostemum ruppioides, 305 Polygala, 326 adenophora, 329 var. gracilis, 329 var. robusta, 329 alopecurus, 331 angustifolia, 332 appressa, 329, 333 subsp. cumbrensis, 329 var. gracillima, 329 var. kavanayena, 329 aristeguietae, 329 asperuloides, 329, 333 blakeana, 329, 334 brevialata, 329 brizoides, 332 caracasana, 329, 333 columbica, 329 diversifolia, 339 exigua, 330 var. fendleri, 330, 333 fendleri, 330 funckii, 331 galioides, 330 glochidiata, 330 var. glochidiata, 330, 332 gracilis, 331 hygrophila, 330, 335 incarnata, 329 leptocaulis, 331 longicaulis, 330, 335 maguirei, 331 microspora, 330, 332 mollis, 332 monticola, 332 var. bryzoides, 332 paludosa, 331 paniculata, 330, 334 retifera, 329

866

I NDEX

[Polygala] sanariapoana, 330, 334 savannarum, 331, 335 sipapoana, 331 spectabilis, 331, 336 spruceana, 331, 336 subsecunda, 329 subtilis, 331, 336 tenella, 331, 335 timoutoides, 331 var. maguirei, 331, 334 timoutou, 332 trichosperma, 332 variabilis, 332 violacea, 332, 334 Polygalaceae, 316 Polygonaceae, 347 Polygonum, 363 acuminatum, 363, 364 var. glabrecens, 363 var. weddelii, 363 declinatum, 353 densiflorum, 365 ferrugineum, 363 glabrum, 365 hydropiperoides, 365 portoricense, 365 punctatum, 365 Polypogon spicatus, 112 Pontederia, 374 azurea, 372 cordifolia, 376 crassipes, 372 rotundifolia, 375, 376 triflora, 375, 376 Pontederiaceae, 371 Portlandia hexandra, 568 Portulaca, 377 cinerea, 381 elatior, 379 fruticosa, 382 halimoides, 379 insignis, 379 marginata, 379 mucronata, 379, 380 oleracea, 379, 380 paniculata, 383 pilosa, 379 pusilla, 380, 380 pygmaea, 380 sedifolia, 381

teretifolia, 381 triangularis, 382 umbraticola, 381 Portulacaceae, 376 Posoqueria, 702 gracilis, 703 latifolia, 703 subsp. gracilis, 703 subsp. latifolia, 704, 706 longiflora, 704, 706 panamensis, 704 subsp. grandiflora, 704 williamsii, 704, 705 Potoruco, 354 Primulaceae, 383 Proteaceae, 384 Prunus, 492 accumulans, 493 amplifolia, 493 espinozana, 494 lichoana, 494 myrtifolia, 495 var. accumulans, 493 var. myrtifolia, 493, 495 sphaerocarpa, 495 wurdackii, 494, 495 Pseudochimarrhis, 542 Psychotria, 706 acuminata, 721 adderleyi, 721 adenophora, 722 alba, 722 alboviridula, 755 aligera, 662 amazonica, 686 amplectans, 722, 766 amplinoda, 663 var. ayangannae, 663 anartiothrix, 723 anceps, 723 apoda, 724 arcuata, 730 arenaria, 732 astrellantha, 724 atabapoensis, 758 atricapilla, 742 aubletiana, 724 var. andina, 725 var. cacuminis, 725 f. tomentella, 725 auyantepuiensis, 742 avia, 810

axillaris, 802 bahiensis var. cornigera, 733 barbiflora, 740 barcellana, 754 barnebyana, 725 berryi, 725 berteroana, 725 blakei, 726 blepharophylla, 747 bolivarensis, 726 boliviana, 727 borjensis, 727 bostrychothyrsus, 728 brachybotrya, 739 bracteata var. intermedia, 746 var. latifolia, 746 var. tenuifolia, 746 bracteocardia, 728 brazaoi, 728 breweri, 759 cacuminis, 725, 738 calviflora, 733 campyloneura, 729 campylopoda, 729, 768 capitata, 729 subsp. inundata, 730, 765 var. roraimensis, 730 var. septentrionalis, 730 cardiomorpha, 730 subsp. perpusilla, 730 carrenoi, 731 carthagenensis, 731 casiquiaria, 731, 769 celiae, 732 ceratantha, 732 cerronis, 732 chionantha, 746 chlorantha, 723 chondroloma, 756 cincta, 744 colorata, 733 concinna, 733 cordifolia, 730 subsp. perpusilla, 730 cornifolia, 807 cornigera, 733 corymbifera, 684 coussareoides, 734, 767 var. ciliata, 734

I NDEX

crassa, 662 f. alba, 662 f. angustior, 662 crocea, 685 crocochlamys, 734, 769 cubitalis, 730 cupularis, 735 davidsei, 725 deflexa, 735 subsp. campyloneura, 729 subsp. venulosa, 763 deinocalyx, 736 dichotoma, 752 duidana, 736 duricoria, 736 var. duricoria, 737 var. longiloba, 737, 768 durifolia, 737, 768 edaphothrix, 684 egensis, 737 erecta, 802 f. fluctuans, 802 ernestii, 737 erythrophylla, 740 everardii, 738, 768 subsp. neblinensis, 738 subsp. sipapoensis, 737 subsp. tatei, 759 fanshawei, 738 fastigiata, 685 fimbriata, 807 flavovirens, 745 flexuosa, 750 fluctuans, 802 var. angustifolia, 802 fockeana, 731 foetidiflora, 727 foveolata, 731 franquevilleana, 739 glandulicalyx, 739 subsp. glandulicalyx var. densipubens, 739 glaucescens, 741 gracilenta, 739 grandifolia, 686 guanchezii, 740 guianensis, 687 hancorniifolia, 762 var. angustifolia, 762 var. longifolia, 762 hemicephaelis, 740, 766 heterocarpa, 730

heteroneura, 740 subg. Heteropsychotria, 708 hoffmannseggiana, 740 var. erythrophylla, 740 homoplastica, 758 horizontalis, 741 var. glaucescens, 741 var. psilophylla, 741 humboldtiana, 741, 770 var. ornata, 741 imthurniana, 742, 770 f. ptariensis, 742 inundata, 730, 807 involucrata, 749 iodotricha, 742, 771 subsp. atricapilla, 742 irwinii, 743 jauaensis, 743, 770 kappleri, 743 kukenanensis, 727 lancigera, 687 latifolia, 802 leiantha, 743 longicuspis, 744 longiflora, 687 longistipula, 755 longistipulata, 688 loretensis, 744 lourteigiana, 744 lupulina, 744 subsp. lupulina, 745, 771 subsp. rhodoleuca, 745 var. maypurensis, 745 f. pubens, 745 maguirei, 810 maguireorum, 746 mapourioides, 746 mapuria, 746 martiana, 752 maturacensis, 746 maypurensis, 745 medusula, 747, 772 microbotrys, 747 microcephala, 740, 754 var. tripotamica, 754 microdon, 747 var. meridionalis, 747 multiramosa, 662 nana, 748 necopinata, 750 nematostachya, 748

867

nitida, 746 nitidella, 688 sect. Notopleura, 661 oblita, 748 oblonga, 748 occidentalis, 749 officinalis, 749 sect. Oppositiflorae, 801 pacimonica, 750 palicoureoides, 747 pallidinervia, 750 paniculata, 750 paradoxa, 722 parimensis, 750 parvibractea, 751 paupertina, 751 pectinata, 751 pendula subsp. trinitensis, 662 perpapillifera, 662 persimilis, 745 phaneroloma, 751, 772 var. angustior, 751 phaneroneura, 738 phelpsiana, 752, 772 pilosa, 752 platypoda, 752 podocephala, 753, 774 poeppigiana, 753 subsp. barcellana, 754 subsp. poeppigiana, 754, 773 polycephala, 754, 775 prunifolia, 754, 773 subg. Psychotria, 719 ptariensis, 742 quadrifolia, 690 racemifera, 754 racemosa, 755 remota, 755 rhodoleuca, 745 rigida, 690 romboutsii, 722 ronaldii, 727 rosea, 755 var. rosea f. calvescens, 755 sancti-caroli, 688 sandwithiana, 663 schomburgkii, 756 sessilis, 762 var. angustifolia, 762 setifera, 730

868

I NDEX

[Psychotria] sipapoensis, 756 sororiella, 760 sororopanensis, 762 spadicea, 756, 767 speluncae, 757, 774 subsp. brevicalyx, 757 subsp. exserta, 757 sphaerocephala, 757 spicata, 758 spiciflora, 758 sprucei, 686 steinii, 743 stipulosa, 758, 775 subcuspidata, 733 subsessilis, 802 subundulata, 758 tapajozensis, 663 tapirapecoana, 758 tatei, 759 tepuiensis, 759, 773 thesceloantha, 663 transiens, 759 trichotoma, 760 triphylla, 690 turbinella, 760 var. sororiella, 760 uliginosa, 663 ulviformis, 760, 766 urophylla, 721 vareschii, 761 variegata, 761 vasivensis, 762, 774 vaupesana, 758 vellosiana, 762, 775 venezuelensis, 763 ventuariana, 763 venulosa, 763 vernicifolia, 738 verrucosa, 684 viburnoides, 723 vichadensis, 764, 771 viridis, 764 wurdackii, 764, 767 xanthocephala, 754 yapacanensis, 764, 771 yavitensis, 765 yutajensis, 732 Psychotrophum, 719 Punteral, 549, 778, 779, 804, 808 Punteral negro, 615, 778 Puruí, 513

—Q— Quiina, 399 acutangula, 405 albiflora, 401 attenuata, 400 crenata, 397 cruegeriana, 400 florida, 401, 402 gracilis, 405 guianensis, 401, 402 indigofera, 401, 403 longifolia, 401, 404 macrophylla, 401 macrostachya, 397 oblanceolata, 401 obovata, 401 oiapocensis, 405 poeppigiana, 401 pteridophylla, 403, 405 rhytidopus, 405 spruceana, 397 tinifolia, 404, 405 wurdackii, 404, 405 yatuensis, 405 Quiinaceae, 393 —R— Rabo de alacrán, 332 Rabo de guaca, 846 Rabo de ratón, 421 Racimo de cambur, 844 Raddia nana, 248 Raddiella, 248 esenbeckii, 248, 249 aff. kaieteurana, 249, 249 maipuriensis, 249 nana, 248 aff. potaroensis, 249 Rafflesiaceae, 407 Raíz de cachicamo, 332 Randia, 776 aculeata, 777 var. dasyphylla, 777 amazonasensis, 777 armata, 777, 778 brevipes, 778 caracasana, 778 dioica, 778 formosa, 805 var. densiflora, 804

var. nitida, 805 hebecarpa, 778 orinocensis, 805 spinosa, 777 venezuelensis, 779 Ranunculaceae, 410 Rapatea, 434 angustifolia, 435, 438 aracamuniana, 435 chimantensis, 435, 437 circasiana, 435, 439 fanshawei, 436, 437 longipes, 436, 438 var. modesta, 436 modesta, 436 paludosa, 436 var. paludosa, 440, 441 scabra, 441 schultesiana, 436 spruceana, 439, 441 steyermarkii, 436, 441 unilateralis, 451 viscosa, 447 yapacana, 441 Rapateaceae, 413 Rastrojero, 478 Rebalsera, 830 Reimaria acuta, 251 Reimarochloa, 250 acuta, 250, 251 brasiliensis, 251 Relbunium, 604 croceum, 604 glaberrimum, 604 hypocarpium, 604 subsp. nitidum, 604 nitidum, 604 Remijia, 779 amazonica, 781 var. amazonica, 781 aracamuniensis, 781 argentea, 781, 786 berryi, 784 delascioi, 782 densiflora, 782 subsp. densiflora, 782 var. densiflora, 782 f. glabricalyx, 782 var. minima, 782 subsp. stenopetala, 782, 786 firmula, 782

I NDEX

globosa, 782, 787 hispida, 783, 788 longifolia, 783 maguirei, 783 marahuacensis, 783 morilloi, 783, 788 pacimonica, 783, 789 pedunculata, 784, 790 pilosinervula, 784, 792 reducta, 784 roraimae, 784 var. adpressa, 784 var. auyantepuiensis, 784 var. guianensis, 784, 791 var. roraimae, 784 sessilis, 785 sipapoensis, 785, 790 spruceana, 782 steyermarkii, 785, 785 ulei, 785 uniflora, 785 wurdackii, 791, 792 Requena, 339 Retiniphyllum, 792 sect. Ammianthus, 792 angustiflorum, 794 cearense, 796 chloranthum, 794, 797 sect. Commianthus, 792 concolor, 794, 798 discolor, 795 erythranthum, 796 laxiflorum, 795 var. laxiflorum, 795, 799 var. longilobum, 795 martianum, 794 pallidum, 800 pauciflorum, 795 pilosum, 795, 798 scabrum, 796, 799 var. ayangannense, 796 var. erythranthum, 796 schomburgkii, 796, 797 subsp. occidentale, 796 var. hirticalyx, 796 var. angustiflorum, 796 secundiflorum, 796, 799 tepuiense, 797, 800 truncatum, 799, 800 var. angustifolium, 800

Reuma, 478 Reussia, 374 triflora, 376 Rhamnaceae, 473 Rhamnus, 479 acuminata, 480 chimantensis, 480, 481 longipes, 481 marahuacensis, 481 neblinensis, 481 polygamus, 479 psilocarpa, 481 sipapoenis, 481 ulei, 481, 482 Rhipidocladum, 251 aff. racemiflorum, 252 sibilans, 252, 253 sp. A, 252 sp. B, 252 Rhizophora, 486 harrisonii, 488 mangle, 487, 488 var. racemosa, 489 racemosa, 488, 489 Rhizophoraceae, 484 Rhynchelytrum, 252 repens, 254, 255 roseum, 255 Rhyncholacis, 311 applanata, 312 var. applanata, 312 coronata, 312, 313 divaricata, 312 flagellifolia, 312, 313 hydrocichorium, 312, 313 oligandra, 314 var. oligandra, 314 penicillata, 313, 314 Rhytachne, 255 guianensis, 256 rottboellioides, 256, 257 subgibbosa, 256 Richardia, 800 scabra, 800, 801 Richardsonia scabra, 800 Roblito, 341 Ronabea, 801 erecta, 802 latifolia, 802, 802 var. hispidula, 802 Rondeletia, 802 capitatum, 554

869

microdon, 747 orinocensis, 803, 803 repens, 610 rupicola, 835 var. chiribiquetana, 835 Rosaceae, 490 Rosenbergiodendron, 804 densiflorum, 804 formosum, 805 var. nitidum, 805, 805 Roshevitzia, 97 glomerata, 100 japonica, 100 Rottboellia aurita, 76 loricata, 256 subsp. glaberrima, 256 subsp. subgibbosa, 256 sanguinea, 265 Roupala, 389 chimantensis, 390, 391 complicata, 391 dentata, 391 dissimilis, 391 griotii, 390 macropoda, 391 minima, 390, 392 montana, 391 ß complicata, 391 var. dentata, 391 obtusata, 391, 393 paruensis, 393 schomburgkii, 393 sororopana, 392, 393 suaveolens, 392, 393 Rubia nitida, 604 Rubiaceae, 497 Rubus, 495 floribundus, 495 guyanensis, 496, 496 robustus, 495 schomburgkii, 496 urticifolius, 496 Rudgea, 805 avia, 810 berryi, 809 bolivarensis, 808 cardonae, 807, 813 carolina, 809 cornifolia, 807, 812 corocoroensis, 808 crassiloba, 808, 812

870

I NDEX

[Rudgea] duidae, 810 fimbriata, 807 fissistipula, 809 hostmanniana, 808 subsp. hostmanniana, 808, 813 subsp. maypurensis, 808, 811 klugii, 809, 814 lanceifolia, 809, 814 laurifolia, 810 maypurensis, 808 morichensis, 810 ovalifolia, 809 phaneroneura, 809, 815 schomburgkiana, 808 sclerocalyx, 810, 815 sipapoensis, 809 standleyana, 809 stipulacea, 810, 812 tillettii, 734 woronowii, 810, 811 wurdackii, 815, 816 Ruprechtia, 365 amentacea, 367 concina, 367 coriacea, 366 hamanii, 367 howardiana, 366, 367 laxiflora, 366 ramiflora, 366 tangarana, 367 tenuiflora, 366, 367 —S— Sabañon, 615 Sabicea, 816 amazonensis, 817, 820 aspera var. glabrescens, 818 var. velutina, 819 bariensis, 817 brachycalyx, 818, 821 cinerea, 818 edulis, 513 glabrescens, 818 var. oblongifolia, 818 grandifolia, 818, 820 guianensis, 819 hirsuta, 822 var. adpressa, 822

leucotricha, 819 morillorum, 818, 821 oblongifolia, 818 tillettii, 819 velutina, 819 subsp. chimantensis, 819 subsp. duidensis, 819 var. oblongifolia, 818 subsp. velutina, 819, 820 venezuelensis, 819, 821 villosa, 822 var. adpressa, 822 var. villosa, 822 Saccharum, 256 asperum, 259 contractum, 133 giganteum, 259 repens, 255 sagittatum, 113 trinii, 258, 259 Sacciolepis, 259 angustissima, 260, 261 myuros, 260 otachyrioides, 260 pungens, 260 striata, 260, 261 vilvoides, 260, 261 Sádu, 287 Saeta, 92 Sakau-yék, 672 Saldanha illustris, 620 Santa Maria, 369, 370 Saquí ya rojo, 728 Saranda, 838 Sardina, 665, 830 Saxofridericia, 441 aculeata, 443 australis, 443 compressa, 443, 445 duidae, 443, 445 subsp. marahuacensis, 443 grandis, 443, 446 inermis, 443, 446 pandanoides, 443 petiolata, 444 regalis, 444, 447 spongiosa, 447 subcordata, 443 sp. A, 447

Scepseothamnus, 558 Schizachyrium, 260 brevifolium, 262, 263 condensatum, 262, 263 hirtiflorum, 265 microstachyum, 263 riedelii, 265 sanguineum, 264, 265 semiberbe, 265 tenerum, 264, 265 Schoenocephalium, 447 coriaceum, 449 cucullatum, 448, 449 sipapoanum, 428 teretifolium, 448, 449 Schoenus flavus, 421 Schradera, 822 bipendunculata, 823 brevipes, 824 hilliifolia, 824, 825 maguirei, 825 marahuacensis, 825 nilssonii, 824, 825 polycephala, 824, 825 spicata, 838 yutajensis, 825 Schwenkfeldia aurea, 554 Securidaca, 337 bialata, 339, 342 cacumina, 339, 342 complicata, 340 cordata, 341 coriacea, 339, 343 densiflora, 341 divaricata, 339 diversifolia, 339 ecrista, 339 fendleri, 339 fruticans, 340 hostmannii, 340 longifolia, 340, 343 maguirei, 340 major, 341 marginata, 340, 342 mollis, 339 orinocensis, 340 paniculata, 340 var. lasiocarpa, 340 var. paniculata, 340, 344 pendula, 340, 345

I NDEX

prancei, 341 pubescens, 341, 345 pyramidalis, 341 retusa, 341, 345 rivinifolia, 339 var. parvifolia, 339 savannarum, 341 scandens, 341, 346 speciosa, 341 tenuifolia, 341 uniflora, 341, 344 venosa, 339 volubilis, 341, 346 warmingiana, 344, 347 Semaphyllanthe, 538 obovata, 539 venezuelense, 540 Senites, 294 Seréu-ka-yek, 740 Setaria, 265 geniculata, 266 gracilis, 266 macrostachya, 266 parviflora, 266 poiretiana, 266 tenacissima, 266, 267 tenax, 266, 267 vulpiseta, 266, 268 Shinatü, 397 Sickingia, 825 orinocensis, 701 pisoniiformis, 826 Siempre viva, 830 Siempre viva sabanera, 330 Simarjo, 810 Simayo, 629 Simira, 825 erythroxylon var. saxicola, 827 ignicola, 826 pisoniiformis, 826, 827 rubescens, 826, 827 Simiuri, 400 Simiyurí, 401 Sinátomo, 368 Sipanea, 828 acinifolia, 830 biflora, 829, 831 carrenoi, 829, 832 colombiana, 830 dichotoma, 830 galioides, 829, 831 glomerata, 829, 832

var. paucinervia, 829 hispida, 830 var. hispida, 830, 832 palustris, 638, 831 pratensis, 830 var. dichotoma, 830, 832 f. brachycarpa, 830 f. breviflora, 830 f. glabriloba, 830 f. glabrior, 830 pubinoda, 830 radicans, 829 rupicola, 835 setacea, 830 spraguei, 830 veris, 830, 832 wilson-brownei, 830, 831 Sipaneopsis, 833 foldatsii, 833, 834 huberi, 835 maguirei, 834, 835 morichensis, 835 pacimoniensis, 835 rupicola, 834, 835 wurdackii, 835 Solena gracilis, 703 latifolia, 703 Soowo, 131 Sorghastrum, 269 parviflorum, 269 setosum, 269, 270 Sorghum, 269 bicolor, 271, 272 parviflorum, 269 vulgare, 272 Sorteja amarilla, 684 Sortija, 721 Spartina, 273 alterniflora, 272, 273 brasiliensis, 273 Spathanthus, 449 bicolor, 450, 451 unilateralis, 451 var. unilateralis, 450, 451 Spermacoce alata, 528 apiculata, 575 assurgens, 532 aturensis, 529 caerulescens, 530

871

capitata, 528 capitellata, 532 densiflora, 530 diffusa, 654 exilis, 532 ferruginea, 529 frigida, 654 guianensis, 532 hirta, 654 hyssopifolia, 575 latifolia, 530 linearis, 575 mauritiana, 532 ocymifolia, 576 orinocensis, 529 prostrata, 531 radula, 576 remota, 532 rigida, 575 spinosa, 530 tenella, 529 verticillata, 533 villosa, 654 Sphinctanthus, 835 rupestris, 837 striiflorus, 836, 837 Sporobolus, 273 cubensis, 274, 275 effusus, 274 indicus var. pyramidalis, 274 jacquemontii, 274 minutiflorus, 274 muralis, 274 pyramidalis, 274 var. jacquemontii, 274 tenuissimus, 274, 275 Sprucea rubescens, 826 Stachyarrhena, 837 duckei, 838, 838 longifolia, 838 penduliflora, 838 reticulata, 838 spicata, 838 var. multinervis, 839 Stannia grandiflora, 704 panamensis, 704 Stegolepis, 451 aemulans, 430 albiflora, 455, 462 angustata, 456, 463

872

I NDEX

[Stegolepis] breweri, 456, 462 cardonae, 456, 463 celiae, 456 choripetala, 456 gleasoniana, 457 grandis, 457, 464 subsp. jauensis, 457 subsp. phelpsiae, 457 guianensis, 457, 464 hitchcockii, 457 subsp. hitchcockii, 457 subsp. morichensis, 458, 465 huberi, 458 humilis, 458, 466 jauaensis, 458, 465 ligulata, 458, 462 linearis, 458 maguireana, 458 membranacea, 459 microcephala, 459, 467 minor, 459 neblinensis, 459, 468 parvipetala, 459, 469 subsp. chimantensis, 459 pauciflora, 459 perligulata, 459, 468 ptaritepuiensis, 460, 464 pulchella, 460 pungens, 460, 470 squarrosa, 460, 471 steyermarkii, 461, 471 terramarensis, 461, 472 vivipara, 461, 467 wurdackii, 461 subsp. chimantensis, 461, 466 subsp. wurdackii, 461 Steinchisma, 276 decipiens, 276 laxa, 276 stenophylla, 276, 277 Steirachne, 278 barbata, 277, 278 Sterigmapetalum, 489 chrysophyllum, 489 exappendiculatum, 489 guianense, 489 subsp. guianense, 490 subsp. ichunense, 490, 490

resinosum, 490 Steyermarkochloa, 278 angustifolia, 278, 279 unifolia, 278 Stipa spicata, 288 Strempelia fimbriata, 807 Streptachne, 28 scabra, 35 tenuis, 35 Streptogyna, 280 americana, 280, 281 Streptostachys, 280 asperifolia, 282, 282 Suduchu, 124 Symmeria, 368 paniculata, 368, 368 Synisoon schomburgkianum, 795 Syntherisma, 84 —T— Tabirau-yék, 669 Tabquillo, 818 Tadiji, 489 Talinum, 381 fruticosum, 382, 382 paniculatum, 382, 383 triangulare, 382 Taparita, 514 Tapogomea, 708 alba, 760 glabra, 747 purpurea, 761 tomentosa, 753 violacea, 724 Taquari, 137 Tetramerium sessifolium, 595 Thieleodoxa lanceolata, 513 nitidula, 514 Thrasya, 282 axillaris, 283, 284 paspaloides, 283, 285 petrosa, 284, 287 robusta, 287 setosa, 283 stricta, 286, 287 trinitensis, 286, 287 Tin-tin, 368

Tobagoa, 839 maleolens, 839, 839 Tocoyena, 840 brevifolia, 841, 843 guianensis, 841, 842 var. communis, 841 var. glabriuscula, 841 neglecta, 841 orinocensis, 841, 843 pendulina, 842 Tontanea, 553 guianensis, 555 Toronjillo, 704 Tortolito, 808 Totumillo, 516 Totumito, 808 Tournefortia tenuiflora, 570 Touroulia, 405 amazonica, 406, 407 guianensis, 406, 407 jenmanii, 397 pteridophylla, 405 Trachypogon, 287 angustifolius, 288 canescens, 288 capensis, 289 subvar. mollis, 289 densus, 291 dissoluta, 289 durus, 291 filifolius, 289 glaucescens, 291 gouinii, 289 gracilis, 289 var. ciliatus, 289 var. hirtus, 289 involutus, 291 karwinskyi, 291 ligularis, 289 macroglossus, 289 mayaensis, 291 micans, 289 mollis, 288 montufarii, 288 var. bolivianus, 291 var. grandiflorus, 289 var. mollis, 288 var. pauciflorus, 289 var. pilosus, 289 muelleri, 289 palmeri, 291 parviflorus, 291

I NDEX

plumosus, 288 var. montufarii, 288 subvar. secundus, 289 var. secundus, 289 polymorphus, 291 var. bolivianus, 291 var. canescens, 288 subvar. capensis, 289 subvar. dactyloides, 289 var. dissolutus, 289 var. filifolius, 289 var. gouinii, 291 subvar. gracilis, 289 var. karwinskyi, 291 var. ligularis, 289 var. macroglossus, 289 subvar. mollis, 289 var. montufarii, 288 var. plumosus, 288 subvar. secundus, 289 var. thollonii, 291 var. truncatus, 289 var. vestitus, 289 preslii, 289 f. secundus, 289 ramosus, 291 renvoizei, 291 rigidifolius, 291 rufus, 123 spicatus, 288, 290 thollonii, 291 truncatus, 289 vestitus, 289 Trichachne, 84 insularis, 85 sacchariflora, 85 tenuis, 85 Tricholaena rosea, 255 Tricuspis latifolia, 107 Tridens, 291 flaccidus, 291, 292 Triodia flaccida, 291 Tripa de pollo, 401 Tripa de sardina, 401 Triplaris, 369 americana, 369 caracasana, 369 var. genuina, 369 var. vargasii, 369 coriacea, 366

cumingiana, 369 felipensis, 369 pavonii, 369 ramiflora, 366 schomburgkiana, 369 surinamensis, 370 weigeltiana, 370, 370 Tripogon, 293 spicatus, 293, 293 Tripsacum, 294 australe, 294, 295 hermaphroditum, 26 Trisecus frangulaefolius, 478 Tristicha, 314 trifaria, 314, 315 Trompillo, 513 Tson, 240 Tulipán, 728, 730, 754 Tulipán montañero morado, 728 Tulipán morado, 728 Tulipán rojo montañero, 754 —U— Uaka-dame, 493 Ueni-yu, 86 Uirín, 305 Uiri-yu-yek, 135 Ukánaha, 339 Uña de gato, 354, 356 Uña de gavilán, 844 Uña de murciélago, 353 Uncaria, 844 guianensis, 844, 844 tomentosa, 844, 844 Uragoga, 708 alba, 722, 761 glabra, 747 kappleri, 743 oblonga, 748 purpurea, 761 tomentosa, 753 uliginosa, 663 violacea, 724 Uralepis flaccida, 291 Urceolaria, 822 Uríyu, 118 Urochloa arrecta, 64

873

decumbens, 64 fasciculata, 66 fusca, 67 humidicola, 67 maxima, 186 mollis, 67 mutica, 67 plantaginea, 67 Uverillo, 353 Uvero, 353, 354, 355 Uvero blanco, 354 Uvero negro, 355 —V— Valantia hypocarpia, 604 Vanallo, 564 Verdolaga, 379 Verdolaga de cabra, 382, 383 Vieja, 670 Viejita, 763 Vilfa jacquemontii, 274 minutiflora, 274 pyramidalis, 274 Virecta biflora, 829 —W— Wairá-yu-yek, 751 Waiye enuda, 401 Wakinapaye, 386 Waná, 124 Wana’ ipun, 155, 266 Wana’ ipun kawëmën, 263 Wanak, 212 Waná-ká-puh, 330 Warszewiczia, 846 coccinea, 846, 847 elata, 847 obovata, 539 schomburgkiana, 846 schwackei, 847 Wayo wayo, 478 Weddellina, 315 squamulosa, 315, 316 Wiri-yu, 287 Wondaiyek, 405 Won-ta-yek, 795

874

I NDEX

—Y— Yaerute ukwae, 594 Yarí-kuto-orainó, 330 Yayabare, 581 Yejama, 724 Yurima, 830 Yutajea, 624 liesneri, 625

—Z— Zajarito montañero, 777 Zeugites, 294 sp. A, 296, 297 Zizyphus, 482 cinnamomum, 484 mauritiana, 483, 484 saeri, 483, 484